New Material of Apocopodon Sericeus Cope, 1886

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Santana et al., New material of Apocopodon sericeus PalArch’s Journal of Vertebrate Palaeontology, 8(6) (2011)

NEW MATERIAL OF APOCOPODON SERICEUS COPE, 1886 (MYLIOBATIFORMES, MYLIOBATIDAE) FROM THE PARAÍBA BASIN (NORTHEASTERN BRAZIL), AND SOUTH CAROLINA (USA) WITH A REANALYSIS OF THE SPECIES

Felipe Ribeiro de Santana,# David J. Cicimurri* & José Antonio Barbosa#

# Universidade Federal de Pernambuco DGEO-LAGESE-UFPE, Rua Acadêmico Hélio Ramos, s/n. Recife-PE, Brazil, 50740-530 * Corresponding author. South Carolina State Museum, 301 Gervais Street, Columbia 29201, [email protected]

Felipe Ribeiro de Santana, David J. Cicimurri & José Antonio Barbosa. 2011. New material of Apocopodon sericeus Cope, 1886 (Myliobatiformes, Myli- obatidae) from the Paraíba Basin (Northeastern Brazil) and South Caro- lina (USA) with a reanalysis of the species. – Palarch’s Journal of Verte- brate Palaeontology 8(6) (2011), 1-20. ISSN 1567-2158. 20 pages + 6 ﬁ gures.

Keywords: Myliobatiformes, Myliobatidae, Apocopodon, Paraíba Basin, Maria Farinha Formation, Rhems Formation, Paleocene.

ABSTRACT

Myliobatiformes (Elasmobranchii: Batoidei) is circumglobally distributed and consists of 26 extant genera within ten families. The oldest records of the group occur in upper Cretaceous deposits, and one extinct species, Apocopodon sericeus, is found in the Dani- an (lower Paleocene) Maria Farinha Formation of the Paraíba Basin, Pernambuco state, northeastern Brazil. This taxon is known from isolated teeth and several partial dentitions in various states of completeness, and herein we describe a new, incomplete (lingual portion) Brazilian dental plate. Previously considered to be endemic to the Paraíba Basin, Apocopodon was recently identiﬁ ed from South Carolina, United States. The sample consists of a nearly complete upper(?) dentition and numerous isolated teeth, and the material is indistinguishable from A. sericeus. The precise stratigraphic position and age of the South Carolina fossils is unknown, but based on the other associated Paleocene vertebrate fossils, we believe that the fossils originated from the Danian Rhems Forma- tion. The South Carolina occurrence of Apocopodon represents a signiﬁ cant geographic range extension of more than 7,000 km to the north of the type area, and the occurrence of this ray in such widely separated areas demonstrates the dispersion potential of fossil elasmobranch species. Analysis of the new specimens, along with reanalysis of all previously known Apocopodon dentitions maintained in Brazilian institutions, resulted in a revision of the morphological characteristics used to identify the taxon.

Introduction the small quarries from which the local population extracted limestone blocks. Teeth, dermal denticles and caudal fi n spines of Apocopodon sericeus has been considered to Myliobatiformes are known from upper Creta- be endemic to the Paraíba Basin (e.g. Silva et al., ceous strata of Europe, Africa, North and South 2001; Silva et al., 2002), but some isolated teeth America, and Asia (Cappetta, 1987; Marmi et found in Texas, USA were tentatively attributed al., 2010). In Brazil, remains of Myliobatiformes to the genus by Meyer (1974). However, our in- are found in the upper Cretaceous (Maastrich- terpretation of the specimens he illustrated is tian) Itapecuru Formation of the São Luís Ba- that the Texas material can be assigned to other sin (Eugênio, 1994), Maastrichtian Gramame taxa. The Maastrichtian specimen that Meyer (Silva, 2007) and lower Paleocene (Danian) Ma- (1974) identifi ed as Apocopodon? sp. 1 (p. 160, ria Farinha formations of the Paraíba Basin (Re- fi g. 50) actually represents Brachyrhizodus bouças & Silva Santos, 1956; Silva, 1993; 1994), wichitaensis, and his Cenomanian Apocopodon? and Miocene Pirabas Formation of the Pará-Ma- sp. 2 (p. 162, fi g. 51) appears to be an aberrant ranhão Basin (Silva Santos & Travassos, 1960; Pseudohypolophus mcnultyi. Silva Santos & Salgado, 1971). A nearly complete dentition collected from Fossil Myliobatiformes have been known to Williamsburg County, South Carolina (United occur within the Paraíba Basin since the 19th States), is virtually indistinguishable from a century and are represented by the extinct specimen of A. sericeus discussed by Silva et genera Rhombodus and Apocopodon, and ex- al. (1995), and this specimen (in addition to tant genera Rhinoptera and Myliobatis (Silva isolated teeth) represents the fi rst undeniable Santos & Gomes, 1987; Silva, 1993; 1994; Sil- record of Apocopodon outside of the type area va, 2007). Rhombodus binkhorsti was reported and extends the paleogeographic distribution by Rebouças & Silva Santos (1956) from the of the genus more than 7,000 km to the north. Gramame Formation, whereas the other three Herein we present morphological descriptions genera occur in the Maria Farinha Formation of the new South Carolina and Brazilian fossils, (e.g. Cope, 1886; Woodward, 1907; Silva et al., and we provide the results of a morphological 2001; Silva et al., 2002). Rhinoptera prisca was analysis of all known Apocopodon dentitions in erected by Woodward (1907) and the type ma- museum collections. Additionally, we discuss terial includes an incomplete dental plate. Apo- geographic distribution, environmental prefer- copodon sericeus was based on an isolated tooth ence, and other biological aspects of the genus. and three articulated teeth (Cope, 1886; later re- vised by Woodward, 1907; see also Silva Santos & Gomes, 1987). Until the 1980s, reference to Geological and Paleoenvironmental Apocopodon has appeared sporadically in the Settings literature and only in reference to the type ma- The coastal zone of the Paraíba Basin corre- terial (e.g. Cappetta, 1987). However, thanks to sponds to that region of the platform margin extensive studies of the fossiliferous strata in between the Pernambuco Shear Zone (PESZ), Pernambuco over the past 25 years, new speci- near the city of Recife, and the Mamanguape mens of A. sericeus have come to light, includ- Fault, an offshoot of the Patos Shear Zone ing additional isolated teeth and several denti- (PASZ), located north of the city of João Pessoa, tions (e.g. Silva et al., 1995; Silva, 2007; Santana Paraíba state (fi gure 1C, D). The coastal basin is et al., 2009). divided into three sub-basins, Olinda, Alhandra All of the Brazilian Apocopodon material, in- and Miriri (Mabesoone & Alheiros, 1993; Bar- cluding a new specimen described in this work, bosa & Lima Filho, 2006; Barbosa, 2007), and was collected from coastal plain deposits that the new Brazilian specimen described herein occur in the Paraíba Basin, Pernambuco State, was collected from the Poty Quarry (Olinda predominantly in the Poty Quarry near Paulista sub-basin) in the southern region of the Paraíba city (around 30 km to north of Recife) but also Basin (fi gure 1C). in small limestone quarries around Paulista (fi g- The sedimentary succession in the Paraíba ure 1). The Poty Quarry did not exist at the time Basin is divided into fi ve formations, including Cope obtained his specimens, and the fossils he the Coniacian(?) to Santonian Beberibe, upper examined were probably collected from one of Campanian to lower Maastrichtian Itamaracá,

