Systematics and Phylogenetic Interrelationships of the Enigmatic Late Jurassic Shark Protospinax annectans Woodward, 1918 with Comments on the Shark-Ray Sister Group Relationship.

The Late Jurassic elasmobranch Protospinax annectans is often regarded as a key species to our understanding of crown group elasmobranch interrelationships and the evolutionary history of this group. However, since its first description more than 100 years ago, its phylogenetic position within the Elasmobranchii (sharks and rays) has proven controversial, and a closer relationship between Protospinax and each of the posited superorders (Batomorphii, Squalomorphii, and Galeomorphii) has been proposed over the time. Here we revise this controversial taxon based on new holomorphic specimens from the Late Jurassic Konservat-Lagerstätte of the Solnhofen Archipelago in Bavaria (Germany) and review its skeletal morphology, systematics, and phylogenetic interrelationships. A data matrix with 224 morphological characters was compiled and analyzed under a molecular backbone constraint. Our results indicate a close relationship between Protospinax, angel sharks (Squatiniformes), and saw sharks (Pristiophoriformes). However, the revision of our morphological data matrix within a molecular framework highlights the lack of morphological characters defining certain groups, especially sharks of the order Squaliformes, hampering the phylogenetic resolution of Protospinax annectans with certainty. Furthermore, the monophyly of modern sharks retrieved by molecular studies is only weakly supported by morphological data, stressing the need for more characters to align morphological and molecular studies in the future.

Are shark teeth proxies for functional traits? A framework to infer ecology from the fossil record.

Modern sharks have an evolutionary history of at least 250 million years and are known to play key roles in marine systems, from controlling prey populations to connecting habitats across oceans. These ecological roles can be quantified based on their functional traits, which are typically morphological (e.g., body size) or behavioural (e.g., feeding and diet). Nonetheless, the understanding of such roles of extinct sharks is limited due to the inherent incompleteness of their fossil record, which consists mainly of isolated teeth. As such, establishing links between tooth morphology and ecological traits in living sharks could provide a useful framework to infer sharks' ecology from the fossil record. Here, based on extant sharks from which morphological and behavioural characteristics are known, the authors assess the extent to which isolated teeth can serve as proxies for functional traits. To do so, they first review the scientific literature on extant species to evaluate the use of shark dental characters as proxies for ecology to then perform validation analyses based on an independent data set collected from museum collections. Their results reveal that 12 dental characters have been used in shark literature as proxies for three functional traits: body size, prey preference and feeding mechanism. From all dental characters identified, tooth size and cutting edge are the most widely used. Validation analyses suggest that seven dental characters - crown height, crown width, cutting edge, lateral cusplets, curvature, longitudinal outline and cross-section outline - are the best proxies for the three functional traits. In particular, tooth size (crown height and width) was found to be a reliable proxy of all three traits; the presence of serrations on the cutting edge was one of the best proxies for prey preference; and tooth shape (longitudinal outline) and the presence of lateral cusplets were among the best indicators of feeding mechanism. Overall, the authors' results suggest that in the absence of directly measurable traits in the fossil record, these seven dental characters (and different combinations of them) can be used to quantify the ecological roles of extinct sharks. This information has the potential to provide key insights into how shark functional diversity has changed through time, including their ecological responses to extinction events.

A diverse assemblage of Ptychodus species (Elasmobranchii: Ptychodontidae) from the Upper Cretaceous of Ukraine, with comments on possible diversification drivers during the Cenomanian.

New isolated teeth from the Upper Cretaceous of Ukraine and belonging to the extinct durophagous shark Ptychodus are described here. The taxonomic identification of the examined material reveals a quite diverse Cenomanian shark fauna which comprised both cuspidate and un-cuspidate species of Ptychodus from the coastal areas at the north-western margin of the Ukrainian Massif. In addition, P. latissimus from the Turonian of Ukraine is reported here for the first time. The revision of the Ukrainian record of Ptychodus revealed that most specimens described here are the oldest so far known from this part of the European Peri-Tethys. Moreover, the present study highlights the co-occurrence of cuspidate and un-cuspidate Ptychodus and a variety of shelled macroinvertebrates, which inhabited coastal and offshore areas of the European epicontinental seas during the Late Cretaceous. The availability of different prey items is proposed here as one of the possible drivers, in addition to abiotic environmental factors, for the diversification of shark tooth morphologies, and possible trophic partitioning between cuspidate and un-cuspidate species of the genus Ptychodus.

Heterodonty and ontogenetic shift dynamics in the dentition of the tiger shark Galeocerdo cuvier (Chondrichthyes, Galeocerdidae).

