Content uploaded by David John Ward
Author content
All content in this area was uploaded by David John Ward on Jul 24, 2015
Content may be subject to copyright.
Additions to and a review of the Miocene Shark and Ray fauna of Malta.
... Teeth of P. benedenii have been reported from world regions as disparate as Europe (Belgium, Germany, Hungary, Italy, Malta, the Netherlands, Portugal, Slovakia and Switzerland), Africa (Angola and South Africa), Macaronesia (the Azores and the Canary Islands), North America (USA, along both the eastern and western coasts of the country, and Mexico), South America (Ecuador and Peru), Indonesia (Sumatra) and the western Pacific (South Korea, Japan, New Caledonia; Australia and New Zealand) [3,4,[17][18][19][20][21], as well as from the floor of the Indian and Pacific oceans, where this otherwise uncommon taxon appears to be surprisingly abundant [22,23]. As regards the peri-Mediterranean area, finds of P. benedenii come from the Miocene of Malta [24], southern Spain [25] and southern Italy [6,26], as well as from the Pliocene of the Balearic Islands [17] and many Italian localities [27]. In particular, the Pliocene marine deposits of Tuscany (central Italy) have yielded teeth of P. benedenii for a very long time. ...
... Consisting of 114 teeth, the largest of which is slightly shorter than 60 mm (viz the height of GAMPS-00876b), this tooth set allowed Kent and Powell [56] to reconstruct the dentition of P. benedenii with 14 upper files (including a reduced intermediate) and 13 lower files. Emendations to this scheme focusing on the identification of symphyseal and intermediate teeth have been proposed by Purdy et al. [15], Ward and Bonavia [24] and Kent [4]. The same North Carolinian tooth set was used by Kent [57] to extrapolate a total body length of 7.6 m based on the reconstructed upper jaw perimeter. ...
The extinct “false mako” shark, Parotodus benedenii (Lamniformes: Otodontidae), is essentially known from large, robust teeth that are widespread but overall rare in Oligocene to Pliocene deposits worldwide. More than 150 years after its description, this species still represents a palaeontological conundrum, as very little is known about its body aspect and palaeoecology. Here, we describe new specimens of P. benedenii from the Pliocene of Tuscany, central Italy. These new finds comprise some of the geologically youngest finds of P. benedenii worldwide, witnessing to the survival of false makos until the Late Pliocene at least, which in turn suggests that P. benedenii may have been the latest surviving member of the family Otodontidae. Building upon a thorough literature review, we provide an updated synthesis of the palaeobiology of P. benedenii. In light of the morphological evidence, and considering previously published suggestions, P. benedenii may be reconstructed as a large-sized, carnivorous shark that dwelt in pelagic settings and fed primarily on large, soft prey and scavenging items. Thus, some ecological partitioning did likely exist between P. benedenii and other elasmobranch apex predators of the Neogene mid-latitude seas (including, in Pliocene times, the extant species Carcharodon carcharias, Carcharhinus leucas and Galeocerdo cuvier).
... Although the identity of the other teeth mentioned by Woodward (1889) remains a mystery, the occurrence of G. aduncus elsewhere is generally restricted to the Miocene (see Cappetta, 2012). Woodward (1889) also reported the occurrence of Galeocerdo contortus Gibbes, 1849 teeth within Eocene deposits in Clarke County, Alabama, but this species was subsequently referred to the genus Physogaleus by Ward & Bonavia (2001). In the same work, Woodward (1889) reported teeth from Eocene deposits in Alabama that he referred to "Galeocerdo(?) minor." ...
