APPENDIX
Below, we provide a list of all elasmobranch species found in the studied fossil sites, with information regarding their tooth morphology alongside palaeoecological data gleaned from the literature of both fossil and recent related forms. Taxonomic information is also provided when necessary.
The species are presented in systematic order (classification scheme follows Compagno et al., 2005, Cappetta, 2012), with genera and species within the same family ordered alphabetically for ease of use.
The trophic level (TL) data for the modern taxa were delineated by Cortés (1999) and also as available from Froese and Pauly (2019). For skates (Raja spp.), the work of Ebert and Bizzarro (2007) was consulted.
ORDER HEXANCHIFORMES De Buen, 1926
Family Chlamydoselachidae Garman, 1884
Genus Rolfodon Cappetta, Morrison and Adnet, 2021
Rolfodon bracheri (Pfeil, 1983)
Tooth type: Clutching.
This species was originally placed within the recent genus Chlamydolselachus Garman, 1884. However, Cappetta et al. (2021) transferred the species to the extinct genus Rolfodon Regardless of that, both genera have similar trident-shaped teeth and Chlamydoselachus seems to be the obvious nearest recent relative of Rolfodon. There are two recent species of Chlamydoselachus: C. anguineus Garman, 1884, living at depths between 17 and 1520 m. (Ebert et al., 2021) and C. africana Ebert and Compagno, 2009, living from 300 to 1400 m depth (Ebert et al., 2021). Both these species have very similar teeth (see illustrations in Ebert et al., 2021: p. 97). For C. anguineus , Cortés (1999) defined a TL of 4.2 and listed cephalopods as its staple food. Ebert and Compagno (2009) and Ebert et al. (2021) listed members of the Scyliorhinidae and bony fishes as diet for both species. According to the latter authors, C. anguineus also feeds on midwater crustaceans.
Family Hexanchidae Gray, 1851
Genus Heptranchias Rafinesque, 1810
Heptranchias sp.
Tooth type: Cutting-clutching.
Only one living species is known in this genus, namely H. perlo (Bonnaterre, 1788), which lives from 0 to 1000 m depth (Ebert et al., 2021). Cortés (1999) assigned it a TL of 4.2, mentioning bony fishes and cephalopods as staple foods.
Genus Paraheptranchias Pfeil, 1981
Paraheptranchias repens (Probst, 1879a)
Tooth type: Cutting-clutching.
The genus is extinct. Based on tooth morphology, the species is most similar to the recent Heptranchias perlo (see Heptranchias sp. above).
Genus Notorynchus Ayres, 1855
Notorynchus primigenius Agassiz, 1835
Tooth type: Cutting-clutching.
The only extant relative is N. cepedianus (Péron, 1807), which very often feeds on other elasmobranchs such as Myliobatis spp., Dasyatis spp. and Squalus spp. (Ebert, 1991). Other prey items are bony fishes, squids (Loligo spp.), octopuses, crustaceans, gastropods, marine mammals (cetaceans and pinnipeds), agnathans and carrion (Ebert, 1991, 2003, Ebert et al., 2021). According to Cortés (1999), marine mammals and elasmobranchs are the staple food for this species, which has a TL of 4.7. This shark is the dominant elasmobranch in the nearshore marine environment, living from the surf line (less than 1 m depth) to at least 570 m depth, but mostly in less than 100 m depth (Ebert, 2003, Ebert et al., 2021).
ORDER SQUALIFORMES Goodrich, 1909
Family Echinorhinidae Gill, 1862
Genus Echinorhinus Blainville, 1816
Echinorhinus pfauntschi Pfeil, 1983
Tooth type: Cutting-clutching.
There are two recent species of Echinorhinus: E. cookei Pietschmann, 1928 and E. brucus (Bonnaterre, 1788), living from 4 to 1100 m and from 10 to 1214 m depth (Ebert et al., 2021), respectively. The TL of E. cookie is circa 4.4 ±0.76 (Froese and Pauly, 2019) and that of E. brucus is 4.4 (Cortés, 1999). Both bony and cartilaginous fishes have been indicated as staple foods of the recent species (Cortés, 1999, Ebert et al., 2021).
Family Squalidae Blainville, 1816
Genus Squalus Linnaeus, 1758
Squalus sp.
Tooth type: Cutting.
Ebert et al. (2021) listed 34 recent Squalus species, the biology of some of which are poorly known. Most extant species include bony fishes and several kinds of invertebrates in their diet (Last et al., 2007; Ebert et al., 2021). Cortés (1999) assigned TL values between 3.9 and 4.2 to five Squalus species, all with bony fishes as staple food. The members of this genus can be found from 0 to more than 1500 m depth (Ebert et al., 2021).
Family Centrophoridae Bleeker, 1859
Genus Centrophorus Müller and Henle, 1837
Centrophorus sp.
Tooth type: Cutting-clutching.
