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.

Non-reef environments impact the diversification of extant jacks, remoras and allies (Carangoidei, Percomorpha).

Various factors may impact the processes of diversification of a clade. In the marine realm, it has been shown that coral reef environments have promoted diversification in various fish groups. With the exception of requiem sharks, all the groups showing a higher level of diversity in reefs than in non-reef habitats have diets based predominantly on plankton, algae or benthic invertebrates. Here we explore the pattern of diversification of carangoid fishes, a clade that includes numerous piscivorous species (e.g. trevallies, jacks and dolphinfishes), using time-calibrated phylogenies as well as ecological and morphological data from both extant and fossil species. The study of carangoid morphospace suggests that reef environments played a role in their early radiation during the Eocene. However, contrary to the hypothesis of a reef-association-promoting effect, we show that habitat shifts to non-reef environments have increased the rates of morphological diversification (i.e. size and body shape) in extant carangoids. Piscivory did not have a major impact on the tempo of diversification of this group. Through the ecological radiation of carangoid fishes, we demonstrate that non-reef environments may sustain and promote processes of diversification of different marine fish groups, at least those including a large proportion of piscivorous species.

The evolutionary origin of the durophagous pelagic stingray ecomorph.

Studies of the origin of evolutionary novelties (novel traits, feeding modes, behaviours, ecological niches, etc.) have considered a number of taxa experimenting with new body plans, allowing them to occupy new habitats and exploit new trophic resources. In the marine realm, colonization of pelagic environments by marine fishes occurred recurrently through time. Stingrays (Myliobatiformes) are a diverse clade of batoid fishes commonly known to possess venomous tail stings. Current hypotheses suggest that stingrays experimented with a transition from a benthic to a pelagic/benthopelagic habitat coupled with a transition from a non-durophagous diet to extreme durophagy. However, there is no study detailing macroevolutionary patterns to understand how and when habitat shift and feeding specialization arose along their evolutionary history. A new exquisitely preserved fossil stingray from the Eocene Konservat-Lagerstätte of Bolca (Italy) exhibits a unique mosaic of plesiomorphic features of the rajobenthic ecomorph, and derived traits of aquilopelagic taxa, that helps to clarify the evolutionary origin of durophagy and pelagic lifestyle in stingrays. A scenario of early evolution of the aquilopelagic ecomorph is proposed based on new data, and the possible adaptive meaning of the observed evolutionary changes is discussed. The body plan of †Dasyomyliobatis thomyorkei gen. et sp. nov. is intermediate between the rajobenthic and more derived aquilopelagic stingrays, supporting its stem phylogenetic position and the hypothesis that the aquilopelagic body plan arose in association with the evolution of durophagy and pelagic lifestyle from a benthic, soft-prey feeder ancestor.

The Phylogeny of Rays and Skates (Chondrichthyes: Elasmobranchii) Based on Morphological Characters Revisited.

Elasmobranchii are relatively well-studied. However, numerous phylogenetic uncertainties about their relationships remain. Here, we revisit the phylogenetic evidence based on a detailed morphological re-evaluation of all the major extant batomorph clades (skates and rays), including several holomorphic fossil taxa from the Palaeozoic, Mesozoic and Cenozoic, and an extensive outgroup sampling, which includes sharks, chimaeras and several other fossil chondrichthyans. The parsimony and maximum-likelihood analyses found more resolved but contrasting topologies, with the Bayesian inference tree neither supporting nor disfavouring any of them. Overall, the analyses result in similar clade compositions and topologies, with the Jurassic batomorphs forming the sister clade to all the other batomorphs, whilst all the Cretaceous batomorphs are nested within the remaining main clades. The disparate arrangements recovered under the different criteria suggest that a detailed study of Jurassic taxa is of utmost importance to present a more consistent topology in the deeper nodes, as issues continue to be present when analysing those clades previously recognized only by molecular analyses (e.g., Rhinopristiformes and Torpediniformes). The consistent placement of fossil taxa within specific groups by the different phylogenetic criteria is promising and indicates that the inclusion of more fossil taxa in the present matrix will likely not cause loss of resolution, therefore suggesting that a strong phylogenetic signal can be recovered from fossil taxa.

