0000000000496527
AUTHOR
Héctor Botella
Categorical versus geometric morphometric approaches to characterizing the evolution of morphological disparity in Osteostraci (Vertebrata, stem Gnathostomata)
Morphological variation (disparity) tends to be evaluated through two non-mutually exclusive approaches: (i) quantitatively, through geometric morphometrics, and (ii) in terms of discrete, ‘cladistic’, or categorical characters. Uncertainty over the comparability of these approaches diminishes the potential to obtain nomothetic insights into the evolution of morphological disparity, and the few benchmarking studies conducted so far show contrasting results. Here, we apply both approaches to characterising morphology in the stem-gnathostome vertebrate clade Osteostraci, in order to assess congruence between these alternative methods as well as to explore the evolutionary patterns of the grou…
The palaeontological virtual collection of the University of Valencia’s Natural History Museum: a new tool for palaeontological heritage outreach
The rise of new digitalization technologies is changing the way to access to the fossil collections for palaeontology outreach, providing new tools to preserve our important palaeontological heritage. In this sense, museums and palaeontological institutions,aware of the advantages of applying these new technologies for the accomplishment of their functions, have started to develop their own online repositories to facilitate the access to the most representative fossil of their collections. Following this trend, the aim of this work is to present the creation of the new “Virtual 3D Collection” of the University of Valencia’s Natural History Museum, showing, as an example, the “Ichnofossil Co…
Late Devonian (Famennian) chondrichthyes from Mexico
The Paleozoic vertebrate fossil record from Mexico is very scarce and strongly biased by rock exposure, composed mainly of upper Paleozoic (Carboniferous and Permian) outcrops (e.g., Sánchez-Zavala et al., 1999; Poole et al., 2005; González- Rodríguez et al., 2013). In particular, the Mexican Paleozoic fish fossil record comprises a few isolated chondrichthyan scales and some semiarticulated symphysial tooth whorls belonging to the iconic shark Helicoprion, with ages ranging from the Late Carboniferous to the early Permian. The first reported Paleozoic fish from Mexico belongs to a semiarticulated symphysial tooth whorl of Helicoprion mexicanus from the Permian of Coahuila, near Las Delicia…
First occurrence of fossil vertebrates from the Carboniferous of Colombia
Data concerning Paleozoic vertebrates from the South American continent are still scarce. In Colombia, occurrences were until now restricted to the Late Devonian fish assemblage from Floresta and, ...
First Perigondwanan record of actinolepids (Vertebrata: Placodermi: Arthrodira) from the Lochkovian (Early Devonian) of Spain and its palaeobiogeographic significance.
Different palaeogeographic models have been proposed for the position of Laurussia ( including Baltica) and Gondwana-derived microcontinents ( including Ibero-Armorica) during Ordovician to Late Carboniferous times. Principal differences concern the presence and duration of a large ocean, the Rheic Ocean, acting as a faunal barrier between these areas. The timing of the collision of Laurussia with Gondwana and/or Gondwana-derived terranes continues to be debated. Here we present new faunal data revealing close biogeographical relations between Ibero-Armorica ("Perigondwanan" or Gondwanan derivate terranes) and Podolia (SE margin of Baltica, in Laurussia). The placoderm assemblage found in t…
A NEW EARLY DEVONIAN THELODONT FROM CELTIBERIA (SPAIN), WITH A REVISION OF SPANISH THELODONTS
: A revision of thelodont scales from the Lower Devonian of the Iberian Chains enables their grouping into two taxa: Turinia pagei and T. nachoi sp. nov. These taxa are clearly distinguishable by morphological and histological features; they also have a different stratigraphic range (T. pagei is restricted to Lochkovian strata, whereas T. nachoi sp. nov. occurs within lower–middle Pragian rocks). The new species is represented by head, transitional (cephalopectoral) and trunk scales.