The Centrophorus teeth from the OMM were sometimes identified as belonging to Centrophorus cf. granulosus (Bloch and Schneider, 1801) (Pfeil., 1991, Vialle et al., 2011, Pollerspöck and Beaury, 2014; Pollerspöck et al., 2022). Centrophorus granulosus is a recent species, which feeds mainly on bony fishes and cephalopods (Cortés, 1999), has a TL of 4.1 (Cortés, 1999), and lives in depths from 50 to 1500 m, mostly deeper than 600 m (Ebert et al., 2021).
Family Etmopteridae Fowler, 1934
Genus Etmopterus Rafinesque, 1810a
Etmopterus sp.
Tooth type: Cutting-clutching.
Ebert et al. (2021) listed 42 recent species of Etmopterus , the biology of most of which is poorly known. Cortés (1999) listed nine recent species, with TL values ranging from 3.8 to 4.2, which have bony fishes and cephalopods as their staple food. Representatives of this genus can be found from 0 to more than 2490 m depth (Ebert et al., 2021).
Family Somniosidae Jordan, 1888
Genus Centroscymnus Barbosa Du Bocage and De Brito Capello, 1864
Centroscymnus sp.
Tooth type: Cutting-clutching.
Two extant species are known: C. coelolepis Barbosa Du Bocage and De Brito Capello, 1864 (128 and 3675 m deep) and C. owstoni Garman, 1906 (150 to 1459 m deep) (Ebert et al., 2021). Both species live on or near the bottom (Ebert et al., 2021) and have diets composed of bony fishes and cephalopods, resulting in a TL of 4.2 (Cortés, 1999).
Family Dalatiidae Gray, 1851
Genus Isistius Gill, 1865
Isistius triangulus (Probst, 1879)
Tooth type: Cutting-clutching.
There are only two extant species within this genus: I. brasiliensis (Quoy and Gaimard, 1824) and I. plutodus Garrick and Springer, 1964. According to de Figueiredo Petean and de Carvalho (2018), members of this genus may not be good swimmers, given that they are ectoparasites of large fishes and cetaceans. Wounds made by Isistius spp. have been found on marlins, mackerels, tunas, sharks, rays, seals, whales, and dolphins (de Figueiredo Petean and de Carvalho, 2018). Strasburg (1963) also described squid retrieved from stomach contents and Cortés (1999) listed cephalopods as staple prey of I. brasiliensis. Extant I. brasiliensis has a TL of. 4.3 (Cortés, 1999), whereas I. plutodus has a TL of 4.2 ±0.4 (Froese and Pauly, 2019). Isistius spp. live in the oceanic realm, from epipelagic to bathypelagic regions (Ebert et al. 2021). For details on the feeding apparatus of the recent I. brasilianensis see Shirai and Nakaya (1992).
Squaliolus sp.
Both recent species, S. aliae Teng, 1959 and S. laticaudus Smith and Radcliffe, 1912, feed on squids and bony fishes (Ebert et al., 2021); the former has a TL of 4.4 ±0.57 and the latter, 4.2 ±0.73 (Froese and Pauly, 2019). S. aliae lives from 0 to 2000 m deep and S. laticaudus from 10 to 750 m (Ebert et al., 2021).
ORDER PRISTIOPHORIFORMES Berg, 1958
Family Pristiophoridae Bleeker, 1859
Genus Pristiophorus Müller and Henle, 1837
Pristiophorus suevicus Jaekel, 1890
Tooth type: Clutching.
There are seven recent species within this genus, some of which with unknown biology (Ebert et al., 2021). Those whose biology is known feed on small fishes and crustaceans (Ebert et al., 2021). The members of this genus can be subdivided into two groups by habitat (Ebert and Wilms, 2013): one of temperate waters and mostly coastal and continental shelf species, and another of tropical, deep-waters, continental and insular slope species, living near the bottom (maximum known depth of 630 m; Compagno et al., 2005). TL values for the seven recent species range from 3.8 to 4.2 (Froese and Pauly, 2019).
ORDER SQUATINIFORMES De Buen, 1926
Family Squatinidae Bonaparte, 1838
Genus Squatina Dumeril, 1806
Squatina sp.
Tooth type: Clutching.
Ebert et al. (2021) listed 22 recent species of Squatina , the biology of some of which are poorly known. All the species with better known biology have bony fishes as their staple food, with TL values ranging from 4.0 to 4.2 (for six of the living species; Cortés, 1999). Squatina spp. live on or near the bottom, from shallow waters to a depth of 1290 m (Ebert et al., 2021).
ORDER ORECTOLOBIFORMES Applegate, 1974
Family Ginglymostomatidae Gill, 1862
Genus Ginglymostoma Müller and Henle, 1837
Ginglymostoma delfortriei Daimeries, 1889
Tooth type: Clutching.