Diversity, palaeoecology and palaeoenvironmental significance of the Eocene chondrichthyan assemblages of the Bolca Lagerstätte, Italy.

Over the last few years, the morphology, taxonomy and systematics of the cartilaginous fish taxa of the two main sites of the Bolca Lagerstätte, Italy, (Pesciara and Monte Postale sites) have been extensively discussed in a series of papers, resulting in a complete revision of this neglected component of the Eocene Tethyan ichthyofauna. Here, we provide a comprehensive overview of the diversity, palaeoecology and palaeoenvironmental significance of the two chondrichthyan assemblages of the Pesciara and Monte Postale sites. The assemblages include 14 shark species (Lamniformes and Carcharhiniformes) and batoids (Torpediniformes, Rhinopristiformes, Myliobatiformes, Platyrhinidae and Zanobatidae), as well as a single putative chimaeriform. The Pesciara and Monte Postale sites are characterized by eight chondrichthyan taxa each, but the taxonomic compositions are distinctly different reflecting the dissimilarities in the overall composition of both fish assemblages. Palaeoecological interpretations and habitat preferences of the two chondrichthyan assemblages are consistent with previously hypothesized palaeoenvironmental settings based on sedimentological, palaeontological and geochemical evidence. The chondrichthyan assemblages of the two sites appear to be constituted by ecologically vicariant taxa, with both characterized by a predominance of benthic species with durophagous/cancritrophic feeding modes. Taxonomic composition, habitat preferences and palaeobathymetric analyses support the hypothesis that both assemblages occupied tropical marine shallow waters (likely up to 50 m deep) of the inner portion of the Lessini Shelf. The taxonomic composition of both sites is considerably different from that of any other contemporaneous Tethyan and Boreal chondrichthyan assemblages.

Rise and fall of †Pycnodontiformes: Diversity, competition and extinction of a successful fish clade.

†Pycnodontiformes was a successful lineage of primarily marine fishes that broadly diversified during the Mesozoic. They possessed a wide variety of body shapes and were adapted to a broad range of food sources. Two other neopterygian clades possessing similar ecological adaptations in both body morphology (†Dapediiformes) and dentition (Ginglymodi) also occurred in Mesozoic seas. Although these groups occupied the same marine ecosystems, the role that competitive exclusion and niche partitioning played in their ability to survive alongside each other remains unknown. Using geometric morphometrics on both the lower jaw (as constraint for feeding adaptation) and body shape (as constraint for habitat adaptation), we show that while dapediiforms and ginglymodians occupy similar lower jaw morphospace, pycnodontiforms are completely separate. Separation also occurs between the clades in body shape so that competition reduction between pycnodontiforms and the other two clades would have resulted in niche partitioning. Competition within pycnodontiforms seemingly was reduced further by evolving different feeding strategies as shown by disparate jaw shapes that also indicate high levels of plasticity. Acanthomorpha was a teleostean clade that evolved later in the Mesozoic and which has been regarded as implicated in driving the pycnodontiforms to extinction. Although they share similar body shapes, no coeval acanthomorphs had similar jaw shapes or dentitions for dealing with hard prey like pycnodontiforms do and so their success being a factor in pycnodontiform extinction is unlikely. Sea surface temperature and eustatic variations also had no impact on pycnodontiform diversity patterns according to our results. Conversely, the occurrence and number of available reefs and hardgrounds as habitats through time seems to be the main factor in pycnodontiform success. Decline in such habitats during the Late Cretaceous and Palaeogene might have had deleterious consequences for pycnodontiform diversity. Acanthomorphs occupied the niches of pycnodontiforms during the terminal phase of their existence.

Revision of the Eocene 'Platyrhina' species from the Bolca Lagerstätte (Italy) reveals the first panray (Batomorphii: Zanobatidae) in the fossil record.