The lifelong tooth replacement in elasmobranch fishes (sharks, rays and skates) has led to the assemblage of a great number of teeth from fossil and extant species, rendering tooth morphology an important character for taxonomic descriptions, analysing phylogenetic interrelationships and deciphering their evolutionary history (e.g. origination, divergence, extinction). Heterodonty (exhibition of different tooth morphologies) occurs in most elasmobranch species and has proven to be one of the main challenges for these analyses. Although numerous shark species are discovered and described every year, detailed descriptions of tooth morphologies and heterodonty patterns are lacking or are only insufficiently known for most species. Here, we use landmark-based 2D geometric morphometrics on teeth of the tiger shark Galeocerdo cuvier to analyse and describe dental heterodonties among four different ontogenetic stages ranging from embryo to adult. Our results reveal rather gradual and subtle ontogenetic shape changes, mostly characterized by increasing size and complexity of the teeth. We furthermore provide the first comprehensive description of embryonic dental morphologies in tiger sharks. Also, tooth shapes of tiger sharks in different ontogenetic stages are re-assessed and depicted in detail. Finally, multiple cases of tooth file reversal are described. This study, therefore, contributes to our knowledge of dental traits across ontogeny in the extant tiger shark G. cuvier and provides a baseline for further morphological and genetic studies on the dental variation in sharks. Therefore, it has the potential to assist elucidating the underlying developmental and evolutionary processes behind the vast dental diversity observed in elasmobranch fishes today and in deep time.

A new cuspidate ptychodontid shark (Chondrichthyes; Elasmobranchii), from the Upper Cretaceous of Morocco with comments on tooth functionalities and replacement patterns.

The first articulated dentition of †Ptychodus from Africa is described herein. The specimen, likely coming from the Turonian of the Asfla area (Goulmima region, southeastern Morocco), exhibits a well-preserved lower dental plate of a second-level predator. A new species, †P. maghrebianus sp. nov., is erected herein based on this durophagous dentition characterised by imbricated cuspidate teeth. We employed for the first time in †Ptychodus multiple quantitative analyses and statistical parametric and non-parametric tests to process biometrical data taken from articulated, associated and isolated teeth. The quantitative approach (morphospace analysis) is exploited herein to support the traditional taxonomic identification (qualitative examination) of †P. maghrebianus sp. nov. and to separate it from the similar cuspidate species, †P. mortoni. Morphospace reconstructions confirm a marked lower dental heterodonty (mesio-distal patterns) for both species. The analysis protocol employed here also allows assigning indeterminate teeth as belonging to †P. mortoni. The reconstruction of the entire lower dental plate of †P. maghrebianus sp. nov. shows a cuspidate dentition probably able to reduce tooth damages when crushing thin-shelled prey. Both dental morphologies and tooth wear patterns suggest a peculiar food processing and a diet mainly consisting of bivalves, decapods and small fish for this durophagous predator. Trophic reconstructions of the Turonian ichthyofauna inhabiting the middle to outer ramp environment of the Asfla area emphasize that †P. maghrebianus sp. nov. and the batoid †Tingitanius most likely represented second-level consumers, whereas the sclerorhynchiforms †Asflapristis and †Ptychotrygon represented third-level predators. Top positions within the food web were occupied by larger predaceous elasmobranchs (e.g., †Squalicorax).

Evolution, diversity, and disparity of the tiger shark lineage Galeocerdo in deep time.

Sharks have a long and rich fossil record that consists predominantly of isolated teeth due to the poorly mineralized cartilaginous skeleton. Tiger sharks (Galeocerdo), which represent apex predators in modern oceans, have a known fossil record extending back into the early Eocene (ca. 56 Ma) and comprise 22 recognized extinct and one extant species to date. However, many of the fossil species remain dubious, resulting in a still unresolved evolutionary history of the tiger shark genus. Here, we present a revision of the fossil record of Galeocerdo by examining the morphological diversity and disparity of teeth in deep time. We use landmark-based geometric morphometrics to quantify tooth shapes and qualitative morphological characters for species discrimination. Employing this combined approach on fossil and extant tiger shark teeth, our results only support six species to represent valid taxa. Furthermore, the disparity analysis revealed that diversity and disparity are not implicitly correlated and that Galeocerdo retained a relatively high dental disparity since the Miocene despite its decrease from four to one species. With this study, we demonstrate that the combined approach of quantitative geometric morphometric techniques and qualitative morphological comparisons on isolated shark teeth provides a useful tool to distinguish between species with highly similar tooth morphologies.

Diversity and paleoenvironmental implications of an elasmobranch assemblage from the Oligocene-Miocene boundary of Ecuador.