Herein we describe a small but relatively diverse assemblage of fossil fishes derived from the lower Oligocene (Rupelian) Red Bluff Clay at site AMo-9 in Monroe County, Alabama, USA. Identified amongst the remains are 15 unequivocal taxa representing 11 families within five orders, and one additional taxon represents an unknown order and family. Taxa identified include Eostegostoma sp., Otodus (Carcharocles) sp., Mitsukurinidae/Carchariidae indet., Macrorhizodus praecursor, Galeorhinus sp., Negaprion gilmorei, Physogaleus sp., “Sphyrna” sp., Galeocerdo sp., cf. “Aetobatus” sp., Sphyraena sp., Xiphiorhynchus kimblalocki, Xiphiorhynchus sp., Cylindracanthus ornatus, and C. rectus. Several additional fossils could not be identified beyond Lamniformes, Carcharhiniformes, and Teleostei, but they likely belong to one of the identified taxa within this paleofauna. All of the fishes previously reported from the Red Bluff Clay within the entirety of the Gulf Coastal Plain of the USA are otolith-based, and each of the 15 unequivocal taxa reported herein are important new records for this lithostratigraphic unit. In particular, the Eostegostoma sp. and Xiphiorhynchus spp. specimens represent the first occurrences of these taxa in Alabama. The specimens of C. ornatus, Eostegostoma sp., and X. kimblalocki are stratigraphic and temporal range extensions from the middle and late Eocene into the Rupelian Stage of the Oligocene. Other described taxa may represent transitional forms between those described from the late Eocene and late Oligocene within the region. This study provides a tantalizing preliminary view into faunal transitions that occurred amongst marine fishes across the Eocene/Oligocene boundary within the Gulf Coastal Plain of the USA.
... Whether H. serra is the direct ancestor to the Recent H. elongata (Klunzinger, 1871) is questionable. Based on histological differences of the teeth compared to those of extant H. elongata (Klunzinger, 1871), Ward and Bonavia (2001) Based on δ 66 Zn composition, H. serra from the Early Miocene of Malta occupied a higher trophic position than individuals from the Early Miocene of Baden-Württemberg, Germany. This is the same relative result recovered for individuals of Carcharodon hastalis between the two localities; different prey availability or a shorter trophic chain in the German Molasse Basin may also be driving the pattern in this case. ...
In the last years, new findings and new methods (stable isotopes of oxygen, zinc and nitrogen, 2D and 3D modelling, geometric morphometric analyses of the teeth) have enhanced our knowledge of the Neogene shark fauna and its palaeobiology. Several papers deal with the large Otodus (Megselachus) species, including the construction of a 3D model as well as insights into lifestyle and diet. In addition, skeletal remains of Carcharias gustrowensis, Carcharodon hastalis, Keasius parvus and a natural tooth set of Carcharodon hubbelli have been described in the last 13 years, and the dentition of the Neogene species Carcharoides catticus, Megachasma applegatei and Parotodus benedenii have been reconstructed. Stable isotope analyses of the teeth from the Neogene species of Araloselachus, Carcharias, Carcharodon, Galeocerdo, Hemipristris, and Mitsukurina have given insights into the trophic position of these genera during the Neogene, and shark teeth preserved near skeletal remains of prey animals (mammals) and shark bite traces on these remains provide direct evidence of trophic interactions. Tooth shape, fossil locality and palaeoenvironment have been used to better understand the taxa Carcharhinus dicelmai, Megalolamna paradoxodon, Pachyscyllium dachiardii and P. distans. Among extant species, Galeorhinus galeus can be traced back to the Eocene. The following taxa can be traced back to the Oligocene: ?Alopias superciliosus, and Rhincodon typus. Species already present in the Miocene include: Alopias vulpinus, Carcharhinus amblyrhynchoides, C. amblyrhynchos, C. albimarginatus, C. amboinensis, C. brachyurus, C. brevipinna, C. falciformis, C. glaucus, C. leucas, C. limbatus, C. longimanus, C. macloti, C. obscurus, C. perezi, C. sealei, ?Carcharodon carcharias, Centrophorus granulosus, Cetorhinus maximus, Dalatias licha, Deania calcea, Galeocerdo cuvier, , Glyphis glyphis, Heptranchias perlo, Isurus paucus, Lamna nasus, Negaprion brevirostris, Odontaspis ferox, Pseudocarcharias kamoharai, Sphyrna media, S. mokarran. First appearing in the Pliocene are: Scymnodon ringens, Somniosus rostratus, Zameus squamulosus. For some extant species (Carcharias taurus, Hexanchus griseus, Isurus oxyrinchus, Notorynchus cepedianus, Sphyrna zygaena) it is not clear if the assigned Neogene teeth represent the same species. Applying these new methods to more fossil shark taxa, a detailed search for shark fossils, as well as better knowledge of the dentition of extant species (especially those with minute-sized teeth) will further enhance knowledge of the evolution and palaeobiology of sharks.