The fossil-Lagerstätte of Bolca (Italy) is well known for the diversity and exquisite preservation of its bony and cartilaginous fishes documenting tropical shallow-water marine environments associated with coral reefs in the western Tethys during the early Eocene. In this study, the taxonomic, systematic and phylogenetic position of two batoid species traditionally assigned to the living thornback ray genus Platyrhina is re-evaluated. †Platyrhina bolcensis Heckel, 1851 is recognized as a separate species of the Platyrhinidae because of its plate-like antorbital cartilage with an irregular outline and a small horn on the nasal capsules. Also, the rostral cartilage does not reach the anterior border of the disc. Support for the placement of this species within the new genus †Eoplatyrhina gen. nov. is based on a combination of morphological and meristic features (e.g. nasal capsules at right angles to the rostrum; large space between the hyomandibulae and mandibular arch; approximately 132 vertebral centra; 15-16 rib pairs; 81-87 pectoral radials; 18-21 pelvic radials; short, straight and stout claspers; 40-50 caudal-fin radials; thorns absent). A second species, †Platyrhina egertoni (De Zigno, 1876), is more closely related to the living panray Zanobatus than Platyrhina and is assigned here to †Plesiozanobatus gen. nov. because of a combination of characters that support its placement within the family Zanobatidae (tail stout and short, distinctly demarcated from disc; two dorsal fins and complete caudal fin; small dermal denticles and scattered thorns covering disc and tail; rostral cartilage absent; nasal capsules without horn-like processes; mesopterygium absent). The systematic position of a third taxon, †Platyrhina gigantea (Blainville, 1818), is currently impossible to establish due to the poor preservation of the only known specimen, and therefore we propose to consider it a nomen dubium. Palaeoecological and biogeographic features of the Eocene platyrhinids and zanobatids from Bolca are also discussed. http://zoobank.org/urn:lsid:zoobank.org:pub:B4C7A979-7972-409B-B489-A6DDD5E35FE5.

Evolutionary trends of the conserved neurocranium shape in angel sharks (Squatiniformes, Elasmobranchii).

Elasmobranchii (i.e., sharks, skates, and rays) forms one of the most diverse groups of marine predators. With a fossil record extending back into the Devonian, several modifications in their body plan illustrate their body shape diversity through time. The angel sharks, whose fossil record dates back to the Late Jurassic, some 160 Ma, have a dorsoventrally flattened body, similar to skates and rays. Fossil skeletons of this group show that the overall morphology was well established earlier in its history. By examining the skull shape of well-preserved fossil material compared to extant angel sharks using geometric morphometric methods, within a phylogenetic framework, we were able to determine the conservative skull shape among angel sharks with a high degree of integration. The morphospace occupation of extant angel sharks is rather restricted, with extensive overlap. Most of the differences in skull shape are related to their geographic distribution patterns. We found higher levels of disparity in extinct forms, but lower ones in extant species. Since angel sharks display a highly specialized prey capture behaviour, we suggest that the morphological integration and biogeographic processes are the main drivers of their diversity, which might limit their capacity to display higher disparities since their origin.

Sharks, rays and skates (Chondrichthyes, Elasmobranchii) from the Upper Marine Molasse (middle Burdigalian, early Miocene) of the Simssee area (Bavaria, Germany), with comments on palaeogeographic and ecological patterns.

Elasmobranch remains are quite common in Miocene deposits and were the subject of numerous studies since the middle of the nineteenth century. Nevertheless, the taxonomic diversity of the Marine Molasse sharks, rays and skates is still largely unknown. Here, we describe 37 taxa from the lower Miocene of the Molasse Basin: 21 taxa could be identified at species level, whereas 15 taxa could only be assigned to genus and one taxon is left as order incertae sedis. The material was collected from deposits of the Auwiesholz Member of the Achen Formation (middle Burdigalian, middle Ottnangian age, ca. 17.8 Ma) exposed near Simssee, Upper Bavaria. This faunal assemblage is a mixture of shallow marine, near-coastal, pelagic and deep-water taxa. The fauna from Simssee displays different biogeographic dynamics at local and regional scales, possibly related to the intense climatic, oceanographic and tectonic events that occurred during the Eggenburgian-Ottnangian stages. The faunal relationships of the early Miocene chondrichthyan faunas from the Mediterranean Sea and Paratethys with others regions are established on the basis of qualitative (presence/absence) data. The beta diversity (Sørensen-Dice coefficient) of the Miocene Molasse elasmobranchs was used to characterize the taxonomic differentiation between localities and regions. According to our results, the fauna from Simssee shows close similarities with those from Switzerland, Austria, France and northern Germany. Faunal similarities and differences are mainly related to tectonic events and oceanographic variables (i.e. migration through seaway passages) or might represent collecting biases.