Jaws and teeth of the earliest bony fishes
Extant jawed vertebrates, or gnathostomes, fall into two major monophyletic groups, namely chondrichthyans (cartilaginous fishes) and osteichthyans (bony fishes and tetrapods). Fossil representatives of the osteichthyan crown group are known from the latest Silurian period, 418 million years (Myr) ago, to the present. By contrast, stem chondrichthyans and stem osteichthyans are still largely unknown. Two extinct Palaeozoic groups, the acanthodians and placoderms, may fall into these stem groups or the common stem group of gnathostomes, but their relationships and monophyletic status are both debated. Here we report unambiguous evidence for osteichthyan characters in jaw bones referred to th…
The chondrichthyan fauna from the Middle Triassic (Ladinian) of the Iberian Range (Spain)
ABSTRACT Here we present for the first time a detailed taxonomic study of a diverse chondrichthyan fauna from the Middle Triassic of the Iberian Range (Spain). The assemblage consists of isolated remains of seven species of five non-neoselachian shark genera (Palaeobates, Hybodus, Pseudodalatias, Prolatodon, gen. nov., and Lissodus), including a new species of hybodontiform shark, Hybodus bugarensis, sp. nov. In addition, a new homalodontid genus, Prolatodon, sp. nov., is erected for the taxa ‘Polyacrodus’ bucheri and ‘Polyacrodus’ contrarius. The chondrichthyans of the Iberian Range represent a heterogeneous group from a paleogeographic point of view made up of common components of Middle …
Lonchidion derenzii, sp. nov., a new lonchidiid shark (Chondrichthyes, Hybodontiforms) from the Upper Triassic of Spain, with remarks on lonchidiid enameloid
Lonchidiidae Herman, 1977, represents one of the most diverse and controversial families of Hybodontiformes, the sister group of Neoselachii (i.e., modern sharks, skates, and rays). It was initially erected as a monogeneric family including only Lonchidion Estes, 1964, a genus of small euryhaline hybodonts from the Mesozoic. Recently, Cappetta (2012) recognized up to eight genera within the family: Baharyodon, Diplolonchidion, Vectiselachos, Hylaeobatis, Isanodus, Parvodus,Lissodus, andLonchidion, although the content of the family is still under discussion (see, e.g., Rees, 2008; Khamha et al., 2016). Major discrepancies concern the phylogenetic relationships between Lonchidion and Lissodu…
On the tail ofErrivaspisand the condition of the caudal fin in heterostracans
Articulated caudal fins of heterostracans are preserved in very rare cases. Their morphology and internal structure are completely unknown for the majority of species. One of the few preserved caudal fins belongs to the Early Devonian pteraspidid Errivaspis waynensis from the Welsh Borderland, UK. There are two different reconstructions of the tail: (1) strongly asymmetric, with a considerably longer ventral lobe, and a rather concave posterior margin (e.g. White 1935); and (2) almost symmetrical with an only slightly longer ventral lobe and a small notch in the posterior margin (e.g. Blieck 1984). A new reconstruction of the Errivaspis caudal fin, based on a detailed study of White's speci…
Biomechanics of Machaeracanthus pectoral fin spines provide evidence for distinctive spine function and lifestyle among early chondrichthyans
Acanthodians are a major group of Paleaozoic jawed vertebrates that constitute a paraphyletic assemblage of stem-chondrichthyans (Brazeau and Friedman, 2015). Representatives of this group are characterized, among other traits, by the presence of bony spines in front of all paired and median fins except the caudal (Denison, 1979), which has given rise to their colloquial name of 'spiny sharks'. The occurrence of pectoral fin spines is recognized as a potential gnathostome synapomorphy (Miller et al., 2003) or symplesiomorphy (Coates, 2003), being also present in other major groups of Paleaozoic jawed vertebrates, including placoderms (Young, 2010), 'non-acanthodian' chondrichthyans (Miller …
Pseudodalatias henarejensisnov. sp. a new pseudodalatiid (Elasmobranchii) from the Middle Triassic of Spain
ABSTRACT Pseudodalatiids, a chondrichthyan family of uncertain phylogenetic affinities, have been hitherto exclusively known from the tooth-based species Pseudodalatias barnstonensis (Sykes, 1971), which has a stratigraphic range restricted to the Upper Triassic of Europe. Pseudodalatias presents a characteristic dentition which allows it to hold and cut its prey, showing a neoselachian design, but lacking the triple-layered enameloid microstructure of neoselachian teeth. The discovery of Pseudodalatias henarejensis nov. sp. in the Ladinian of Spain extends the stratigraphical range and the palaeogeographical distribution of this family. This new species also demonstrates that a cutting-clu…
Network dynamics of eukaryotic LTR retroelements beyond phylogenetic trees
Abstract Background Sequencing projects have allowed diverse retroviruses and LTR retrotransposons from different eukaryotic organisms to be characterized. It is known that retroviruses and other retro-transcribing viruses evolve from LTR retrotransposons and that this whole system clusters into five families: Ty3/Gypsy, Retroviridae, Ty1/Copia, Bel/Pao and Caulimoviridae. Phylogenetic analyses usually show that these split into multiple distinct lineages but what is yet to be understood is how deep evolution occurred in this system. Results We combined phylogenetic and graph analyses to investigate the history of LTR retroelements both as a tree and as a network. We used 268 non-redundant …
Supplementary methods, figures, tables and information from Use of nursery areas by the extinct megatooth shark Otodus megalodon (Chondrichthyes: Lamniformes)
Nursery areas are fundamental for the success of many marine species, particularly for large, slow-growing taxa with low fecundity and high age of maturity. Here, we examine the population size-class structure of the extinct gigantic shark Otodus megalodon in a newly described middle Miocene locality from Northeastern Spain, as well as in eight previously known formations (Temblor, Calvert, Pisco, Gatun, Chucunaque, Bahía Inglesa, Yorktown and Bone Valley). In all cases, body lengths of all individuals were inferred from dental parameters and the size-class structure was estimated from Kernel probability density functions and Gaussian mixture models. Our analyses support the presence of fiv…
The oldest fossil evidence of a dental lamina in sharks
(2006). The oldest fossil evidence of a dental lamina in sharks. Journal of Vertebrate Paleontology: Vol. 26, No. 4, pp. 1002-1003.