Ginglymostoma cirratum (Bonnaterre, 1788) is the only recent species known, living between 0 and 130 m depth, but mostly at depths shallower than 40 m (Ebert et al., 2021). Its staple foods are crustaceans, bony fishes and cephalopods, and its TL value is 3.8 (Cortés, 1999). For details on the feeding apparatus of the recent Ginglymostoma cirratum see Motta et al. (2008).
ORDER LAMNIFORMES Berg, 1937
Family Odontaspididae Müller and Henle, 1839
Genus Araloselachus Glikman 1964
Araloselachus cuspidatus (Agassiz, 1843)
Tooth type: Tearing.
The genus is extinct. The teeth present a general odontaspidid morphology and can be found nearly worldwide in neritic Oligocene and Miocene sediments. The genus affiliation of this species is under discussion (see Hovestadt 2020). Based on δ66Zn istopes from teeth of the OMM of Rengetsweiler and Walbertsweiler, McCormack et al. (2022) found out that this species was placed much higher in the food pyramid than the other piscivore taxa, together with Carcharias, Pseudocarcharias, Hemipristris, Mitsukurina and also Carcharodon (see also their fig. 2). That conclusion is also supported by the more robust teeth of A. cuspidatus. Furthermore, according to McCormack et al. (2022), A. cuspidatus had a higher trophic level than Hemipristris and Carcharodon from the studied OMM deposits, which would mean that A. cuspidatus hunted fishes that had a higher position in the trophic chain.
Genus Carcharoides Ameghino, 1901
Carcharoides catticus (Philippi, 1846)
Tooth type: Tearing.
The genus is extinct. Teeth of C. catticus are found in neritic sediments and have an odontaspidid morphology, being also similar to those of extant Triaenodon obsesus (Rüppel, 1837) (family Carcharhinidae). Because of this similarity, Purdy et al. (2001) placed Carcharoides in synonymy with T. obsesus , but Ward and Bonavia (2001) later showed that they are different taxa. In spite of the similar tooth shape suggests that C. catticus had a similar diet to T. obsesus , which consists of bony fish and squids (Cortés, 1999; Ebert et al., 2021).
Genus Odontaspis Agassiz, 1838
Odontaspis molassica (Probst, 1879a)
Tooth type: Tearing.
As already mentioned, there are two recent species of Odontaspis, O. noronhai and O. ferox; the teeth of the latter are most similar to O. molassica (Bass et al., 1975; Pfeil, 1991). According to Pfeil (1991), O. molassica is the ancestor of O. ferox (Risso, 1810). The latter species feeds on small bony fishes, including rockfish, as well as squid and shrimp, living between 10 and 1015 m depth (Ebert et al., 2021) and having a TL value of 4.2 ±0.56 (Froese and Pauly, 2019). Odontaspis ferox has a cosmopolitan distribution across warm-temperate and tropical waters, and although it is essentially demersal, it has also been captured pelagically in the mid-ocean (Fergusson et al., 2008).
Family Carchariidae Müller and Henle, 1838
Genus Carcharias Rafinesque, 1810
Carcharias contortidens (Agassiz, 1843)
Tooth type: Tearing.
The only recent species is C. taurus Rafinesque, 1810, which has very similar teeth to the above fossil taxon (Reinecke et al., 2011). It feeds mainly on bony fishes and elasmobranchs, including members of the families Carcharhinidae, Dasyatidae, Hexanchidae, Rajidae, and Rhinobatidae (Cortés, 1999; Smale, 2005; Lucifora et al., 2009). It lives near the bottom from 1 to 232 m depth, but mostly between 15 to 25 m (Ebert et al., 2021). TL for recent C. taurus is 4.4 (Cortés, 1999).
Family Pseudocarchariidae Taylor, Compagno and Struhsaker, 1983
Genus Pseudocarcharias Cadenat, 1963
Pseudocarcharias kamoharai (Matsubara, 1936)
Tooth type: Tearing.
The OMM fossils have been assigned to P. kamoharai , which is an extant species. It lives offshore (occasionally closer to the shore), from the surface down to at least 590 m depth (Compagno, 1984; Nelson, 2006; Ebert et al., 2021). It feeds on small pelagic bony fishes, squids as well as shrimps (Compagno 1984; Estupiñán-Montaño and Galván-Magaña 2021) and has a TL value of 4.2 ±0.60 (Froese and Pauly, 2019).
Family Mitsukurinidae Jordan, 1898
Genus Mitsukurina Jordan, 1898
Mitsukurina lineata (Probst, 1879)
Tooth type: Tearing.
The only living species is M. owstoni Jordan, 1898, which feeds primarily on bony fishes (Yano et al., 2007), having a TL of 4.1 ±0.62 (Froese and Pauly, 2019). This species lives mostly offshore, from 100 to 1300 m depth, rarely approaching the surface (Compagno, 1984; Yano et al., 2007; Ebert et al., 2021). According to Ebert (2003), M. owstoni appears to forage away from the bottom and may in fact occupy more of a midwater habitat than generally assumed.