Figure 1. Geographic and stratigraphic occurrences of Apocopodon fossils. A) Map of contiguous United States showing location of South Carolina; B) Enlarged view of the fossil area (Kingstree) showing regional geology (modiﬁ ed from Weems & Bybell, 1998); C) Geographic map of Brazil and stratigraphic section of the Poty Mine (adapted from Barbosa, 2007); D) Enlarged view of Pernambuco State showing regional geology of the Paraíba Basin. Arrow in C indicates stratigraphic position of Apocopodon sericeus.

Maastrichtian Gramame, Danian Maria Farinha, deposition occurred in a nearshore marine, pos- and Plio-Pleistocene Barreiras formations (Sou- sibly deltaic environment (Weems & Bybell, za, 1998; Barbosa, 2004; 2007). The Danian de- 1998), and this interpretation is supported by posits that occur in the Olinda sub-basin are vertebrate fossils attributed to the formation characterized by marl and marly limestone that (Erickson, 1998; Hutchinson & Weems, 1998; was deposited in a shallow platform environ- Sawyer, 1998). ment dominated by mud infl ux, probably in the proximity of estuaries (Barbosa et al., 2006) Age of Apocopodon Fossils (fi gure 1). These deposits represent lateral fa- cies variations related to the regressive event Cope (1886) thought that Brazilian strata yield- that began at the end of the Maastrichtian, with ing Apocopodon were of Maastrichtian age and regions near the paleo-coastline undergoing a correlative to the Fox Hills Formation of the change from a shallow platform environment United States Western Interior (also Branner, to a sub-coastal environment, and increased 1890). Upper Cretaceous elasmobranch taxa energy and infl ux of terrigenous material (in like Rhombodus binkhorsti, Scapanorhynchus, the form of sand) in the shallowest regions Cretalamna biauriculata, Squalicorax kaupi and (Barbosa et al., 2006). Overall, the Poty Quarry S. pristodontus occur in the Paraíba Basin, but deposits exhibit a coarsening upward sequence these fossils are derived from the Campanian- that refl ects this increase of terrigenous mate- Maastrichtian Itamaracá and Gramame forma- rial and possible growing infl uence of estuaries. tions (Silva, 2007; Silva et al., 2007). The Maria The Danian deposits in the Poty Quarry are Farinha Formation, the source of Apocopodon rich in fossils of marine animals like mollusks, fossils, is now known to have been deposited echinoids, fi sh, marine crocodiles, and sea tur- during the early Paleocene (Danian Stage; see tles (Muniz, 1993; Silva et al., 2001; Barbosa, Silva, 1994; Albertão et al., 1994; Silva et al., 2004; Barbosa et al., 2008). A large portion of the 1995; Stinnesbeck & Keller, 1996; Gallo et al., Danian invertebrate paleofauna is represented 2001; Silva et al., 2002; Silva, 2007; Barbosa by colonizing species that occupied the basin et al., 2008), and Cope may have examined a after a major biotic reduction at the end of the mixed assortment of fossils that lacked strati- Cretaceous (K/T event) (Stinnesbeck, 1989; Al- graphic provenience. The temporal occurrence bertão et al., 1994; Stinnesbeck & Keller, 1996; of Apocopodon was incorrectly reported as Barbosa, 2004; Neumann et al., 2009). Most of Thanetian (late Paleocene) by Cappetta (2006) the colonizing biota, mainly mollusks that in- and Cleason et al. (2010). habited the basin during the post-K/T recovery The source strata of the South Carolina Apo- period, appears to have emigrated from the copodon sample are not known with certainty, north in the Gulf of Mexico region and North as the fossils are part of a temporally mixed ver- America (Barbosa, 2004). tebrate assemblage that occurs in a lag depos- Unfortunately, less can be said about the it. Cretaceous taxa include dinosaurs and the South Carolina Apocopodon occurrence. The elasmobranchs Squalicorax kaupi, Cretalamna fossils were recovered from a lag deposit con- biauriculata, Serratolamna serrata, Schizo- taining Cretaceous and Paleocene fossils that rhiza stromeri, and Rhombodus binkhorsti, an were probably mixed together during the Plio- assemblage indicative of a Maastrichtian age Pleistocene, likely as the result of ancient coast- (Case, 1979; Padian, 1984). The Paleocene fos- al processes (based on the association of shark sils are believed to have been derived from the teeth like Carcharodon carcharias and Galeo- Rhems Formation (Erickson, 1998; Hutchinson cerdo cuvier with terrestrial vertebrate remains & Weems, 1998), a lithostratigraphic unit that like Nanippus sp.). The fossil site, located along- formed during the Danian Stage (calcareous nan- side a creek inside the city limits of Kingstree, nofossil zone NP 1, possibly as young as NP 3; Williamsburg County, is within the Maastrich- see Weems & Bybell, 1998). Considering the tian Peedee Formation (or equivalent) outcrop fact that Apocopodon is only known from the belt, and the lower Paleocene Rhems Forma- Danian of Brazil, it is parsimonious to consider tion also occurs in the area (Weems & Bybell, the South Carolina material to be of equivalent 1998). Lithostratigraphic and biostratigraphic age (not Cretaceous or Plio-Pleistocene). Sup- analyses of the Rhems Formation indicate that porting evidence for this hypothesis includes

