Redescription of two lungfish (Sarcopterygii: Dipnoi) tooth plates from the Late Cretaceous Bauru Group, Brazil
Introduction
The dipnoans, or lungfishes, form a well-defined clade of sarcopterygian fishes with a fossil record since the Early Devonian. Today, they are represented by just three extant freshwater genera distributed in three continents: Lepidosiren (South America), Protopterus (Africa) and Neoceratodus (Australia) (e.g., Schultze, 1992).
Fossil dipnoans in the Paleozoic are well known from articulated specimens, including three-dimensional skulls and isolated tooth plates, sometimes attached to mandibles and maxillary bones (Denison, 1968). There are a few rare exceptions of articulated Mesozoic specimens from the Jurassic of Kyrgyzstan (Nessov and Kaznyshkin, 1985), the Cretaceous of Thailand (Cavin et al., 2007), and several specimens from Australia (White, 1926; Kemp, 1993, 1994, 1996, 1998); most of the post-Triassic fossil record is composed exclusively of tooth plates (e.g., Kemp, 1997b).
This notable shift in the preservation of fossil dipnoans through time is directly related to the observed evolutionary trend, from heavily ossified skeletons with dermal bones, sometimes covered by cosmine, in the Devonian dipnoans, to the weakly ossified skeletons with thin dermal skull bones embedded in the skin present in Mesozoic and Cenozoic dipnoans (Martin, 1982). In parallel, tooth plates became stronger through increasing mineralization with specialized dentines (Martin, 1982; Kemp, 2001; Cavin et al., 2007).
Despite the relatively frequent occurrence of lungfish tooth plates in South America (e.g., Toledo, 2006; Apesteguía et al., 2007; Cione et al., 2007; Soto and Perea, 2010), only six nominal species were identified in Brazil: Ceratodus africanus Haug, 1905 (see Cunha and Ferreira, 1980; Martin, 1984); Lepidosiren megalos Silva Santos, 1987; Asiatoceratodus cf. tiguidiensis (see Dutra and Malabarba, 2001; Castro et al., 2004); Ceratodus humei Priem, 1914 (see Toledo, 2006) (for a discussion of the taxonomy of this species, see Churcher and De Iuliis, 2001); Ptychoceratodus cf. philipsi (see Richter and Toledo, 2008); and Equinoxiodus alcantarensis Toledo et al. (2011).
The lack of morphological data, apart from tooth plates, makes difficult a precise classification for much of the post-Triassic dipnoan fossil record. The recent discoveries of new material in the Mesozoic, specifically in eastern and western Gondwana (e.g., Churcher and De Iuliis, 2001; Castro et al., 2004; Churcher et al., 2006; Cavin et al., 2007; Cione et al., 2007; Gottfried et al., 2009; Soto and Perea, 2010; Toledo et al., 2011), allowed a more accurate review of some dipnoan fossil groups (e.g., Kemp, 1994; Churcher et al., 2006; Apesteguía et al., 2007; Cavin et al., 2007; Cione et al., 2007; Agnolin, 2010; Soto and Perea, 2010). Based on these reviews, we present here a redescription of two specimens collected during fieldwork in the Adamantina and Marília formations (Fig. 1; the localities of Santo Anastácio and Uberaba towns, respectively), of the Bauru Group. Both specimens were originally identified by Gayet and Brito (1989) as Neoceratodus sp. (also published by Brito et al., 2006). However, a re-examination of the specimens provided new anatomical information that allows them to be assigned to other genera (see Discussion).
Section snippets
Geological settings
The specimens described here were collected in 1988 from two localities within the Bauru Basin: Loc. 99, near Santo Anastácio municipality, São Paulo state, and at km 24 on the highway linking Uberaba and Uberlândia in Minas Gerais state (Fig. 1; see also Bertini et al., 1993).
The Bauru Basin was developed in the southeastern part of the South American platform through thermo-mechanical subsidence during the Late Cretaceous (Fernandes and Coimbra, 1996). This continental basin accumulated an
Material and methods
The specimens described here comprise two partially preserved dipnoan tooth plates that are housed in the Universidade do Estado do Rio de Janeiro, under the numbers UERJ-PMB 156 and DGM 1315-P. They are compared with other genera that have been described in the literature. Measurements and terminology follow Churcher and De Iuliis (2001). The following diagnostic criteria were used: dimension, shape and disposition of ridges and mesiolingual angle. We follow Kemp (1997b) in regarding tooth
Systematic paleontology
Sarcopterygii Romer, 1955
Dipnoi Müller, 1845
Ceratodontiformes Berg, 1940
Asiatoceratodontidae Vorobyeva, 1967
Asiatoceratodus Vorobyeva, 1967
Asiatoceratodus sp.
Fig. 2A, B
Type species. Asiatoceratodus sharovi Vorobyeva, 1967, Early Triassic, Russia.
Material. DGM 1315-P, right lower partially preserved tooth plate from the lower part of Adamantina Formation at Loc. 99 (see Bertini et al., 1993) close to the town of Santo Anastácio, São Paulo State and about 584 km west of the city of São Paulo.
Discussion
The phylogenetic interrelationships and taxonomy of Mesozoic lungfishes remains are problematic, and a comprehensive synthesis of the diversity of the group and its paleogeographical distribution during Mesozoic has yet to be achieved (e.g., Cavin et al., 2007; Gottfried et al., 2009; Agnolin, 2010). As observed above, the robust development of specialized dentines (petrodentine of any derived dipnoans, and also change of organization of dentine compared to other derived dipnoans without
Conclusions
Both Ceratodus and Asiatoceratodus are shown to have been present in the Late Cretaceous Bauru Group of Brazil, thus improving our understanding of the global distribution of these taxa. The occurrence of asiatoceratodontids in the Adamantina Formation indicates that this taxon survived in South America longer than previously thought; it extends the temporal limit of this taxon from Cenomanian to Turonian–Santonian. We document the first occurrence of Ceratodus in the Maastrichtian of South
Acknowledgments
We thank Federico Agnolin (Museo Argentino de Ciencias Naturales) and an anonymous reviewer for suggestions that led to improvements of the manuscript as well as David Martill (University of Portsmouth) and Miguel Furtado (Universidade Federal do Rio de Janeiro) for improving the English. We are grateful to Leonardo Borghi and the LAGESED (Laboratório de Geologia Sedimentar/UFRJ) staff, and Thiago Marinho (UFRJ) for help with photography, Carlos Eduardo Vieira Toledo for information about South
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