Family Alopiidae Bonaparte, 1838
Genus Alopias Rafinesque 1810
Alopias exigua (Probst, 1879)
Tooth type: Tearing.
Out of the three recent species of Alopias listed in Ebert et al. (2021), the teeth of A. superciliosus (Lowe, 1840) display the strongest similarity with those of A. exigua (Bigelow and Schroeder, 1948; Bass et al., 1975; Herman et al., 2004; Ebert et al., 2021). Cortés (1999) indicated cephalopods as the main food item of that species, followed by bony fishes, and established a TL of 4.2. Alopias superciliosus lives from close inshore to the open ocean, from the surface to 955 m depth, but mostly over 100 m depth (Compagno et al., 2005, Ebert et al., 2021).
Family Cetorhinidae Gill, 1862
Genus Keasius Welton, 2013
Keasius parvus (Leriche, 1908)
Tooth type: Clutching.
This is an extinct genus of basking sharks. Based on the morphology of teeth and gill rakers, the only recent relative taxon is Cetorhinus maximus (Gunnerus, 1765), a plankton feeder that lives on the continental shelf and slope (Compagno et al., 2005; Ebert et al., 2021). Cetorhinus maximus can dive down to 1264 m depth (Ebert et al., 2021). During summer time, basking sharks can be found near the surface, whereas in winter they move to deeper waters (Compagno et al., 2005). Cetorhinus maximus has a TL of 3.2 (Cortés, 1999).
Family Lamnidae Müller and Henle, 1838
Genus Carcharodon Smith, 1838
Carcharodon hastalis (Agassiz, 1843)
Tooth type: Tearing with tendency towards the cutting type.
According to Collareta et al. (2017b), adults of the extinct species C. hastalis were in large part piscivorous; they lived in tropical to temperate seas worldwide, probably on the inner shelf (Cappetta, 1987; Bor et al., 2012). The recent white shark C. carcharias feeds on bony and cartilaginous fishes as well as on marine mammals (Cortés,1999; Ebert, 2003), having a TL of 4.5 (Cortés, 1999). If C. hastalis also fed on marine mammals can not be said with certainty.
Genus Isurus Rafinesque, 1810
Isurus oxyrinchus Rafinesque, 1810
Tooth type: Tearing.
This is a living species that feeds on bony and cartilaginous fishes, having a TL of 4.3 (Cortés, 1999). Large specimens also feed on small cetaceans (Ebert et al., 2021). This species inhabits the open ocean and coastal waters, from the surface to 888 m depth (Ebert et al., 2021).
Isurus retroflexus (Agassiz, 1843)
Tooth type: Tearing with tendency towards the cutting type.
The genus level of this species is highly debated. A recent species with similar dental morphology is I. paucus Guitart-Manday, 1966, which feeds primarily on schooling fishes and pelagic cephalopods (Ebert, 2003), having a TL of 4.5 (Froese and Pauly, 2019). The blade-like teeth of I. retroflexus , it might also feed on marine mammals. I. paucus is an epipelagic species that can be found from the surface to 1752 m deep (Compagno 1984; Ebert 2003; Ebert et al., 2021).
Family Otodontidae Glikman, 1964
Genus Megalolamna Shimada, Chandler, Lam, Tanaka and Ward, 2016
Megalolamna paradoxodon Shimada, Chandler, Lam, Tanaka and Ward, 2016
Tooth type: Anterior teeth tearing or grasping type, lateral teeth cutting type (Shimada et al. 2016).
In 2016, Shimada et al. designated this new species and genus for teeth from the early Miocene (Aquitanian-Burdigalian) of Japan, Peru and the USA (see Shimada et al. 2016: fig. 2). Pfeil (1991: 200, pl. 2, fig. 8) mentioned and illustrated a tooth from Messkirch-Walbertsweiler as Lamna sp. When comparing the illustration of “Lamna sp.” with the ones of Megalolamna paradoxodon, it became obvious that it belongs to the latter as per the tooth shape and inclination and position of the lateral cusplets. Shimada et al. (2016) assumed bony fishes as prey for this species.
Genus Otodus Agassiz, 1843
Subgenus Megaselachus Glikman, 1964
Otodus (Megaselachus) sp.
Tooth type: Cutting.
Both the genus and the subgenus are extinct, but its species are thought to have inhabited the neritic realm of warm-temperate to tropical oceans worldwide, feeding on marine mammals (Purdy, 1996). Collareta et al. (2017a) proposed that C. megalodon was an apex predator whose trophic spectrum was still focused on small-sized baleen whales. They may also have fed on other sharks and rays, similar to recent Carcharodon carcharias. According to Kast et al. (2022) teeth of Otodus (Megaselachus) megalodon are characterized by a broad range of δ15 N values, which could be explained by the individuals feeding across many prey types and different trophic levels. Also, according to those authors, teeth of O. (Megaselachus) chubutensis and O. (Megaselachus) megalodon display similar isotope signatures. In the OMM, Otodus (Megaselachus) sp. did likely feed at the highest trophic levels.