© PalArch Foundation 4 Santana et al., New material of Apocopodon sericeus PalArch’s Journal of Vertebrate Palaeontology, 8(6) (2011) occurrences of the elasmobranchs Palaeocar- Systematic Paleontology charodon and Hypolophites hutchinsi, taxa reported from Danian strata elsewhere in the Elasmobranchii Bonaparte, 1838 USA (see Case, 1989; 1993; 1996; Kent, 1994). Neoselachii Compagno, 1977 Unfortunately, indurated matrix obtained from Batomorphii Cappetta, 1980 recesses within the aboral surface of the South Myliobatiformes Compagno, 1973 Carolina dentition did not yield microfossils for Myliobatidae Bonaparte, 1838 relative age determination of the source deposit Apocopodon Cope, 1886 (J. Self-Trial personal communication, 2010). Type and only known species – Apocopodon sericeus Cope (1886: 2). See also Woodward Institutional Abbreviations (1888: 297; 1889: 132; 1907: 194-195, pl. VII, fi gs. 4-5; 1912: 229, text fi g. 73); Zittel (1887: LAGESE - Laboratory of Sedimentary Geology, ý Federal University of Pernambuco, Brazil; 101), Branner (1890: 429), Pelack (1895: 2), DGEO-CTG-UFPE - Geology Department, Fed- Jordan (1907: 215), Fossa-Mancini (1921: 210), eral University of Pernambuco; Casier (1953: 34-36), Cappetta (1987: 171, fi g. UFRPE - Biology Department, Rural Federal 144 D-H), Silva Santos & Gomes (1987: 38-47; University of Pernambuco; pl. 1, fi gs. 1-5), Silva (1994: 317-319, pl.1, fi gs. MN - Museu Nacional, Universidade Federal do 1-3), Silva et al. (1995), Gallo et al. (2001: 269, Rio De Janeiro; fi g. 8; 271), Silva et al. (2002: 7), Cappetta (2006: ChM - Charleston Museum, Charleston, South 277), Silva (2007: 43-44, 71-74, text fi g. VI.17, pl. Carolina, USA; II, fi g. 4; pl. VII, fi gs. 2-3), Silva et al. (2007: 88), NHM - Natural History Museum, London, Eng- Santana et al. (2009: 10), Cleason et al. (2010: land. 660, 666, 668, fi g. 8C). Type locality and stratigraphic horizon – Near Paulista city, Paraíba Basin, Pernambuco, Materials and Methods Brazil; Maria Farinha Formation; Paleocene Se- The new Brazilian specimen was mechani- ries, Danian Stage. cally prepared to free it from limestone matrix General remarks – Cope named A. sericeus and then photographed. South Carolina Apo- in an address to the American Philosophical So- copodon specimens were recovered by screen- ciety on 17 April, 1885, but the description of washing the fossiliferous lag deposit with sieves the type material did not appear in print until down to 0.25 mm. We have limited our analysis the January, 1886 Transactions of the American of Apocopodon sericeus to the articulated denti- Philosophical Society (vol. 23, no. 121). As such, tions described herein because other material is the year attributed to the naming of the genus often imperfectly preserved, and we have also and type species is variously listed as 1885 (e.g. found that Apocopodon teeth are very similar Cappetta, 1987; 2006) or 1886 (Silva, 2007). The in gross morphology to lateral teeth of Rhinop- correct date should appear as 1886 according to tera prisca, a taxon also occurring in the Maria Article 8.4 of the International Code of Zoologi- Farinha Formation. Cope’s collection of ablated cal Nomenclature. Depictions of the type material specimens was discussed by Silva (1994), and did not appear until 1907, when Woodward illus- her descriptions of the fossils were taken into trated the fossils (pl. VI, fi gs. 4-5) as part of a dis- account in our diagnosis of Apocopodon. She cussion on Cretaceous fossils from Brazil. Cope’s listed as syntypes MN 2555/1-V (isolated tooth; specimens do not actually reside at MN, but at pl. 1, fi gs. 2A-B) and MN 2555/2-V (three articu- the NHM where Woodward (e.g. 1907) examined lated teeth; pl. 1, fi gs. 1A-B), and an additional them, and specimens have two catalog numbers specimen consisting of four articulated teeth associated with them; MN 2555/2-V is the same was included (MN 2555/3-V; pl. 1, fi gs. 3A-B). as NHM P24670. Material examined – LAGESE V-00014, lingual portion of dentition; DGEO-CTG-UFPE 5680, partial dentition; DGEO-CTG-UFPE SN (not curated), partial dentition; UFRPE 3223, nearly complete dentition; ChM VP7709, partial dentition.