Discovery of a Lochkovian flora (Lower Devonian) in the Iberian Peninsula
ABSTRACT The Lower Devonian represents an important episode in plant life history, which was marked by the diversification of land plants. Unfortunately, remains of early Devonian plants in the Iberian Peninsula are scarce. In the present paper, we describe a small assemblage of early land plants from the Lochkovian (Lower Devonian) of the Teruel Province in Spain. The main element consists of Taeniocrada-like sterile stems that are predominandy dichotomous and ribbon-like with a narrow central strand. An unidentified fossil formed by dichotomous axes together with an uncertain globular structure were also observed. This finding increases our knowledge of the palaeogeographical distribution…
Squamation and ecology of thelodonts
Thelodonts are an enigmatic group of Paleozoic jawless vertebrates that have been well studied from taxonomical, biostratigraphic and paleogeographic points of view, although our knowledge of their ecology and mode of life is still scant. Their bodies were covered by micrometric scales whose morphology, histology and the developmental process are extremely similar to those of extant sharks. Based on these similarities and on the well-recognized relationship between squamation and ecology in sharks, here we explore the ecological diversity and lifestyles of thelodonts. For this we use classic morphometrics and discriminant analysis to characterize the squamation patterns of a significant num…
Supplementary methods, figures, tables and information from Use of nursery areas by the extinct megatooth shark Otodus megalodon (Chondrichthyes: Lamniformes)
Nursery areas are fundamental for the success of many marine species, particularly for large, slow-growing taxa with low fecundity and high age of maturity. Here, we examine the population size-class structure of the extinct gigantic shark Otodus megalodon in a newly described middle Miocene locality from Northeastern Spain, as well as in eight previously known formations (Temblor, Calvert, Pisco, Gatun, Chucunaque, Bahía Inglesa, Yorktown and Bone Valley). In all cases, body lengths of all individuals were inferred from dental parameters and the size-class structure was estimated from Kernel probability density functions and Gaussian mixture models. Our analyses support the presence of fiv…
Supplementary material from Functional assessment of morphological homoplasy in stem-gnathostomes
The Osteostraci and Galeaspida are stem-gnathostomes, occupying a key phylogenetic position for resolving the nature of the jawless ancestor from which jawed vertebrates evolved more than 400 million years ago. Both groups are characterized by the presence of rigid headshields that share a number of common morphological traits, in some cases hindering the resolution of their interrelationships and the exact nature of their affinities with jawed vertebrates. Here, we explore the morphological and functional diversity of osteostracan and galeaspid headshields using an innovative approach that combines geometric morphometrics and computational fluid dynamics, thereby constraining the underlyin…
Functional assessment of morphological homoplasy in stem-gnathostomes
Osteostraci and Galeaspida are stem-gnathostomes, occupying a key phylogenetic position for resolving the nature of the jawless ancestor from which jawed vertebrates evolved more than 400 million years ago. Both groups are characterized by the presence of rigid headshields that share a number of common morphological traits, in some cases hindering the resolution of their interrelationships and the exact nature of their affinities with jawed vertebrates. Here, we explore the morphological and functional diversity of osteostracan and galeaspid headshields using an innovative approach that combines geometric morphometrics and computational fluid dynamics, thereby constraining the underlying fa…
Enameloid microstructure in the oldest known chondrichthyan teeth
Botella, H., Donoghue, P.C.J and Martinez-Perez, C. 2009. The enameloid microstructure of the oldest known chondrichthyan teeth. — Acta Zoologica (Stockholm) 90 (Suppl. 1): 103‐108 Previous studies on tooth enameloid microstructure in several chondrichthyan taxa spanning the phylogeny of the group provided support for the homology of chondrichthyan tooth enameloid. This hypothesis requires that a single crystallite enameloid (SCE) monolayer must be present in the teeth of the most primitive chondrichthyan. However, the dental microstructure of the earliest sharks has yet to be investigated. We have studied the tooth enameloid microstructure of the two oldest tooth-bearing shark species curr…
First fossil record (Middle Miocene) of the viper shark Trigonognathus Mochizuki and Ohe, 1990, in the Mediterranean realm
The genus Trigonognathus Mochizuki and Ohe, 1990, is a monospecific taxon of `lantern sharks¿ (i.e., family Etmopteridae), a group of small-sized bioluminescent deep-sea chondrichthyans, ranging in mature male specimens between 42¿47 cm total length, and at least 52 cm for females (Ebert et al., 2021). This shark inhabits the upper continental slopes as well as the uppermost slope of seamounts, often at the bottom, at depths ranging between 250¿1000 m, but has been caught at 150 m and 270 m in deep open waters (Mochizuki and Ohe, 1990; Compagno et al., 2005; Ebert et al., 2021). Only two species have been described thus far, the extant species Trigonognathus kabeyai Mochizuki and Ohe, 1990,…