A new Miocene skate from the Central Paratethys (Upper Austria): the first unambiguous skeletal record for the Rajiformes (Chondrichthyes: Batomorphii).

A new fossil skate, Ostarriraja parva gen. et sp. nov., represented by a single partial articulated skeleton collected from the early Miocene fish-bearing strata of Upper Austria, is described here in detail. This taxon exhibits a unique combination of skeletal and dental features (e.g. nasal capsules broad and oval; presence of pectoral arch; compound radial articulated with single radial segments in serial fashion; separated pelvic girdle condyles; reduced catenated calcification of radials; about 86 pectoral radials; 20-21 pelvic-fin radials; 65-70 predorsal vertebrae) that clearly support its assignment to a new genus of the order Rajiformes, and the phylogenetic analyses reveal its basal position within the group. The comparison between Ostarriraja and the holomorphic batoids from Late Cretaceous of Lebanon traditionally aligned with skates concurs to suggest that this Neogene occurrence represents unquestionably the first known skeletal record for the group. The morphological and phylogenetic affinities of Ostarriraja with the living skates suggest a close association of this taxon with the temperate-cold water environments hypothesized for the Central Paratethys during the early Miocene. http://zoobank.org/urn:lsid:zoobank.org:pub:8BB8F0F3-35C5-47FA-AE3C-2CBF445C4BCA.

An Eocene paraclupeid fish (Teleostei, Ellimmichthyiformes) from Bolca, Italy: the youngest marine record of double-armoured herrings.

A new double-armoured herring of the clupeomorph order Ellimmichthyiformes, Eoellimmichthys superstes gen. et sp. nov., is described herein based on a single partially complete specimen from the early Eocene Pesciara site of the Bolca Konservat-Lagerstätte, north-eastern Italy. The fossil documented herein exhibits a unique combination of features (e.g. ornamentation of the skull bones; medial fusion of the contralateral halves of the neural arches of abdominal vertebrae; teeth on endopterygoid, parhypural fused to the first preural centrum; presence of a short series of 6 (or 7) predorsal scutes increasing in size posteriorly; postpelvic scutes bearing very prominent spines), which clearly supports its recognition as a new genus and species of the family Paraclupeidae. Evidence is provided to show that Eoellimmichthys gen. nov. is most closely related phylogenetically to the Early Cretaceous genus Ellimmichthys. The new taxon described herein represents the youngest marine occurrence of the family Paraclupeidae and, more generally, of the order Ellimmichthyiformes, suggesting that the shallow marine settings of the Tethys might have favoured the persistence of certain fish lineages that were severely affected by the end-Cretaceous mass extinction.

Mosaic of plesiomorphic and derived characters in an Eocene myliobatiform batomorph (Chondrichthyes, Elasmobranchii) from Italy defines a new, basal body plan in pelagic stingrays.

BACKGROUND: End-Cretaceous niche-filling by benthic Mesozoic survivors resulted in a prominent increase of durophagous fish families, resulting in the appearance of the earliest representatives of several extant fish lineages, including the pelagic durophagous stingrays, a monophyletic clade of myliobatiform batoids that is characterized by a derived swimming mode and feeding habits. Although the earliest members appeared in the Late Cretaceous, most of the crown genera date back to the Eocene. RESULTS: In this study, we re-examine the anatomy of the Eocene eagle ray Promyliobatis gazolai (de Zigno), represented by two nearly complete and articulated specimens from the world-famous Ypresian Konservat-Lagerstätte of Bolca, in detail. This taxon exhibits a mosaic of plesiomorphic and derived characters (e.g. tail sting displaced posteriorly on the tail, at about 50-60% of tail length; pectoral fins joining in front of the head; anterior and posterior pectoral fin margins nearly straight; compagibus laminam absent; single, unfragmented mesopterygium) that clearly define a new body plan within the pelagic durophagous stingrays. CONCLUSIONS: The significant morphological differences between Promyliobatis and extant representatives of Myliobatidae, Aetobatidae, Rhinopteridae, and Mobulidae, support its placement as separate stem group member. The phylogenetic placement of Promyliobatis, based on skeletal and dental characters, strongly supports its basal position within pelagic stingrays. However, its position within the Myliobatiformes becomes unstable when stingray taxa known by fossil teeth only are included. A comparative analysis of the skeletal and tooth morphologies, as well as of the evolutionary trends of pelagic stingrays is also discussed.