ORDER CARCHARHINIFORMES Compagno, 1977
Family Scyliorhinidae Gill, 1862
Genus Pachyscyllium Reinecke, Moths, Grant and Breitkreuz, 2005
Pachyscyllium dachiardii (Lawley, 1876)
Tooth type: Clutching.
The genus is extinct. Based on odontological analogies, the diet of this species was similar to that of the extant members of Scyliorhinidae. Ebert (2003) listed small bony fish, small sharks, crabs, squid and other invertebrates as prey items for members of Scyliorhinidae. Jacobsen and Bennett (2013) proposed an average TL of 3.9 for scyliorhinids. Members of this family can be found from shallow waters to 825 m depth (Ebert and Fowler, 2014).
Pachyscyllium distans (Probst, 1879a)
The genus is extinct. See P. dachiardii above for details.
Genus Scyliorhinus Blainville, 1816
Scyliorhinus fossilis (Leriche, 1927)
Tooth type: Clutching.
This genus contains different recent species, some with poorly-known biology (Compagno et al., 2005; Froese and Pauly, 2019). Cortés (1999) mentioned bony fishes, cephalopods and crustaceans as staple food for five of the extant species and gave a TL of 4.0 to four of them [3.6 to S. canicula (Linnaeus, 1758)]. Members of this demersal genus live between 1 and 825 m depth (Ebert et al., 2021).
Scyliorhinus sp.
See S. fossilis above.
Family Triakidae Gray, 1851
Genus Iago Compagno and Springer, 1971
Iago angustidens (Cappetta, 1973)
Tooth type: Clutching.
There are two recent species, namely, Iago garricki Fourmanoir, 1979 and I. omanensis (Norman, 1939) (Ebert et al., 2021), both of which forage on cephalopods (Ebert et al. 2021). Cortés (1999) assigned a TL of. 4.1 to I. omanensis. I. garricki can be found from 250 to 477 m depth and I. omanensis from circa 90 m to possibly 2195 m depth (Ebert et al., 2021).
Iago sp.
See I. angustidens above.
Genus Triakis Müller and Henle, 1838
? Triakis sp.
Tooth type: Clutching.
There are two teeth from Meßkirch-Rengetsweiler that possibly belong to Triakis (see Höltke et al., 2022). They show similarities to recent T. scyllium Müller and Henle, 1839 and T. semifasciata Girard, 1855 (Herman et al., 1988). Both these extant species feature bony fishes and invertebrates in their diet and live near the bottom in the neritic realm (Ebert et al., 2021).
Family Hemigaleidae Hasse, 1879
Genus Chaenogaleus Gill, 1862
Chaenogaleus affinis (Probst, 1879)
Tooth type: Cutting-clutching.
The only recent congener is C. macrostoma (Bleeker, 1852), which lives on continental and insular shelves down to 160 m depth (Compagno et al., 2005). Its biology is poorly known (Ebert et al., 2021), but C. macrostoma probably feeds on small fishes, cephalopods and crustaceans (Compagno, 1984). Froese and Pauly (2019) indicated a TL of 4.2 ±0.57 for C. macrostoma.
Genus Hemipristris Agassiz, 1843
Hemipristis serra Agassiz, 1843
Tooth type: Cutting-clutching.
The only recent congener is H. elongata (Klunzinger, 1871), which feeds on bony fishes and cephalopods and has a TL of 4.3 (Cortés, 1999). It lives on continental and insular shelves down to 132 m depth (Ebert et al., 2021). However, the Miocene Hemipristis serra had larger teeth than extant H. elongata , which may suggest different prey items and, maybe, a slightly higher TL for the extinct species.
Hemipristis sp.
See H. serra above.
Genus Paragaleus Budker, 1935
Paragaleus tenuis (Probst, 1878)
Tooth type: Cutting-clutching.
There are four extant species of Paragaleus , living in the neritic realm. They can be found down to a depth of 100 m (Compagno et al., 2005; Ebert et al., 2021). Only one species, P. pectoralis (Garman, 1906), has a known diet: it is a specialist feeder on cephalopods but may also take small fishes (Ebert et al., 2021). It has a TL of 4.3 ±0.64 (Froese and Pauly, 2019). The three other species as well as the extinct P. tenuis have similarly structured teeth (Probst, 1878, pl. 1, figs. 68-70; Herman et al., 1991, pl. 17 and 18; Pfeil, 1991, pl. 3, fig. 8; Ebert et al., pp. 499-500), hence equivalent feeding habits can be assumed.