Use of nursery areas by the extinct megatooth shark Otodus megalodon (Chondrichthyes: Lamniformes)
Nursery areas are fundamental for the success of many marine species, particularly for large, slow-growing taxa with low fecundity and high age of maturity. Here, we examine the population size-class structure of the extinct gigantic shark Otodus megalodon in a newly described middle Miocene locality from Northeastern Spain, as well as in eight previously known formations (Temblor, Calvert, Pisco, Gatún, Chucunaque, Bahía Inglesa, Yorktown and Bone Valley). In all cases, body lengths of all individuals were inferred from dental parameters and the size-class structure was estimated from kernel probability density functions and Gaussian mixture models. Our analyses support the presence of fi…
Ecomorphological inferences in early vertebrates: reconstructing Dunkleosteus terrelli (Arthrodira, Placodermi) caudal fin from palaeoecological data
Our knowledge about the body morphology of many extinct early vertebrates is very limited, especially in regard to their post-thoracic region. The prompt disarticulation of the dermo-skeletal elements due to taphonomic processes and the lack of a well-ossified endoskeleton in a large number of groups hinder the preservation of complete specimens. Previous reconstructions of most early vertebrates known from partial remains have been wholly based on phylogenetically closely related taxa. However, body design of fishes is determined, to a large extent, by their swimming mode and feeding niche, making it possible to recognise different morphological traits that have evolved several times in no…
New insights into Late Devonian vertebrates and associated fauna from the Cuche Formation (Floresta Massif, Colombia)
New vertebrate remains are reported from the Late Devonian (?Frasnian) Cuche Formation of northeastern Colombia, including a new taxon of antiarch placoderm (Colombialepis villarroeli, gen. et sp. nov., previously reported as Asterolepis) and a new taxon of arthrodiran placoderm (Colombiaspis rinconensis, gen. et sp. nov.). We also report evidence of a stegotrachelid actinopterygian, a diplacanthid acanthodian (cf. Florestacanthus morenoi), a second antiarch placoderm (Bothriolepis sp.), a putative megalichthyid, and a putative tristichopterid. The absence of typical Euramerican markers, e.g., Asterolepis and Strepsodus, in this assemblage suggests that faunal interchange between Euramerica…
Patterns of ecological diversification in thelodonts
Here we explore the spatial, temporal and phylogenetic patterns of ecological diversification for the entire clade of thelodonts, one of the earliest groups of vertebrates and longest lasting of the Palaeozoic agnathans in the fossil record. Parsimony and maximum-likelihood methods are used to reconstruct ancestral states of their geographical distributions, habitats and lifestyles. Our results support the concept that thelodonts originated during the Middle?-Late Ordovician probably in marine open waters of Laurasia, with a demersal lifestyle on hard substrates being the ancestral condition for the whole clade. Later, thelodonts underwent a complex ecological diversification and palaeobiog…
Assessing metabolic constraints on the maximum body size of actinopterygians: locomotion energetics of Leedsichthys problematicus (Actinopterygii, Pachycormiformes)
Maximum sizes attained by living actinopterygians are much smaller than those reached by chondrichthyans. Several factors, including the high metabolic requirements of bony fishes, have been proposed as possible body‐size constraints but no empirical approaches exist. Remarkably, fossil evidence has rarely been considered despite some extinct actinopterygians reaching sizes comparable to those of the largest living sharks. Here, we have assessed the locomotion energetics of Leedsichthys problematicus, an extinct gigantic suspension‐feeder and the largest actinopterygian ever known, shedding light on the metabolic limits of body size in actinopterygians and the possible underlying factors th…
Sucesión de Polygnátidos (Conodontos) del Emsiense (Devónico Inferior) en los Pirineos Centrales Españoles
A comprehensive conodont study of five sections of Emsian age, carried out in the Spanish Central Pyrenees, has revealed an important succession of polygnathids that can be used for identifying basal boundaries of globally recognized biozones. This succession consists of Polygnathus excavatus excavatus, Po. gronbergi, Po. nothoperbonus, Po. mashkovae, Po. laticostatus, and the new species Polygnathus luciae that allows the identification of the excavatus, nothoperbonus and laticostatus Zones. The nothoperbonus Zone is further subdivided into Lower and Upper nothoperbonus subzones, relying upon the lowest occurrence of Po. mashkovae. These biostratigraphic data from the Spanish Central Pyren…
New data on the Lower Devonian chondrichthyan fauna from Celtiberia (Spain)
Lower Devonian microvertebrate remains from Spain, including localities in Celtiberia and Cantabrian Mountains, have been extensively studied during recent decades (e.g., Mader, 1986; Wang, 1993; ...