... In older literature, the genus Isurus was attributed to the species hastalis and planus. While, the generic assignment of these closely-related species remains in flux (see Purdy et al. 2001;Ward and Bonavia 2001;Ehret et al. 2012;Kriwet et al. 2015;Yun 2022), for the time being, we follow Ehret et al. (2012) and Kent (2018) by considering it most parsimonious to place these shark species in the genus Carcharodon. C. hastalis had a global distribution and occurred from the late Oligocene to the lower Pliocene (Purdy et al. 2001;Kent 2018) whereas Carcharodon planus comb. ...
... 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). ...
... That the first Italian fossils of A. grandis come from the outer shelf deposits of the Pietra leccese and "Aturia level" is thus not surprising. Other finds of A. grandis are known from the Miocene of the Mediterranean Basin, and specifically from the "Middle Globigerina Limestone" (Aquitanian -Burdigalian; Foresi et al. 2007) of Malta (Kent 2018), which was also deposited in a rather farshore, open-sea (i.e., upper slope) setting (Ward & Bonavia 2001). ...
... Bor et al., 2012;Everaert et al., 2019), as teeth of both species are morphologically very similar (e.g. Cappetta & Nolf, 1991;Ward & Bonavia, 2001). For a long time, these teeth were attributed to Carcharias acutissima (Agassiz, 1843) (e.g. ...
A large fragment of driftwood was discovered in the marine Terhagen Member (Boom Formation, NP23) at Schelle (Belgium), representing the first well-documented case of wood-fall in the Rupelian of the North Sea Basin. This trunk with a side-branch, identified as Cupressinoxylon sp. (Cupressaceae), caused a large irregularity on the sea bottom, creating a unique microenvironment which allowed colonization by some taxa virtually absent elsewhere in the Boom Formation. The fossils were further concentrated in a silty lens against the trunk by the effects of prolonged wave-driven turbulence. This lens comprised a large set of compartmental plates of the turtle barnacle Protochelonibia hermani Gale sp. nov., possibly part of a single colony originally attached to a turtle. The material includes the best preserved plates of Protochelonibia known to date, yielding new information on the construction of its shell. Additionally, a disarticulated tooth set of 154 teeth of Carcharias contortidens (Agassiz, 1843) was found, the first such discovery in more than 100 years. An articulated dentition of this taxon, initially studied by Leriche (1910), is refigured herein. Some very rare valves of the bivalve Palliolum permistum (Beyrich, 1848) are identified and the gastropod Amblyacrum cf. roemeri (von Koenen, 1867) is reported here for the first time from the Belgian Rupelian. The teleost otolith assemblage comprises ca 30,000 specimens belonging to 11 species only, of which Trachurus reineckei Hoedemakers sp. nov. is new to science and Myoxocephalus primas (Koken, 1891) and Capros siccus Schwarzhans, 2008 are new for the Belgian Rupelian. The new species represents the earliest record of the thermophilic genus Trachurus in the Oligocene of the North Sea Basin. Liparis minusculus Nolf, 1977 is synonymized with Myoxocephalus primas, whereas Erythrocles ohei Schwarzhans, 1994 is transferred to the genus Trachurus.