Family Carcharhinidae Jordan and Evermann, 1896
Genus Carcharhinus Blainville, 1816
Carcharhinus acuarius (Probst, 1879)
Tooth type: Tearing.
The next recent congener is Carcharhinus oxyrhynchus (Müller and Henle, 1839), which feeds on small schooling fishes and has a TL of 4.5 (Ebert et al., 2021; Froese and Pauly, 2019). This Recent species was previously the type species of the monotypic genus Isogomphodon Gill, 1862. Based on molecular genetics, da Silva Rodrigues-Filho et al. (2023) mentioned Isogomphodon as a synonym of Carcharhinus.
This species lives in turbid waters in estuaries, mangroves and river mouths, as well as over shallow banks, in depths between 4 and 40 m (Ebert et al., 2021).
Carcharhinus priscus (Agassiz, 1843)
Tooth type: Cutting-clutching.
A comparison of C. priscus teeth with the dentition of extant Carcharhinus spp. indicates a close morphological relationship with C. limbatus (Müller and Henle, 1841), C. perezi (Poey, 1876) and C. brachyurus (Günther, 1870) (Reinecke et al., 2011; Bor et al., 2012; Andrianavalona et al., 2015; Collareta et al., 2021). That said, the teeth from those three extant species differ in one or more features from C. priscus (Bor et al. 2012). According to this morphological affinity, it can be assumed that C. priscus had a similar diet and habits as the aforementioned extant species. Carcharhinus perezi lives inshore, from the surface to 65 m depth, while C. limbatus and C. brachyurus live in- and offshore, from the surface to a depth of 100 m (Voigt and Weber, 2011). All these species feature bony fishes in their diet (Voigt and Weber, 2011). Cortés (1999) also mentioned cephalopods as a second component of the diet of C. brachyurus and C. limbatus , establishing a TL of 4.2 for them. For C. perezi , in turn, a TL of 4.5 ±0.8 has been proposed (Froese and Pauly, 2019).
Carcharhinus similis (Probst, 1978)
Tooth type: Cutting-clutching.
The teeth of C. similis display closer morphological affinities with those of the extant species C. leucas (Valenciennes, 1839) and C. amboinensis (Müller and Henle, 1839), both of which inhabit the neritic realm, from the surface to a depth of 150 m (Reinecke et al., 2011; Voigt and Weber, 2011). The diet of both these species is composed of bony and cartilaginous fishes, and both have a TL of 4.3 (Cortés 1999). Some authors (Cliff and Dudley, 1991a, 1991b; Tillett et al., 2014; Estupiñán-Montaño et al., 2017) provided further detail on the food items of these two species, which appear to include: Aetobatus spp., Dasyatis spp., Carcharhinus spp., Carcharias spp., Isurus oxyrhynchus , Mobula spp., Rhynchobatus spp., Rhizoprionodon spp., Sphyrna spp., Squatina spp. and Squalidae for C. leucas; and Carcharhinidae, Dasyatidae, Gymnuridae, Myliobatidae, Rhinobatidae, Scyliorhinidae, Sphyrnidae and Squatinidae for C. amboinensis.
Genus Physogaleus Cappetta, 1980
Physogaleus contortus (Gibbes, 1849)
Tooth type: Clutching.
The genus Physogaleus is extinct; that said, P. contortus might belong to the extant genus Galeocerdo J.P. Müller and Henle, 1837 (Kent, 1994; Purdy et al., 2001). We follow Reinecke et al. (2011) in assigning this extinct species to Physogaleus. In contrast to Galeocerdo teeth, the teeth of P. contortus have a weaker serration and a slenderer upper part of the crown (see also Bor et al., 2012). Another difference to Galeocerdo is the mesial cutting edge of the P. contortus teeth, which areslightly twisted in a lingual direction. The mentioned slenderer upper part of the crown in particular is most suitable for a diet more specialized on fish, and possibly cephalopods, in contrast to the omnivorous diet of Galeocerdo. The dental morphology of P. contortus matches the clutching type sensu Cappetta (2012) whereas the dentition of Galeocerdo matches the cutting type. The Teeth of P. contortus can be found in neritic sediments (Reinecke et al., 2011).
Physogaleus singularis (Probst, 1878)
Tooth type: Cutting-clutching.
The generic allocation of this species is debated. Based on illustrations from Voigt and Weber (2011) and Ebert et al. (2021), teeth of the extant species of the Carcharhinus sealei-dussumieri group (White 2012) are structured in a similar way to those of P. singularis. That group also contains the recent species C. coatesi (Whitley, 1939) and C. tjutjot (Bleeker, 1852). They live in the shelf region, and their staple food is represented by bony fishes, followed by cephalopods (Cortés 1999; Ebert et al. 2021). According to its tooth shape, the lifestyle and diet of P. singularis was likely similar to those of the aforementioned recent species.
Physogaleus sp.