Machaeracanthus goujetin. sp. (Acanthodii) from the Lower Devonian of Spain and northwest France, with special reference to spine histology
ABSTRACT We describe here a new machaeracanthid acanthodian (Machaeracanthus goujeti n. sp.), based on isolated spines, scales and scapulocoracoids from the Lower Devonian (Lochkovian-Pragian) of the Nogueras Formation, Celtiberia, Spain. The new taxon also includes a fragmentary spine and isolated scales from the Lower Devonian of northern Spain (Palencia and Cantabrian Mountains) and western France (Saint-Cenere) originally assigned to Machaeracanthus sp. The spines of M. goujeti n. sp. comprise two morphotypes in agreement with the morphofunctional model of a pair of pectoral spines articulating with the pectoral girdle already indicated for M. hunsrueckianum Sudkamp & Burrow, 2007, M. l…
Bioluminescent-like squamation in the galeomorph shark Apristurus ampliceps (Chondrichthyes: Elasmobranchii)
Galeomorph sharks constitute the most taxonomically and ecologically diverse superorder of living selachians. Despite comprising several typically deep-water taxa, no bioluminescent species have been reported in this group so far. Interestingly, the study of shark squamation has been revealed in recent years to be a good proxy for inferring some ecological aspects of poorly known species. In particular, the high morphological specificity of the dermal denticles and the squamation patterns of all currently-known bioluminescent sharks could constitute a potential tool for predicting bioluminescence in both fossil and living taxa. Following this idea, we provide the first evidence supporting t…
FLOW PATTERN AROUND THE RIGID CEPHALIC SHIELD OF THE DEVONIAN AGNATHANERRIVASPIS WAYNENSIS(PTERASPIDIFORMES: HETEROSTRACI)
Palaeozoic armoured agnathans (or ostraco- derms) are characterised by having an external, bone shield enclosing the anterior part of their bodies, which demon- strate great diversity of both forms and sizes. The functional significance of these cephalic shields remains unclear (they may have been a functional analogue of the vertebral col- umn, or merely afforded protection). Here we assess the importance of the cephalic shield in terms of locomotion. In order to do this, we have studied flow patterns of the Devo- nian heterostracan Errivaspis waynensis (White, 1935), using an anatomically correct model of E. waynensis positioned at different pitching angles. The fluid flow was visualised …
New insights into the enameloid microstructure of batoid fishes (Chondrichthyes)
Chondrichthyan teeth are capped with a hypermineralized tissue known as enameloid. Its microstructure displays a hierarchical organization that has increased in structural complexity from a homogenous single-crystallite enameloid (SCE) in early Chondricthyans to the complex multilayered enameloid found in modern sharks (consisting of bundles of crystallites arranged in intriguing patterns). Recent analyses of the enameloid microstructure in batoid fishes, focused on Myliobatiformes and fossil taxa, point to the presence of a bundled (or fibred) multilayered enameloid, a condition proposed as plesiomorphic for Batoidea. In this work, we provide further enameloid analysis for a selection of t…
Grist for Riedl's mill: A network model perspective on the integration and modularity of the human skull
This research project was supported by Grant BFU2008‐00643 to D.R.G. from the Spanish Ministerio de Ciencia e Innovacion as well as project CGL2012‐37279 to M.B., from the Spanish Ministerio de Economia y Competitividad.