Eocene sand tiger sharks (Lamniformes, Odontaspididae) from the Bolca Konservat-Lagerstätte, Italy: palaeobiology, palaeobiogeography and evolutionary significance.

Here we report the first record of one of the most common and widespread Palaeogene selachians, the sand tiger shark Brachycarcharias, from the Ypresian Bolca Konservat-Lagerstätte. The combination of dental character of the 15 isolated teeth collected from the Pesciara and Monte Postale sites (e.g. anterior teeth up to 25 mm with fairly low triangular cusp decreasing regularly in width; one to two pairs of well-developed lateral cusplets; root with broadly separated lobes; upper teeth with a cusp bent distally) supports their assignment to the odontaspidid Brachycarcharias lerichei (Casier, 1946), a species widely spread across the North Hemisphere during the early Palaeogene. The unambiguous first report of this lamniform shark in the Eocene Bolca Konservat-Lagerstätte improves our knowledge concerning the diversity and palaeobiology of the cartilaginous fishes of this palaeontological site, and provides new insights about the biotic turnovers that involved the high trophic levels of the marine settings after the end-Cretaceous extinction.

Reappraisal of the Eocene whiptail stingrays (Myliobatiformes, Dasyatidae) of the Bolca Lagerstätte, Italy.

The Eocene whiptail stingrays of the family Dasyatidae from the Bolca Lagerstätte, NE Italy, are revised herein in detail. The analysis of the anatomical and morphometric features allows us to identify the species "Dasyatis" zigni (Molin, 1861) as a junior synonym of "D." muricatus (Volta, 1796), and to assign it to the new genus Tethytrygon gen. n. This new taxon exhibits a unique combination of features (e.g., rhombic disc wider than long, elongated tail folds fail to reach the tip of the tail, thorns absent, single serrated tail sting, "caniniform" teeth on upper jaw, tooth crown ornamentation absent, 175-179 vertebrae, 108-117 pectoral radials, 24-27 pelvic radials and other features of clasper anatomy) that clearly support its attribution to the subfamily Neotrygoninae of the stingray family Dasyatidae. The morphological and phylogenetic affinities of Tethytrygon gen. n. with the living neotrygonines (Neotrygon and Taeniura) suggest a close association of this taxon with the tropical shallow-water habitats hypothesized for the Bolca palaeoenvironment during the early Eocene. Moreover, the analysis of the fossil occurrences of the neotrygonines provides new insights into the role of the Tethys for the origin and evolutionary history of certain whiptail stingrays.

A bizarre Eocene dasyatoid batomorph (Elasmobranchii, Myliobatiformes) from the Bolca Lagerstätte (Italy) reveals a new, extinct body plan for stingrays.

In the last few years, the detailed revision of the Eocene cartilaginous fishes (Chondrichthyes) from the Bolca Lagerstätte (Italy) has provided new insights into the fish biodiversity of the western Tethys. The morphological analysis of three previously undescribed specimens from the Pesciara deposit of Bolca revealed the existence of a new stingray taxon, †Lessiniabatis aenigmatica gen. et sp. nov., which is unique among the myliobatiform batoids in having the following unique combination of characters: low number of vertebrae posterior to the pelvic girdle (65-68); thoracolumbar synarcual extending backward beyond the pelvic girdle; tail extremely short not protruding from the posterior edge of the pectoral disc; radials proximally fused to each other; pelvic girdle extremely small and strongly arched; dorsal and caudal fins absent; tail stings and cartilaginous tail rod absent; and teeth of dasyatoid morphology with smooth enameloid surface. The phylogenetic analysis suggests that †Lessiniabatis gen. nov. is deeply nested within the benthic stingrays (Dasyatoidea) representing the sister to all dasyatids and potamotrygonids. Its unique anatomy clearly reveals the existence of a new hitherto unknown body plan experimented by benthic stingrays, whose evolution can be possibly linked to the adaptive fish radiation in the aftermath of the end-Cretaceous extinction.