Tooth type: Cutting-clutching (?).
Baier et al. (2004) listed Physogaleus sp. for Ermingen, but without any illustration, so it is problematic to determine its diet. Considering what already observed for Physogaleus spp., at least bony fishes can be surmised.
Genus Rhizoprionodon Whitley, 1929
Rhizoprionodon fischeuri (Joleaud, 1912)
Tooth type: Cutting-clutching.
The teeth of recent Rhizoprionodon spp. are very similar to the fossil representatives (Springer, 1964; Reinecke et al., 2011). Cortés (1999) mentioned bony fishes as staple food for the five extant species, assigning them TL values ranging from 3.8 to 4.2. Members of this genus live in the shelf region as well as offshore, down to depths of 500 m (Ebert et al., 2021).
Rhizoprionodon sp.
See R. fischeuri above.
Family Galeocerdonidae Poey, 1875
Genus Galeocerdo Müller and Henle, 1837
Galeocerdo aduncus (Agassiz, 1843)
Tooth type: Cutting
The only extant congener is G. cuvier (Péron and Lesueur, 1822), an omnivorous opportunistic feeder that eats crabs, lobsters, horseshoe crabs, gastropods, cephalopods, jellyfish, bony fishes, other elasmobranchs, sea turtles, sea snakes, marine mammals, marine birds, and even carrion of terrestrial mammals (Schwartz, 2000; Ebert, 2003; Dicken et al., 2017; Estupiñán-Montaño et al., 2017). Like in other elasmobranch taxa, diet of G. cuvier changes as the animal grows, shifting from bony fishes and cephalopods in juveniles to larger prey items as size increases (Ebert 2003). Large specimens mostly consume elasmobranchs, sea turtles, marine mammals, sea birds and crustaceans (Ebert 2003). The living tiger shark has a TL of 4.1 (Cortés, 1999), living from the intertidal zone to 1136 m depth, moving closer inshore at night but retreating into deeper waters offshore during daytime (Ebert, 2003; Ebert et al., 2021).
Family Sphyrnidae Gill, 1872
Genus Sphyrna Rafinesque, 1810
Sphyrna integra Probst, 1878
Tooth type: Cutting-clutching.
The teeth of S. integra are closest in morphology to those of the recent S. lewini (Griffith and Smith, 1834) and S. media Springer, 1940 (Bor et al., 2012). The diet of the latter is unknown, but it has been assigned a TL of 4.0 ±0.4 (Ebert et al., 2021; Froese and Pauly, 2019); the former mainly feeds on bony fishes, followed by squids, crustaceans and elasmobranchs (Estubiñán-Montaño et al. 2009, 2021; Bornatowski et al., 2014), and has a TL of 4.1 ±0.5 (Cortés, 1999; Froese and Pauly, 2019). Both species live on the continental shelf, but S. lewini can also be found on insular shelves and adjacent waters down to a depth of 1043 m or more (Ebert et al., 2021). Estubiñán-Montaño et al. (2021) mentioned a strong relation of S. lewini to coastal regions.
Sphyrna laevissima (Cope, 1867)
Tooth type: Cutting-clutching.
In light of tooth morphology, the recent species that is closest to S. laevissima is S. zygaena (Linnaeus, 1758). Because of such similarity, Purdy et al. (2001) placed S. laevissima in synonymy with S. zygaena , but Reinecke et al. (2011) demonstrated that these two taxa are indeed different. Sphyrna zyganea feeds mainly on cephalopods, followed by bony fishes, and has a TL of 4.2 (Cortés, 1999). It lives on coastal to offshore waters, from the surface to 200 m deep, possibly also down to 500 m deep (Ebert et al., 2021).
ORDER INCERTAE SEDIS
Family Megascyliorhinidae Pfeil, 1984
Genus Megascyliorhinus Cappetta and Ward 1977
Megascyliorhinus miocaenicus (Antunes and Jonet, 1970)
Tooth type: Clutching.
The genus as well as the family are extinct. A discussion about this species can be found in Manganelli and Spadini (2019). The teeth show odontological analogies to the Scyliorhinidae. See Pachyscyllium dachiardii above for details.
ORDER RAJIFORMES Berg, 1940
Family Rhinidae Müller and Henle, 1841
Genus Rhynchobatus Müller and Henle, 1837
Rhynchobatus pristinus (Probst, 1877)
Tooth type: Crushing.
All eight extant species of Rhynchobatus live in shallow water down to 70 m depth and they all feature bony fishes and invertebrates in their diet (Froese and Pauly, 2019). Apart from one species with a TL of 4.5, all the others range from 3.4 to 3.6 in TL (Froese and Pauly, 2019).
Family Rhinobatidae Müller and Henle, 1837
Genus Rhinobatos Linck, 1790
Rhinobatos sp.
Tooth type: Crushing.