Translating taxonomy into the evolution of conodont feeding ecology
Conodont research has long been divided between utilitarian applications to solve geological problems versus analysis of their palaeobiology. However recent advances in conodont functional analysis allow these independent stands of research to be unified, decoding the functional implications of their morphological variation. We demonstrate this using synchrotron tomography and Finite Element Analysis, informed by occlusal and microwear analyses, to analyze functionally the classic evolutionary sequence of the genus Polygnathus. Our study shows that the evolution of the platform in Polygnathus occurred to accommodate and dissipate the stress accumulation derived from the tooth-like function …
Obruchevacanthus ireneae gen. et sp. nov., a new ischnacanthiform (Acanthodii) from the Lower Devonian of Spain
New disarticulated material of typically ischnacanthid scales, tooth whorls, and ?dentigerous jaw bones that occur recurrently together at many levels of the Lower Devonian of the Iberian Chain (Spain) is described. Based on their stratigraphical occurrence, histological evidence and comparison with similar ischnacanthid assemblages from other localities, we suggest including all of these remains in a unique natural assemblage, Obruchevacanthus ireneae gen. et sp. nov. This new taxon shares some features with Trundlelepis cervicostulata from the Lower Devonian of New South Wales (Australia), as for example the presence of a poorly developed pore canal system in their scales. This feature su…
Pre-Messinian ecological diversity of Mediterranean sharks revealed by the study of their dermal denticles
The Messinian salinity crisis (~5.59 Ma) is regarded as one of the most determining events for the recent evolutionary and palaeobiogeographic history of the Mediterranean biota. The impact of such episode has usually been assessed by evaluating the associated taxonomic turnover, however its consequences have rarely been interpreted from an ecological perspective. Here we assess the functional diversity of shark dermal denticles in a Serravallian locality from southeast Spain, providing a primary view into the pre-Messinian ecological diversity of shark communities from the Western Mediterranean. Our results reveal a high diversity of functional types of dermal denticles including dermal de…
New insights into the diversity dynamics of Triassic conodonts
International audience; In this paper, we examine the diversity trends and the evolutionary patterns of Triassic conodonts through a newly powered large-scale data-set compiled directly from the primary literature. Paleodiversity dynamics analyses have been undertaken by working at the species level and using a system of time units based on biozone subdivisions for a fine temporal level resolution. The role of heterogeneous duration of taxa in diversity estimates has been evaluated through the probabilistic profiles. Results reveal three different stages in the diversity behaviour of Triassic conodonts from standing metrics delimited by two inflections at the mid-Anisian and mid-Carnian. Su…
Exploring the major depletions of conodont diversity during the Triassic
International audience; In this paper, we show that the Triassic fossil record reflects just two great depletions of conodont diversity before the Rhaetian, which occurred in the Smithian (Olenekian, Early Triassic) and in the Julian (Carnian, Late Triassic). By exploring this context, our results highlighted that they respond to different origination?extinction dynamics. Thus, while the Smithian diversity depletion can be interpreted as a consequence of elevated extinction, the Julian diversity depletion was triggered by fluctuations in origination regime. This evidence suggests that, despite the role of extinction on diversity losses, conodonts suffered crucial changes on the origination …
Testing models of dental development in the earliest bony vertebrates, Andreolepis and Lophosteus
Theories on the development and evolution of teeth have long been biased by the fallacy that chondrichthyans reflect the ancestral condition for jawed vertebrates. However, correctly resolving the nature of the primitive vertebrate dentition is challenged by a dearth of evidence on dental development in primitive osteichthyans. Jaw elements from the Silurian–Devonian stem-osteichthyansLophosteusandAndreolepishave been described to bear a dentition arranged in longitudinal rows and vertical files, reminiscent of a pattern of successional development. We tested this inference, using synchrotron radiation X-ray tomographic microscopy (SRXTM) to reveal the pattern of skeletal development preser…
Middle-Late Triassic chondrichthyans remains from the the Betic Range (Spain)
Purpose In the present study, we described, for first time, the chondrichthyan fauna from several Middle-Late Triassic sections in the Betic Domain and compare it with other recent described coeval faunas from the Iberian Ranges. Methods Specimens were retrieved after the dissolution (with 10% acetic acid) of carbonate rocks. Results The assemblage comprises of seven species belonging to six genera (Hybodus plicatilis, Omanoselache bucheri comb. nov., Omanoselache contrarius comb. nov., Lonchidion derenzii, Lissodus aff. L. lepagei, Pseudodalatias henarejensis and cf. Rhomaleodus budurovi), most of them non-nesoselachian. Chondrichthyans remains occur in levels dating from Ladinian to Carni…