Diagnosis – Teeth of Apocopodon sericeus Individual teeth are robust, with crowns of form a pavement-type dentition that consists unworn teeth in medial and paramedial rows of longitudinal rows of teeth. The fi rst row of reaching nearly 2 cm in thickness. The labial teeth is herein called the medial row, and this and lingual crown faces are lingually inclined row was presumably located on the jaw sym- and very coarsely wrinkled parallel to crown physis. A line drawn labio-lingually along the height (fi gure 4E-H), the net effect of these fea- middle of this row will divide the dentition into tures being that the lingual face of the crown more or less symmetrical right and left halves of one tooth overlaps the labial face of the suc- (depending on the number of lateral tooth rows ceeding tooth, and the teeth are very strongly actually preserved). The second tooth row, here- interconnected. The wrinkles on the crown in called the paramedial row, directly adjoins faces merge with fi ne ridges along the occlu- the medial row and is therefore the fi rst lateral sal margins, and the entire occlusal surface of tooth row within the dentition. The remaining unworn teeth bears fi ne ornamentation of bi- lateral rows were numbered in ascending order furcating and anastomosing longitudinal ridges towards the lateral margins of the dentition in a (six ridges per millimeter), overall having a ver- similar way that Leriche (1913) labeled his Hy- micular or even reticulated pattern. The occlu- polophites mayombensis dentition. The second sal crown margin of unworn teeth is somewhat lateral row, L2, articulates with the paramedial rounded, and the contacts between teeth within row mesially and L3, the third lateral row, dis- the unworn portion of the dentition are seen as tally. LAGESE V-00014 and DGEO-CTG-UFPE straight to slightly sinuous furrows. However, SN preserve a single tooth in L4 (= fourth lateral tooth contacts within the worn portion of the row; see fi gure 2A, D and 4A, D), and the mesial dentition have an interfi ngering, sutured ap- sides of these teeth articulate with L3, but the pearance. distal end appears to mark the lateral-most edge Tooth bases are low, only one third the height of the dentition. On all remaining dentitions, of the crowns of medial and paramedial teeth. the distal ends of the teeth in L3 have well de- The labial crown face extends forward of the veloped angles, indicating they articulated with base, and in lingual view the crown is separated at least one additional row of teeth that is not from the base by a thick, rounded and irregular preserved. transverse ridge. In addition, bases are polyau- The teeth in medial and paramedial rows are locorhize, with up to seven nutritive grooves six-sided and generally the largest within the occurring on medial and paramedial teeth, but dentition, being approximately 1.5 times wider generally only two nutritive grooves in more than long. The paramedial teeth are asymmetri- lateral rows (fi gure 2B, 3B, 4B). Some tooth bas- cal with a more rhomboidal outline (when com- es exhibit nutritive grooves that are open at one pared to teeth in the medial row), and within end but closed at the other (fi gure 5), and this the dentition it is apparent that their distal end condition could be described as either vestigial is situated closer to the labial margin of the or even “secondarily roofed over”, a condition plate (e.g. fi gure 2F, 3A). Tooth size decreases also reported on Igdabatis (Cappetta, 1972) and distally, with teeth in L2 being roughly as wide Meridiania (Cicimurri, 2010). as long. Although six-sided, teeth within the remaining L3 and L4 are asymmetrical in be- Description ing longer than wide and having a more rectan- LAGESE V-00014 is the well preserved lingual gular outline. The teeth of each row articulate portion of an upper(?) dental plate that con- with those of adjacent rows via lateral angles, sists of 22 articulated complete and fragmen- and the angles of medial and paramedial teeth tary teeth. The teeth are arranged into eight are approximately 90 degrees (and located near longitudinal rows, but a ninth row appears to the transverse midline on medial teeth). be missing from one side (fi gure 2A, C). Four

Figure 2. Dental plates of Apocopodon sericeus. A-E, LAGESE V-00014. A) Occlusal; B) Basal; C) Schematic drawing of occlusal view; D) Labial; E) Enlargement of area within rectangle of D showing crown ornamentation. F-J, ChM VP7709; F) Occlusal; G) Basal; H) Proﬁ le; I) Schematic drawing in occlusal view; J) Enlargement of area within rectangle of G showing serrated contacts between teeth. Labial is at top except in D and J (right). Abbreviations: l. para., left paramedial row; l. L2, left second lateral row; l. L3, left third lateral row; l. L4, left fourth lateral row; r. para., right paramedial row; r. L2, right second lateral row; r. L3, right third lateral row; r. L4, right fourth lateral row.

© PalArch Foundation 7 Santana et al., New material of Apocopodon sericeus PalArch’s Journal of Vertebrate Palaeontology, 8(6) (2011) teeth (two complete and two partials) are pre- dimensions to the medial teeth, but they have served in the medial row, whereas three com- an obvious rhomboidal appearance in occlusal plete teeth and fragments of a fourth are pre- view due to the mesio-labial and disto-lingual served in both paramedial (fi rst lateral) rows. margins being longer than the mesio-lingual Two complete teeth and two partials make up and disto-labial margins. The lateral angles are the left L2, whereas two complete teeth are pre- sharp on both sides of the crown. Teeth in L2 served in the equivalent row on the right side. are six-sided but have a nearly trapezoidal oc- Two teeth are preserved in L3 of both sides. clusal outline (fi gure 2F, I). Tooth crowns mea- Only one tooth is preserved in a right L4, and sure up to 0.5 cm in thickness. In basal view, the this appears to mark the distal margin of the tooth roots are very worn (a result of post-mor- dentition (fi gure 2A, C). tem abrasion) but obviously polyaulocorhize, In occlusal view, the crown surfaces bear with medial teeth bearing at least three nutri- fi ne longitudinal vermicular ornamentation, tive grooves, at least four grooves on paramedi- and the contacts between teeth appear to be al teeth, and at least two grooves on teeth in L2 smooth (fi gure 2A, E). From the broken labial (fi gure 2G, J). The root is 2-3 mm in height and end, the occlusal surface is convex but the basal located closer to the lingual crown margin (the surface is less so. The crowns of medial teeth labial crown face overhangs the root). ChM are three times the height of the root, with PV7709 (see fi gure 2F) is morphologically indis- height diminishing towards the commissure, tinguishable from UFRPE 3223 (see fi gure 3A), where crown height approximately equals root aside from the additional rows of lateral teeth height (fi gure 2D). In basal view, three deep (ap- preserved on the latter specimen. proximately half of the thickness of the base) UFRPE 3223 also appears to represent an up- nutritive grooves divide the bases of medial per dentition, and it is formed of 45 teeth pre- and paramedial teeth into four lobes, whereas served in six longitudinal rows. There are nine there are only two grooves (three lobes) on the teeth in the medial and both paramedial rows, bases of teeth within the remaining lateral rows whereas there are eight teeth in the left L2 but (fi gure 2B). The nutritive grooves represent the only seven remaining on the right side (fi gure locations where blood vessels entered the tooth. 3A, C). A L3 is preserved on the left side that con- ChM VP7709 appears to represent an upper tains six teeth (fi gure 3B). Overall, the oral sur- dentition that, although abraded, is remarkably face is convex and the basal surface correspond- complete, consisting of 36 articulated teeth (fi g- ingly concave (fi gure 3D), but the anterior one ure 2F, G, I). Only fi ve tooth rows are preserved, third of the dentition is concave where the teeth and eight teeth comprise the medial row. Nine have been worn down through in vivo usage (fi g- teeth (seven complete and two partials) are ure 3A). Although occlusal surfaces are abraded, preserved in the left paramedial row, whereas crowns within the (unworn) lingual portion of seven teeth (fi ve complete and two partials) the dentition are much thicker than those within make up the right paramedial row. Within L2, the worn labial portion (fi gure 3D). seven teeth are preserved on the left side and Tooth bases of medial teeth bear three nutri- fi ve on the right. The teeth are tightly packed tive grooves (four lobes), but nutritive groove together and form a rigid battery. Overall the formation on paramedial teeth is asymmetrical, dentition is convex in profi le, but the labial half with three grooves developed on teeth on one is rather fl at to weakly concave (fi gure 2H, F, reside and four grooves (fi ve lobes) developed on spectively). The crowns of medial teeth are six- the opposite side (fi gure 3B). Nutritive groove sided, wider than long (1.2 cm x 0.72 cm; 1.7:1 development within adjoining L2 is also asym- ratio), with sharp lateral angles that are near metrical, with three grooves (four lobes) found perfect 90 degrees (fi gure 2F). The labial crown on one side and two grooves (three lobes) on margin is weakly convex, whereas the lingual the opposite side (in the opposite order as in margin is weakly concave. Teeth in the parame- the paramedial teeth). Two nutritive grooves are dial row are six-sided and very nearly equal in found on the bases of teeth within L3.