The occurrence and diversity of elasmobranchs from the Oligocene-Miocene boundary from Tropical America is poorly known in comparison with the paleodiversity from younger Neogene intervals of the region. Here we describe a new elasmobranch assemblage from the rich fossil site of Montañita-Olón (Dos Bocas Formation, Santa Elena, Ecuador), where other vertebrates have already been described: for example, sea turtles and cetaceans. We report a total of 27 elasmobranch taxa, 19 of which are new fossil records for Ecuador, 10 new records for the Central Eastern Pacific and four new records for South America. Additionally, in order to reconstruct the environment where these marine remains were deposited, we performed abundance, paleobathymetric and habitat preference analyses, concluding that they were likely deposited in an outer neritic (open shelf) environment. The study of Oligocene and early Miocene marine elasmobranchs faunas in Tropical America is key to addressing the issues in the evolutionary history of this group.

Evolutionary trajectories of tooth histology patterns in modern sharks (Chondrichthyes, Elasmobranchii).

During their evolutionary history, modern sharks developed different tooth mineralization patterns that resulted in very distinct histological patterns of the tooth crown (histotypes). To date, three different tooth histotypes have been distinguished: (i) orthodont teeth, which have a central hollow pulp cavity in the crown, encapsulated by a prominent layer of dentine (orthodentine); (ii) pseudoosteodont teeth, which have their pulp cavities secondarily replaced by a dentinal core of porous dentine (osteodentine), encased by orthodentine; and (iii) osteodont teeth, which lack orthodentine and the whole tooth crown of which consists of osteodentine. The aim of the present study was to trace evolutionary trends of tooth mineralization patterns in modern sharks and to find evidence for the presence of phylogenetic or functional signals. High resolution micro-computed tomography images were generated for the teeth of members of all nine extant shark orders and the putative stem group †Synechodontiformes, represented here by three taxa, to examine the tooth histology non-destructively. Pseudoosteodonty is the predominant state among modern sharks and represents unambiguously the plesiomorphic condition. Orthodonty evolved several times independently in modern sharks, while the osteodont tooth histotype is only developed in lamniform sharks. The two shark orders Heterodontiformes and Pristiophoriformes showed highly modified tooth histologies, with Pristiophorus exhibiting a histology only known from batomorphs (i.e. rays and skates), and Heterodontus showing a histological difference between anterior and posterior teeth, indicating a link between its tooth morphology, histology and durophagous lifestyle. The tooth histotype concept has proven to be a useful tool to reflect links between histology, function and its taxonomic value for distinct taxa; however, a high degree of variation, especially in the pseudoosteodont tooth histotype, demonstrates that the current histotype concept is too simplistic to fully resolve these relationships. The vascularization pattern of the dentine might offer new future research pathways for better understanding functional and phylogenetic signals in the tooth histology of modern sharks.

The Neogene fossil record of Aetomylaeus (Elasmobranchii, Myliobatidae) from the southeastern Pacific.

The presence of eagle rays of the genus Aetomylaeus in the Neogene of the Temperate Pacific coast of South America (TPSA) still is ambiguous, although the fossil record of elasmobranch fishes (sharks, rays, and skates) from this area is quite good. Here, we present the first unmistakable fossil remains of Aetomylaeus from the Neogene of the TPSA. The material comprises 13 dental plates from one site in Peru and six localities in Chile ranging in age from Miocene to Pliocene and was compared with dental plates of extant species. Our study reveals that the number of tooth rows and the shape of lateral teeth in extant species are seemingly very variable and need to be established before fossil specimens can be confidently identified. Consequently, we do not assign the fossil specimens from the Neogene of the TPSA to any species but leave them as Aetomylaeus. Moreover, we recognized that only the shape of medial teeth provides reliable diagnostic characters in our material, whereas the shape and number of lateral teeth are highly variable, similar to the condition seen in extant species.

Micro-computed tomography imaging reveals the development of a unique tooth mineralization pattern in mackerel sharks (Chondrichthyes; Lamniformes) in deep time.

The cartilaginous fishes (Chondrichthyes) have a rich fossil record which consists mostly of isolated teeth and, therefore, phylogenetic relationships of extinct taxa are mainly resolved based on dental characters. One character, the tooth histology, has been examined since the 19th century, but its implications on the phylogeny of Chondrichthyes is still in debate. We used high resolution micro-CT images and tooth sections of 11 recent and seven extinct lamniform sharks to examine the tooth mineralization processes in this group. Our data showed similarities between lamniform sharks and other taxa (a dentinal core of osteodentine instead of a hollow pulp cavity), but also one feature that has not been known from any other elasmobranch fish: the absence of orthodentine. Our results suggest that this character resembles a synapomorphic condition for lamniform sharks, with the basking shark, Cetorhinus maximus, representing the only exception and reverted to the plesiomorphic tooth histotype. Additionally, †Palaeocarcharias stromeri, whose affiliation still is debated, shares the same tooth histology only known from lamniform sharks. This suggests that †Palaeocarcharias stromeri is member of the order Lamniformes, contradicting recent interpretations and thus, dating the origin of this group back at least into the Middle Jurassic.