The evolution of gigantism in active marine predators
A novel hypothesis to better understand the evolution of gigantism in active marine predators and the diversity of body sizes, feeding strategies and thermophysiologies of extinct and living aquatic vertebrates is proposed. Recent works suggest that some aspects of animal energetics can act as constraining factors for body size. Given that mass-specific metabolic rate decreases with body mass, the body size of active predators should be limited by the high metabolic demand of this feeding strategy. In this context, we propose that shifts towards higher metabolic levels can enable the same activity and feeding strategy to be maintained at bigger body sizes, offering a satisfactory explanatio…
Middle Triassic sharks from the Catalan Coastal ranges (NE Spain) and faunal colonization patterns during the westward transgression of Tethys
Abstract Palaeogeographic changes that occurred during the Middle Triassic in the westernmost Tethyan domain were governed by a westward marine transgression of the Tethys Ocean. The transgression flooded wide areas of the eastern part of Iberia, forming new epicontinental shallow-marine environments, which were subsequently colonized by diverse faunas, including chondrichthyans. The transgression is recorded by two successive transgressive–regressive cycles: (1) middle–late Anisian and (2) late Anisian–early Carnian. Here, we describe the chondrichthyan fauna recovered from several Middle Triassic stratigraphic sections (Pelsonian-Longobardian) located at the Catalan Coastal Basin (western…
There is no general model for occlusal kinematics in conodonts
Knowledge of conodont element function is based largely on analysis of morphologically similar P-1 elements of few comparatively closely related species known from abundant articulated remains. From these, a stereotypical pattern of rotational occlusion has been inferred, leading to the suggestion that this may represent a general model for ozarkodinin P-1 elements at the very least. We test the generality of this occlusal model through functional analysis of Pseudofurnishius murcianus P-1 elements which, though superficially similar to homologous elements in gnathodids, evolved their platform morphology independently, through a different mode of morphogenesis, and in a different topologica…
Tooth replacement rates in early chondrichthyans: a qualitative approach
The continuous replacement of teeth throughout their lifetime is a common characteristic of most chondrichthyans. This process was already present in the earliest representatives of the group. It has been well established that different species of extant sharks show rapid tooth replacement rates; however, some authors have suggested that in early chondrichthyans this rate might have been much slower. Here we present a qualitative approach to analyse tooth replacement rates in the Early Devonian shark Leonodus carlsi, the earliest tooth-bearing shark known to date. For this, we have examined 1,103 isolated teeth from Celtiberia, Spain. Our study provides strong evidences of an extremely slow…
Morphometric Discriminant Analysis of isolated chondrichthyan scales for palaeoecological inferences: the Middle Triassic of the Iberian Chain (Spain) as a case of study
[EN] Palaeontological studies on exosqueletal disarticulated remains of chondrichthyans have focused on teeth and only less interest has been paid to scales due their limited taxonomic and systematic significance. However, classical works linking the morphology and the function of the squamation in extant sharks suggest that, despite their limited taxonomic value, the study of isolated scales can be a useful tool for palaeoenvironmental and palaeoecological inferences. Following this idea, we have analyzed the fossil record of shark scales from two Middle Triassic sections of the Iberian Chain (Spain), identifying different functional types by means of a morphometric discriminant analysis. …
Tiburones de aguas profundas en el Mioceno medio de Alicante: implicaciones para la recuperación de los ecosistemas abisales mediterráneos después de la crisis mesiniense
El estudio de la localidad clásica de El Chorrillo del Mioceno Medio (Serravaliense) en la Provincia de Alicante, ha permitido la recuperación de una interesante asociación de microrestos de tiburones caracterizada por al menos siete taxones (Deania calcea, ¿Isistius triangulus, ¿Squaliolus cf. schaubi, ¿Paraetmopterus sp., Pristiophorus sp., Scyliorhinus sp. y un Squaliforme indeterminado) pertenecientes a tres órdenes diferentes (Squaliformes, Pristiophoriformes y Carcharhiniformes). Asociados a estos microrestos se han identificado también macrodientes pertenecientes a Cosmopolitodus hastalis, Isurus sp., Hemipristis serra, Odontaspis sp., Carcharhinus spp. y ¿Otodus (Megaselachus) megal…
Computational Fluid Dynamics Suggests Ecological Diversification among Stem-Gnathostomes.