Figure 3. Dental plates of Apocopodon sericeus. A-D, UFRPE 3223. A) Occlusal; B) Basal; C) Schematic drawing in occlusal view; D) Profi le. E-G, DGEO-CTG-UFPE 5680; E) Occlusal; F) Basal; G) Schematic drawing in occlusal view; H) Profi le. Labial is at top in all views. Abbreviations as in fi gure 2.

DGEO-CTG-UFPE 5680 consists of the labial The number of nutritive grooves on tooth half of what may be a lower dentition. This spec- bases varies within the medial row, ranging imen is composed of seven longitudinal rows from three grooves (four lobes) labially to of teeth (fi gure 3E-G). Five teeth are preserved fi ve grooves (six lobes) lingually (fi gure 4B). within the medial row, whereas the paramedial This trend of increasing numbers of nutritive rows contain six teeth on the right side and fi ve grooves is also seen on the left paramedial row, on the left (fi gure 3G). Within L2, six teeth are but in the right paramedial row the opposite preserved on the right side and fi ve teeth on the is true – nutritive grooves decrease from six to left side. Five teeth are preserved within L3 on four labio-lingually. Within the remaining lat- of both sides (fi gure 3E-G). In profi le the oral eral rows, the number of grooves decreases dis- surface is convex and the basal surface is con- tally from three to two (fi gure 4B). The increase cave (fi gure 3H). The medial teeth are nearly in the number of nutritive grooves within the perfectly hexagonal, whereas the paramedial medial row and left paramedial row also coin- teeth, although also six-sided, are approximate- cides with an increase in crown and tooth base ly twice as wide and oriented with their distal width. ends located closer to the labial margin of the As mentioned previously, lateral teeth of dentition (fi gure 3E, G). Teeth in L2 are slightly Rhinoptera prisca are very similar to Apo- smaller but even more asymmetrical, having a copodon teeth, and DGEO-CTG-UFPE 6158 and roughly rhomboidal outline, and teeth within 6177 are noted here in order to help differenti- L3 are six-sided but very asymmetrical (fi gure ate the two species. DGEO-CTG-UFPE 6117 was 3F). The anterior half of the preserved dentition identifi ed as A. sericeus by Silva (2007), but we is highly worn (in vivo usage), whereas the re- believe that it actually represents R. prisca. This maining half is unworn (fi gure 3E). specimen (fi gure 4E-H), although reminiscent of There are three nutritive grooves (four lobes) A. sericeus, is identical to paramedial teeth in a on bases of medial teeth (fi gure 3F). Nutritive very well preserved, nearly complete upper den- groove formation is asymmetrical on paramedial tition of R. prisca (LAGESE V-00013; currently teeth, with fi ve grooves (six lobes) on the bases under study). We consider DGEO-CTG-UFPE of the right row, but seven grooves (eight lobes) 6158 (fi gure 4I-4L) to belong to A. sericeus, and on the left row. Within L2, there are three nu- distinguishing features may be: 1) it has a more tritive grooves (four lobes) on the right row, but rhomboidal occlusal outline, whereas R. prisca four (fi ve lobes) on the left row. Tooth bases in L3 is more trapezoidal (fi gure 4G vs. fi gure 4K); have two to three grooves (three to four lobes). 2) the lingual tongue of the tongue-and-groove DGEO-CTG-UFPE SN is presumed to be an interlocking mechanism (sensu Cleason et al., upper dentition, and the teeth are distributed in 2010) on Rhinoptera teeth is a straight, rounded eight longitudinal rows. There are fi ve complete a ridge, whereas in Apocopodon it is highly ir- and two incomplete teeth remaining within the regular (fi gure 4F vs. fi gure 4J); 3) tooth bases of medial row, whereas six teeth are preserved in Apocopodon have fewer nutritive grooves and the left paramedial row and eight teeth on the wider laminae (see Claeson et al., 2010) than right. In L2, there are fi ve teeth on the left side equivalent teeth of R. prisca (fi gure 4H vs. fi g- and at least seven on the right side. Within L3, ure 4L), and this is particularly true in medial there are three teeth on the left side and six on teeth; 4) the occlusal ornamentation of R. prisca the right. Only one tooth is preserved in a left is less robust than on Apocopodon. L4 (fi gure 4A-C). The oral surface of the plate is convex and the basal surface is concave (fi gure Discussion 4D). The medial teeth and most of the paramedial teeth are six-sided, roughly equidimension- Dentition al, with their labial margins being convex as op- Previous descriptions of Apocopodon sericeus posed to straight (fi gure 4A, B). Within L2, all dentitions suggested that the contacts between crowns are six-sided, but one row of teeth is ap- the teeth would be serrated (e.g., Silva Santos & proximately twice as wide as the opposite row Gomes, 1987; Silva 1993; 1994; Santana et al., (fi gure 4C). The anterior one third of the plate 2009). Our analysis of Apocopodon dentitions is worn (in vivo usage), whereas the remaining shows that, in occlusal view, the teeth of the un- two thirds is unworn (fi gure 4A). worn lingual portion of the dentition appear to