The recent representatives of Rhinobatos feed on molluscs and crustaceans but will also take small fish (Froese and Pauly, 2019). They live from coastal habitats to offshore regions, down to 350 m depth (Séret et al., 2016). The TL values of the 21 recent species of Rhinobatos range from 3.5 to 4.1 (Froese and Pauly, 2019).
Family Rajidae Blainville, 1816
Genus Raja Linnaeus, 1758
Raja sp.
Tooth type: Crushing (female), Clutching (male).
All recent Raja spp. feed on bony fishes and invertebrates (mostly benthic crustaceans), living on the bottom, from shallow waters down to 800 m depth (Last et al., 2016a). The 24 extant species have TL values between 3.3 and 4.0 (Froese and Pauly, 2019). Ebert and Bizzarro (2007) gave a mean TL value of 3.76 for members of the genus Raja.
ORDER MYLIOBATIFORMES Compagno, 1973
Family Aetobatidae Agassiz, 1858
Genus Aetobatus Blainville, 1816
Aetobatus arcuatus (Agassiz, 1843)
Tooth type: Grinding.
There are five extant species of Aetobatus with different diet habits, albeit all of them include bivalves (White and Last, 2016a). Aetobatus spp. can be between from 1 to 60 m depth (White and Last, 2016a). Four species have a TL of 3.6, but A. narinari (Euphrasen, 1790) has a TL of 4.2 (Froese and Pauly, 2019).
Aetobatus sp.
See A. arcuatus above.
Family Dasyatidae Jordan, 1888
Genus Dasyatis Rafinesque, 1810
Dasyatis probsti Cappetta, 1970
Toothy type: Crushing (female), clutching (male).
There is scarce information on the diet of the recent species of Dasyatis , though all of the, seem to feed on crustaceans (Ebert and Bizzarro, 2007). The 9 recent Dasyatis spp. have been associated to TL values comprised between 3.5 and 4.1 (Froese and Pauly, 2019).
Dasyatis rugosa Probst, 1877
See D. probsti above.
Dasyatis strangulata (Probst, 1877)
See D. probsti above.
Dasyatis sp.
See D. probsti above.
Genus Taeniurops Garman, 1913
Taeniurops cavernosus (Probst, 1877)
Tooth type: Crushing (female), Clutching (male).
The only recent congener, T. meyeni Müller and Henle, 1841, is a benthic organism living between 20 and 60 m depth (Froese and Pauly, 2019). It feeds on bottom fishes, bivalves, crabs and shrimp (Compagno et al., 1989; Froese and Pauly, 2019) and has a TL of 4.2 ±0.69 (Froese and Pauly, 2019).
Family Gymnuridae Fowler, 1934
Genus Gymnura van Hasselt, 1823
Gymnura sp.
Tooth type: Clutching.
Recent Gymnura spp. havedifferent trophic habits, all of which include bony fishes (Yokota et al., 2016). Extant members of this genus live inshore, on the bottom down, to 15 m depth (Yokota et al., 2016), and have TL values ranging from 3.6 and 4.5 (Froese and Pauly, 2019).
Family Myliobatidae Bonaparte, 1835
Genus Myliobatis Cuvier, 1816
Myliobatis sp.
All the extant congeners feed on bivalves and gastropods, as well as on crabs (White and Last 2016c); the TL of the 12 recent species range from 3.2 to 3.6 (Froese and Pauly, 2019). Members of this genus can be found in coastal and offshore waters down to 420 m depth (White and Last, 2016c).
Family Rhinopteridae Jordan and Evermann, 1896
Genus Rhinoptera Jordan and Evermann, 1896
Rhinoptera studeri Agassiz, 1843
Tooth type: Grinding.
Dietary data is only available for two extant species of Rhinoptera, R. bonasus (Mitchill, 1815) and R. brasiliensis Müller, 1836, both of which feed on molluscs (Myers et al., 2007; Last et al., 2016b). That said, Froese and Pauly (2019) assigned TL values to all seven extant species, ranging from 3.2 to 3.8. Members of Rhinoptera live in the benthopelagic zone in the open ocean as well as inshore on the continental shelf, down to a depth of 100 m (Last et al., 2016b).
Rhinoptera sp.
See R. studeri above.
Family Mobulidae Gill, 1893
Genus Mobula Rafinesque, 1810b
Mobula sp.
Tooth type: Clutching.
All recent Mobula spp. feed on zooplankton and have a pelagic lifestyle in coastal and offshore areas (White and Last, 2016b). The 11 species of Mobula have been associated to TL values ranging between 3.1 and 3.9 (Froese and Pauly, 2019).
OTHER TAXA
Other marine taxa apart from sharks and rays are listed in Table 3. Remnants of non-marine taxa (birds, reptiles, and mammals) were also found in the deposits and probably came into the palaeoenvironment as carrion (they are not listed in Table 3, but see Table 2 for references).