Summary The evolutionary assembly of the vertebrate bodyplan has been characterized as a long-term ecological trend toward increasingly active and predatory lifestyles, culminating in jawed vertebrates that dominate modern vertebrate biodiversity [ 1 , 2 , 3 , 4 , 5 , 6 , 7 , 8 ]. This contrast is no more stark than between the earliest jawed vertebrates and their immediate relatives, the extinct jawless, dermal armor-encased osteostracans, which have conventionally been interpreted as benthic mud-grubbers with poor swimming capabilities and low maneuverability [ 9 , 10 , 11 , 12 ]. Using computational fluid dynamics, we show that osteostracan headshield morphology is compatible with a dive…
A Serravallian (Middle Miocene) shark fauna from Southeastern Spain and its palaeoenvironment significance
The study of a new Serravallian (Middle Miocene) locality from the Southeastern Spain has yielded a shark assemblage characterized by microremains of at least seven taxa (Deania calcea, ¿Isistius triangulus, ¿Squaliolus cf. S. schaubi, ¿Paraetmopterus sp., Pristiophorus sp., Scyliorhinus sp. and a cf. Squaliformes indet) of three different orders (Squaliformes, Pristiophoriformes and Carcharhiniformes). In addition, associated macroremains have also been found, including teeth of ¿Cosmopolitodus hastalis, Isurus sp., Hemipristis serra, Odontaspis sp., Carcharhinus spp. and ¿Otodus (Megaselachus) megalodon. The assemblage contains taxa with disparate environmental preferences including not o…
Supplementary methods, figures, tables and information from Use of nursery areas by the extinct megatooth shark Otodus megalodon (Chondrichthyes: Lamniformes)
Nursery areas are fundamental for the success of many marine species, particularly for large, slow-growing taxa with low fecundity and high age of maturity. Here, we examine the population size-class structure of the extinct gigantic shark Otodus megalodon in a newly described middle Miocene locality from Northeastern Spain, as well as in eight previously known formations (Temblor, Calvert, Pisco, Gatun, Chucunaque, Bahía Inglesa, Yorktown and Bone Valley). In all cases, body lengths of all individuals were inferred from dental parameters and the size-class structure was estimated from Kernel probability density functions and Gaussian mixture models. Our analyses support the presence of fiv…
Biomechanics of Machaeracanthus pectoral fin spines provide evidence for distinctive spine function and lifestyle among early chondrichthyans
Biomechanics of Machaeracanthus pectoral fin spines provide evidence for distinctive spine function and lifestyle among early chondrichthyans
R scripts of performed analyses from Use of nursery areas by the extinct megatooth shark Otodus megalodon (Chondrichthyes: Lamniformes)
Nursery areas are fundamental for the success of many marine species, particularly for large, slow-growing taxa with low fecundity and high age of maturity. Here, we examine the population size-class structure of the extinct gigantic shark Otodus megalodon in a newly described middle Miocene locality from Northeastern Spain, as well as in eight previously known formations (Temblor, Calvert, Pisco, Gatun, Chucunaque, Bahía Inglesa, Yorktown and Bone Valley). In all cases, body lengths of all individuals were inferred from dental parameters and the size-class structure was estimated from Kernel probability density functions and Gaussian mixture models. Our analyses support the presence of fiv…
Data from: Patterns of ecological diversification in thelodonts
Here we explore the spatial, temporal and phylogenetic patterns of ecological diversification for the entire clade of thelodonts, one of the earliest groups of vertebrates and longest lasting of the Palaeozoic agnathans in the fossil record. Parsimony and maximum-likelihood methods are used to reconstruct ancestral states of their geographical distributions, habitats and lifestyles. Our results support the concept that thelodonts originated during the Middle?–Late Ordovician probably in marine open waters of Laurasia, with a demersal lifestyle on hard substrates being the ancestral condition for the whole clade. Later, thelodonts underwent a complex ecological diversification and palaeobiog…
New insights into the diversity dynamics of Triassic conodonts
In this paper, we examine the diversity trends and the evolutionary patterns of Triassic conodonts through a newly powered large-scale data-set compiled directly from the primary literature. Paleodiversity dynamics analyses have been undertaken by working at the species level and using a system of time units based on biozone subdivisions for a fine temporal level resolution. The role of heterogeneous duration of taxa in diversity estimates has been evaluated through the probabilistic profiles. Results reveal three different stages in the diversity behaviour of Triassic conodonts from standing metrics delimited by two inflections at the mid-Anisian and mid-Carnian. Survivorship analysis supp…
Illustration for social media and potential cover image from Use of nursery areas by the extinct megatooth shark Otodus megalodon (Chondrichthyes: Lamniformes)
Credits: Hugo Salais (Metazoa Studio)
New insights into Late Devonian vertebrates and associated fauna from the Cuche Formation (Floresta Massif, Colombia)
New vertebrate remains are reported from the Late Devonian (?Frasnian) Cuche Formation of northeastern Colombia, including a new taxon of antiarch placoderm (Colombialepis villarroeli, gen. et sp. nov., previously reported as Asterolepis) and a new taxon of arthrodiran placoderm (Colombiaspis rinconensis, gen. et sp. nov.). We also report evidence of a stegotrachelid actinopterygian, a diplacanthid acanthodian (cf. Florestacanthus morenoi), a second antiarch placoderm (Bothriolepis sp.), a putative megalichthyid, and a putative tristichopterid. The absence of typical Euramerican markers, e.g., Asterolepis and Strepsodus, in this assemblage suggests that faunal interchange between Euramerica…