Figure 4. A-C, dental plate of Apocopodon serieus, DGEO-CTG-FPE SN. A) Occlusal; B) Basal; C) Schematic drawing in occlusal view; D) Proﬁ le. E-H, isolated Rhinoptera prisca tooth, DGEO-CTG-FPE. E) Labial; F) Lingual; G) Occlusal; H) Basal views. I-L, isolated A. sericeus lateral tooth, DGEO-CTG-FPE 6177. I) Labial; J) Lingual; K) Occlusal; L) Basal views. Labial at top in A-D, G-H, K-L. Abbreviations as in Figure 2.

© PalArch Foundation 11 Santana et al., New material of Apocopodon sericeus PalArch’s Journal of Vertebrate Palaeontology, 8(6) (2011) be united by smooth contacts (Santana et al., larger teeth form at the lingual-most margin of 2009), whereas the worn labial portion exhib- the dentition). These features indicate that the its serrated contacts (e.g. fi gure 2A, 3A, 3E, 4A). number of nutritive grooves is an uninforma- The basal surface of ChM VP7709 is free of ma- tive characteristic in diagnosing Apocopodon trix, and the specimen demonstrates that all of (e.g., Silva Santos & Gomes, 1987; Silva, 1994). the teeth are in fact united by serrated contacts (fi gure 2G, J). The lingual portion of the occlu- Taxonomic Remarks sal surface appears to have smooth contacts Woodward (1888; 1889; 1907; 1912) consis- between teeth because the unworn teeth have tently speculated that Apocopodon represents a a beveled edge, which eventually wears down transitional species between Cretaceous Ptycho- to reveal the highly interdigitating wrinkles on dus (a taxon that became extinct by the early the crown faces (resulting in serrated contacts Campanian) and Cenozoic Myliobatidae. This as seen in fi gure 4A). This same phenomenon is hypothesis may have been infl uenced by the observable on the Rhinoptera prisca upper den- belief that Apocopodon was a Cretaceous taxon tition (LAGESE V-00013) that we are studying. (see Cope, 1886; Jordan, 1907) and the fact that It is likely that the interdigitating crown faces the crown faces of Apocopodon teeth are heav- increased the strength and rigidity of the denti- ily wrinkled. However, the resemblance to Pty- tion to resist the forces exerted during mastica- chodus is at best only superfi cial and a result tion (see below). of convergent evolution, and there is no close Previously unnoted in the labial part of Apo- phylogenetic relationship between Ptychodus copodon dentitions, the tooth crowns may be (Hybodontiformes?, Ptychodontidae) and Apo- approximately equal in thickness to the tooth copodon (Myliobatiformes, Myliobatidae). base, whereas crowns within the lingual part Cleason et al. (2010) recently conducted a are two and one half times the thickness of the phylogenetic analysis of Myliobatiformes, and base (LAGESE V-00014; fi gure 2D). This char- the dental characters they utilized led them to acteristic of decreasing crown height towards conclude that Apocopodon is the sister taxon to the labial margin is seen in DGEO-UFRPE 3223 “Crown-Myliobatidae” (which includes, among (fi gure 3D), CTG-UFPE 5680 (fi gure 3H), and others, Rhinoptera, Myliobatis, Weissobatis, Ae- DGEO-CTG-UFPE SN (fi gure 4D), as well as the tobatus). Their analysis relied solely on three R. prisca upper dentition (LAGESE V-00013) articulated teeth described by Cope (1886), and previously mentioned. Crown height therefore we are currently utilizing the fi ve Apocopodon is not of taxonomic importance, and in our dentitions reported herein to test Cleason et opinion crown thickness is related to the pre- al.’s (2010) conclusion. This will be presented sumed diet of Apocopodon (see below). as a companion to our study of the Rhinoptera Other interesting phenomena we observed prisca upper dentition (LAGESE V-000130) pre- include the number of apparently incomplete viously mentioned. nutritive grooves present on the tooth bases Based solely on our observations, Apo- (e.g. fi gure 5), as well as the asymmetrical de- copodon is fi rmly grounded within Mylioba- velopment of grooves between equivalent rows tidae because: 1) the pavement-like dentition on the right and left sides of the dentition (e.g. consists of six-sided teeth; 2) medial teeth are fi gure 4B). At present we are unsure if the in- expanded (although only slightly); 3) teeth do complete nutritive grooves represent a develop- exhibit tongue-and-groove articulation; 4) tooth mental abnormality, the initial formation of ad- bases are polyaulocorhize (e.g., Cleason et al., ditional grooves to improve blood supply to the 2010; see also Silva Santos & Gomes, 1987; Sil- developing tooth, or if the groove is partially va, 1993; 1994). A close phylogenetic relation- “secondarily roofed over” (blood vessel partially ship between Apocopodon and Rhinoptera was enclosed by dentine). As noted above, the lin- suggested by Fossa-Mancini (1921), and Casier gual increase in the number of nutritive grooves (1953) later indicated that Apocopodon could on teeth within some rows of a dentition (e.g. be an evolutionary intermediate between Cre- DGEO-CTG-UFPE SN) also coincides with an taceous Rhombodus and Cenozoic Rhinoptera. increase in crown and tooth base width. Our in- Although there are similarities in tooth mor- terpretation is that this refl ects ontogeny, with phology between these three taxa, the ances- tooth size increasing as the animal grew (new, tor-descendent hypothesis advocated by Fossa-

Figure 5. Detail of LAGESE V-00014 in basal view showing partially closed nutritive groove. Labial is at top.