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.

Eoalosa janvieri gen. et sp. nov., a new clupeid fish (Teleostei, Clupeiformes) from the Eocene of Monte Bolca, Italy.

Fishes of the family Clupeidae are extremely abundant in the Eocene fossiliferous limestone of Monte Bolca representing the most common group from this celebrated locality. A new clupeid from the Pesciara site, Eoalosa janvieri gen. et sp. nov., is described. The new taxon exhibits a unique combination of characters supporting its recognition as a new genus and species of clupeid fish that is tentatively placed in the subfamily Alosinae. The description of this new taxon improves our knowledge of the diversity of clupeoid fishes in the Eocene of Monte Bolca.

Revision of Eocene electric rays (Torpediniformes, Batomorphii) from the Bolca Konservat-Lagerstätte, Italy, reveals the first fossil embryo in situ in marine batoids and provides new insights into the origin of trophic novelties in coral reef fishes.

The Eocene electric ray †Titanonarke Carvalho, 2010 from the Bolca Konservat-Lagerstätte, north-eastern Italy, is redescribed in detail based upon new material from recent excavations. This taxon exhibits a combination of features (large voids between the pectoral and the axial skeleton filled in life by electric organs, anteriorly directed fan-shaped antorbital cartilages, lack of dermal denticles, long prepelvic processes, and rounded basibranchial copula with a small caudal tab) that clearly supports its assignment to the order Torpediniformes. The analysis of new material also demonstrates that the previous apparent absence of typical narcinoid characters used to diagnose †Titanonarke was the result of taphonomic biases. †Titanonarke shares at least three synapomorphies (presence of a rostral fontanelle, low number of ribs, and rostral cartilage connected to the antorbital cartilage through lateral appendices) with the extant genera Benthobatis, Diplobatis, Discopyge and Narcine, with which it forms a clade (family Narcinidae) recognized herein as unquestionably monophyletic. Moreover, based upon a single specimen of †Titanonarke that exhibits a unique combination of morphometric and meristic features, a new species of Eocene numbfish †T. megapterygia sp. nov., is recognized. The presence of several specimens representing different ontogenetic stages of at least two species of numbfishes suggests a close association of this taxon with shallow-water habitats corresponding to coral reefs as hypothesized for the Monte Postale palaeoenvironment. The occurrence of a fossilized marine batoid embryo is reported here for the first time. Moreover, the analysis of the gut contents suggests that the dietary adaptations of †Titanonarke can be related, at least in part, to an opportunistic strategy in the context of abundant larger foraminifera in the Monte Postale palaeobiotope, suggesting that this kind of feeding mode, known to occur in present-day reefs, already was realized 50 million years ago.

A quantitative approach to determine the taxonomic identity and ontogeny of the pycnodontiform fish Pycnodus (Neopterygii, Actinopterygii) from the Eocene of Bolca Lagerstätte, Italy.

BACKGROUND: The pycnodontiform fish Pycnodus is one of the representatives of the highly diverse actinopterygian fish fauna from the early Eocene Bolca Lagerstätte, representing one of the youngest and thus last occurrences of this extinct neopterygian clade. This genus has historically been used as a wastebasket taxon in regards to poorly known pycnodontiform fossils. Authors have argued over the specific status of the Bolca Lagerstätte Pycnodus in terms of how many species are contained within the genus with some arguing for multiple species and others suggesting lumping all Bolca specimens together into one species. METHODS: Here, we use a quantitative approach performing biometric and geometric morphometric analyses on 52 specimens of Pycnodus in order to determine if the morphological variability within the sample might be related to inter- or intraspecific variation. RESULTS: The analyses revealed that the variations of body shape, morphometric and meristic characters cannot be used to distinguish different morphotypes. On the contrary, our results show a remarkable link between shape and size, related to ontogeny. DISCUSSION: Differences in body shape of small (juvenile) and large (adult) individuals is probably related to different microhabitats occupation on the Bolca reef with juveniles sheltering within crevices on the reef and adults being more powerful swimmers that swim above the coral. Taxonomically, we suggest that the Bolca Pycnodus should be referred to strictly as Pycnodus apodus as this was the name given to the holotype. Additionally, an overview of species assigned to Pycnodus is given.