Early Eocene batomorph faunas, represented by isolated teeth from the London Clay Formation, LCF (Division D, lower Sheppey Member, lower NP12) at Burnham-on-Crouch, Essex, UK, and the Tielt Formation (Egemkapel Clay Member) and overlying Hyon Formation (Egem Sand Member, middle to upper NP12) at Egemkapel, western Flanders, Belgium, are described and illustrated. Whereas the Burnham fauna is from an outer neritic, deep-water environment, the faunal assemblages recovered from different stratigraphic levels in the Egemkapel Clay and Egem Sand Members are characteristic of mid to inner neritic settings. We also examined a small batomorph fauna from the upper levels of the Roubaix Clay Member of the Kortrijk Clay Formation exposed in the Koekelberg Clay Pit at Marke, Belgium. The upper Roubaix Clay Member is considered penecontemporaneus with Division D of the LCF, but was deposited in a mid-neritic environment. In all, we identified 13 batomorph taxa in the Sheppey Member, 11 in the Roubaix Clay Member, 24 in two different beds of the Egemkapel Clay Member, and 22 in five beds of the Egem Sand Member. Seven new genera are proposed: Glaucopristis gen. nov., Essexraja gen. nov., Casierabatis gen. nov., Sheppeytrygon gen. nov., Serratodasyatis gen. nov., Belgabatis gen. nov., and Eurasiabatis gen. nov. Three new species are described: Essexraja ypresiensis gen. et sp. nov., Casierabatis lambrechtsi gen. et sp. nov., and Eurasiabatis occlusostriata gen. et sp. nov. Based on detailed comparisons with the dentition and dental morphology of living stingrays, four species previously included in the genus “Dasyatis” are re-allocated to new dasyatoid genera: Casierabatis jaekeli (Leriche, 1905) comb. nov., Sheppeytrygon davisi (Casier, 1966) comb. nov., Serratodasyatis tricuspidata (Casier, 1946) comb. nov., and Belgabatis thierryi (Smith, 1999) comb. nov. The rhinopristiform “Rhinobatus” bruxelliensis Jaekel 1894 is transferred to the new genus Glaucopristis. A stratigraphic range chart for batomorph taxa in the late Paleocene, Ypresian, and Lutetian of the North Sea Basin is presented. This chart may be considered as a starting point for additions and updates in future works, because identifications of some small-toothed rhinopristiform and dasyatoid species reported in the literature appear doubtful and the stratigraphic coverage of
several species, especially those from outer neritic habitats, is insufficiently known.
In recent years, new findings and new methods (stable isotopes of oxygen, zinc, and nitrogen; 2D and 3D modeling; and geometric morphometric analyses of the teeth) have enhanced our knowledge of the Neogene shark fauna and its paleobiology. Several papers deal with the large Otodus (Megaselachus) species, including the construction of a 3D model, as well as insights into its lifestyle and diet. In addition, the skeletal remains of Carcharias gustrowensis, Carcharodon hastalis, and Keasius parvus and a natural tooth set of Carcharodon hubbelli have been described in the last 13 years, and the dentition of the Neogene species Carcharoides catticus, Megachasma applegatei, and Parotodus benedenii has been reconstructed. Stable isotope analyses of the teeth from the Neogene species of Araloselachus, Carcharias, Carcharodon, Galeocerdo, Hemipristris, and Mitsukurina have given insights into the trophic positions of these genera during the Neogene, and shark teeth preserved near the skeletal remains of prey animals (mammals) and shark bite traces on these remains provide direct evidence of trophic interactions. The tooth shape, fossil locality, and paleoenvironment have been used to better understand the taxa Carcharhinus dicelmai, Megalolamna paradoxodon, Pachyscyllium dachiardii, and P. distans. Among extant species, Galeorhinus galeus can be traced back to the Eocene. Alopias superciliosus, Rhincodon typus, and possibly A. vulpinus can be traced back to the Oligocene. Species present by the Miocene include Alopias vulpinus, Carcharhinus amblyrhynchoides, C. amblyrhynchos, C. albimarginatus, C. amboinensis, C. brachyurus, C. brevipinna, C. falciformis, C. glaucus, C. leucas, C. limbatus, C. longimanus, C. macloti, C. obscurus, C. perezi, C. sealei, Centrophorus granulosus, Cetorhinus maximus, Dalatias licha, Deania calcea, Galeocerdo cuvier, Glyphis glyphis, Heptranchias perlo, Isurus paucus, Lamna nasus, Negaprion brevirostris, Odontaspis ferox, Pseudocarcharias kamoharai, Sphyrna media, S. mokarran, and possibly Carcharodon carcharias. First appearing in the Pliocene are Scymnodon ringens, Somniosus rostratus, and Zameus squamulosus. For some extant species (Carcharias taurus, Hexanchus griseus, Isurus oxyrinchus, Notorynchus cepedianus, and Sphyrna zygaena), it is not clear whether the assigned Neogene teeth represent the same species. The application of new methods to more fossil shark taxa, a detailed search for shark fossils, and better knowledge of the dentition of extant species (especially those with minute-sized teeth) will further enhance our knowledge of the evolution and paleobiology of sharks.
ResearchGate has not been able to resolve any references for this publication.