Mancini (1921) and Casier (1953) may have The characteristics outlined above indicate relied more on the belief that Apocopodon was a that the Apocopodon dentition is more simi- Cretaceous taxon that occurred stratigraphical- lar to Rhinoptera than to Myliobatis (all three ly between Rhombodus and Rhinoptera. Howev- genera occur in the Maria Farinha Formation). er, Apocopodon is an early Paleocene taxon that Preliminary examination of the upper R. prisca was coeval with Rhinoptera in both Brazil and dentition (LAGESE V-00013) revealed that the South Carolina. lateral teeth are very similar in size, shape and Morphological features that Apocopodon crown thickness to those of Apocopodon, dem- shares with Rhinoptera (Rhinopterinae) include onstrating the potential difﬁ culty of accurately lateral teeth that are thicker mesially than they identifying isolated teeth to the generic level. are distally and, although the teeth in the me- When comparing the R. prisca dentition to dial and paramedial rows are generally nearly Apocopodon, it is obvious that R. prisca medial equal in size, the teeth in remaining lateral rows teeth are wider (3.12 cm vs. 2 cm). of the Apocopodon dentition decrease in width Histological analyses may prove to be use- towards the commissure (observable in the R. ful in separating morphologically similar tooth bonasus [SC 88.120.1] and R. brasiliensis [LAG- types within Myliobatidae, and in their analysis ESE uncurated] we examined; see also Bigelow of Myliobatis sp. cf. M. dixoni from the Maria & Schroeder, 1953; Cappetta, 1987; Purdy et al., Farinha Formation, Silva et al. (2002) found 2001). In contrast, tooth sizes within all lateral that the histology of Myliobatis differs from rows of a Myliobatis californica (BCGM uncurat- Rhinoptera. Silva Santos & Gomes (1987) dis- ed specimen) dentition are rather consistent to- cussed the tooth histology of A. sericeus, but wards the commissure (see Bigelow & Schroed- their description is very similar to what Silva et er, 1953 for other examples within Myliobatis). al. (2002) reported in Rhinoptera. This leads to

© PalArch Foundation 13 Santana et al., New material of Apocopodon sericeus PalArch’s Journal of Vertebrate Palaeontology, 8(6) (2011) the possible conclusions that Apocopodon is in- the Caribbean, where warm water circulated to deed very closely related to Rhinoptera, or Silva North America (see Haq, 1984; Pinet & Popenoe, Santos & Gomes (1987) actually thin-sectioned 1985). Alternatively, mollusk species within the specimens of Rhinoptera and not Apocopodon. Maria Farinha Formation have affi nities to the Additional histological analysis of Apocopodon Gulf of Mexico and North America (Barbosa, teeth appears to be warranted. 2004), and southward migration of Apocopodon It is interesting to note here that the up- is a possibility to consider. The timing and di- per and/or lower dentitions known for Hypo- rection of migration of Apocopodon may be re- lophites, a genus recently placed as the sister solved when the source strata of Apocopodon taxon to Myliobatidae, have a comparable tooth fossils are identifi ed in South Carolina and dat- morphology, dental arrangement and plate size ed using biostratigraphic markers. to Apocopodon (see Stromer, 1910; Leriche, The Apocopodon-bearing strata within the 1913; White, 1934; Cappetta, 1972). However, Maria Farinha Formation were deposited in an the two genera can be differentiated at least inner littoral, probably deltaic environment (< on the basis that all of the teeth within a Hy- 20 m depth) where water temperature was be- polophites dentition have holaulocorhize bases, tween 25° and 29° C (Silva & Silva, 2007; see also whereas medial and paramedial teeth of Apo- Silva, 2007; Silva et al., 2007). If indeed derived copodon are polyaulocorhize, and only the teeth from the Rhems Formation, the South Carolina in the more distal lateral rows are holauloco- Apocopodon also likely inhabited a nearshore rhize (e.g. L3). marine, deltaic environment (Erickson, 1998; Sawyer, 1998; Hutchinson & Weems, 1998; Paleobiology Weems & Bybell, 1998). Apocopodon sericeus was considered to be Tooth articulation within the Apocopodon endemic to Brazil (Silva et al., 2007) because dentition is complex, with the lingual crown the taxon was heretofore not known to occur face overlapping the labial face of the suc- outside of Pernambuco State, and within this ceeding tooth, and the extensive longitudinal region the ray only occurs within the Maria wrinkling on crown faces allows for increased Farinha Formation. Discovery of Apocopodon surface contact area between teeth through in- in South Carolina was therefore a surprise and terdigitating of the wrinkles. This results in a extends the paleogeographic distribution north- serrated suture and is visible in basal view and wards by more than 7,000 km. The occurrences within the worn labial region of the occlusal demonstrate the dispersion potential of elas- surface. Sutured contacts are not visible at the mobranch species and show that faunal inter- unworn lingual ends of the dentitions because change between these widely separated areas the margins of the occlusal surface appear occurred during the Paleocene. This appears to slightly beveled (e.g. LAGESE V-00014), result- be a pattern continuing at least from the latest ing in a V-shaped furrow between teeth. The Cretaceous because the elasmobranch species serrated contacts become visible as teeth are Rhombodus binkhorsti, Squalicorax pristodon- worn down through occlusion (see below). In tus and Cretalamna biauriculata, occur in these addition, there is an irregular transverse lingual two regions. The discovery of three coeval ba- ridge located at the crown-root junction, and toids in the Maria Farinha Formation (and at the labial crown base has a correspondingly ir- least three early Paleocene taxa in the Kingstree regular furrow. This furrow overlaps the lingual lag deposit) also highlights the rapid diversifi ca- ridge of the preceding tooth. Overlapping and tion of the group shortly after the K/T extinc- highly interdigitating contacts between teeth tion. could explain the good sample of partial and Barbosa et al. (2008) suggested that after nearly complete Brazilian plates, and survival the K/T extinction dyrosaurid (longirostrine) of the nearly complete South Carolina plate crocodilians could have reached North America through at least one episode of reworking. by fi rst crossing the Atlantic Ocean from west- Except for LAGESE V-00014, the lingual part ern Africa to northeastern Brazil. Apocopodon of a dentition, the specimens we examined all is currently unknown from Africa, but the ge- display an arched morphology (seen in profi le). nus could have reached South Carolina via the This arching refl ects the way in which the den- northeastern coastline of Brazil (fi gure 6) into tition wrapped over the jaw cartilages and was

Figure 6. Paleogeographic map showing early Paleocene (Danian) continental distribution (dark gray) and sea level (light gray). Stars indicate Apocopodon occurrences in Brazil (fi ve-pointed star) and South Carolina (four-pointed star), and arrows indicate possible dispersal route of marine vertebrates between these regions (modifi ed from Barbosa et al., 2008). attached via connective tissue. Unfortunately, teeth of nearly the same dimensions, would be we are not sure if the preserved arching repre- upper dentitions. sents the original state or the effects of tapho- The morphology of the Apocopodon denti- nomic processes (e.g. was the arching more tion indicates that the symphyses of the palato- pronounced or less?). In the jaws of Rhinoptera quadrates (upper jaws) and Meckel’s cartilages bonasus (SC 88.120.1), R. brasiliensis (LAGESE (lower jaws) were labio-lingually broad and uncurated) and Myliobatis californica (BCGM fused, a condition observed on the M. califor- uncurated) that we examined, the upper denti- nica, R. bonasus and R. brasiliensis jaws we ex- tions are more tightly arched than the lower amined. Extensive ablation is evident on the dentitions, and the lower medial teeth are not labial portion of four Apocopodon dentitions as wide as the upper medial teeth. If this latter (LAGESE V-00014 represents only the lingual characteristic can be applied to Apocopodon, portion of a dentition). The occlusal surfaces then the nearly symmetrical medial teeth of of the teeth in this area have a rough appear- DGEO-CTG-UFPE 5680 indicate that Gallo et ance because the enameloid is partly or com- al. (2001) were correct in considering the plate pletely worn away (see especially UFRPE 3223 to represent a lower dentition. The remaining and DGEO-CTG-UFPE 5680; fi gure 3A and D, specimens, which have medial and paramedial respectively), and this region is also noticeably

© PalArch Foundation 15 Santana et al., New material of Apocopodon sericeus PalArch’s Journal of Vertebrate Palaeontology, 8(6) (2011) concave. Extant members of Myliobatidae use face of the succeeding tooth, and the labial edge their dentitions to crush and grind up various of the crown base overlaps a lingual transverse shelled invertebrate prey animals like crusta- ridge at the crown/root junction. The occlusal ceans and mollusks, which results in a simi- surface of unworn teeth is slightly convex and lar wear pattern (e.g. Randall, 1967; Cappetta, ornamented with longitudinal anastomosing 1987; Summers et al., 1998; Jardas et al., 2004; and bifurcating ridges, resulting in a vermicular Sasko et al., 2006). We concur with previous and/or reticulated texture. Tooth bases are poly- interpretations that Apocopodon was a duro- aulocorhize, with up to ﬁ ve nutritive grooves phagous predator and the wear seen on den- on medial teeth, seven grooves on paramedial titions is the result of occlusion between the teeth, and two to three grooves in subsequent upper and lower batteries during food process- lateral rows. The anterior one third to one half ing (see Summers, 2000; Gallo et al., 2001). We of four of the dentitions we studied are heavily may also infer that, based on studies of extant worn, with the occlusal surface being concave Myliobatidae having comparable tooth plate and with a rough texture. This wear indicates morphologies, that the dentition-bearing sec- that Apocopodon was a durophagous predator tions of Apocopodon jaws were reinforced by and the dentition was used to crush/grind the multiple layers of calciﬁ ed cartilage at the jaw shells of prey animals during feeding. surface, as well as internal struts (Summers et al., 1998; Summers, 2000). Acknowledgments

Conclusions Thanks go to LAGESE (UFPE) and Human Re- sources Program of the PRH-26/ANP/FINEP for Apocopodon was once considered to be endem- providing logistical and fi nancial support for ic to the Paraíba Basin of Pernambuco, Brazil, the studies carried out in the Paraíba Basin. Al but the discovery of fossils in South Carolina, Sanders (ChM) kindly allowed us to study the USA extends the paleogeographic distribution South Carolina Apocopodon dentition and iso- of this ray more than 7,000 km to the north. lated teeth, and James Knight (South Carolina Apocopodon seems to have been a short-lived State Museum, Columbia) and Chuck Ciampa- taxon because fossils have only been found in glio (Wright State University, Celina, Ohio) pro- deposits of known or suspected early Paleocene vided DJC access to additional isolated teeth un- (Danian) age (Maria Farinha and Rhems for- der their care. Shawn Ham kindly provided DJC mations). It is possible that the apparently fast with a copy of the Fossa-Mancini (1921) paper. evolution of early Cenozoic Myliobatiformes Jean Self-Trial (US Geological Survey, Reston, and distribution of Apocopodon is related to the Virginia) examined the indurated matrix refi lling of ecological niches left vacant after the covered from between the tooth bases of ChM K/T extinction. Tooth shape and organization of VP7709. We are greatful for the editorial sug- the dentition indicates that Apocopodon can be gestions of Kerin Cleason and an anonymous re- placed within Myliobatidae and that the genus viewer, whose efforts helped refi ne and improve has a closer affi nity to Rhinopterinae than to upon an earlier version of this manuscript. Myliobatinae or Mobulinae. Apocopodon has a crushing/grinding den- Cited Literature tition composed of at least eight longitudinal rows of six-sided (up to 1.5 times wider than Albertão, G.A., E.A.M. Koutsoukos, M.P.S. long) and thick-crowned (over 1 cm high) teeth, Regali, M. Attrep, Jr. & P.P. Martins, Jr. and each row consisted of at least up to nine 1994. The Cretaceous-Tertiary teeth. The teeth are very strongly articulated boundary in southern low-latitude re- to each other via longitudinal wrinkles that gions: preliminary study in Pernambuco, interfi nger with each other, forming a tightly northeastern Brazil. – Terra Nova 6, 4: 366- packed and very rigid dentition (contacts be- 375. tween teeth actually appear serrated in basal Barbosa, J.A. 2004. Evolução da Bacia da Paraí- view and in regions of the dentition that are ba durante o Maastrichtiano-Paleoceno: highly worn). In addition, the lingual crown formações Gramame e Maria Farinha, NE face is inclined such that it overlaps the labial do Brasil. Dissertação de Mestrado. Centro

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