Illuminating the evolution of bioluminescence in sharks

The evolutionary context in which shark bioluminescence originated is poorly understood, despite it being critical to uncovering influential factors in the evolutionary history and diversity of living chondrichthyans as well as the mechanisms of deep-water colonization by vertebrates. This study provides the first joint reconstruction of the habitats, lifestyles, and occurrence of bioluminescence in the evolution of squalomorph sharks using ancestral state estimation analysis to resolve the timing of deep-sea colonization, the evolutionary origin of bioluminescence and the ancestral ecologies of this group. The results suggest that most squalomorphs originated in neritic environments from where they colonized deep waters on several independent occasions during the Late Jurassic and Early Cretaceous, predating most of the previous estimates of the timing of this event. The colonization of the deep sea took place via the benthic zone, in contrast to the view that an intermediate mesopelagic stage occurred during this ecological transition. Finally, the analyses accounting for uncertainty of the presence of bioluminescence strongly support that this trait evolved only once among sharks in a bathydemersal ancestor. This study reveals that shark bioluminescence evolved in a complex scenario that combines elements of several previous proposals, and enriches our perspective on the sequence of events that characterized the vertebrate conquest of the deep sea

Evidence of endothermy in the extinct macropredatory osteichthyan <i>Xiphactinus audax</i> (Teleostei, Ichthyodectiformes)

Xiphactinus audax is the largest macropredatory osteichthyan ever known (Everhart et al., 2010). Some of the largest specimens exceed 5 m in total length, although the discovery of a few large, isolated teeth suggests that this teleost could reach even larger body sizes (Vavrek et al., 2016, and references therein). Fossil remains of this species have only been reported from the Upper Cretaceous of North America, across the Western Interior Basin (Schwimmer et al., 1997; Vavrek et al., 2016). The discovery of several virtually complete individuals in this area has provided valuable information about the anatomy, the dimensions, and the ecology of this species (Cope, 1872; Bardack, 1965). Xiphactinus audax displayed a tarpon-like body plan with a semilunate caudal fin and large caniniform teeth (Hay, 1898; Bardack, 1965; Carrillo- Briceño et al., 2012), suggestive of a highly active predatory lifestyle (Cavin et al., 2013). In fact, the emblematic 'fish-within-afish' specimen, containing an entire 2-m-long Gillicus arcuatus, provides evidence of the ability of X. audax to prey upon large, rapidly swimming fishes (Bardack, 1965). Predation entails a high energetic demand (Brown and Kotler, 2004). Ferrón et al. (2017) recently proposed that, because mass-specific metabolic rate decreases with increasing body size, highly active lifestyles (such as macropredation) cannot be maintained by an ectothermic metabolism over a specific body size. From this perspective, they argued that the punctual evolution of gigantism among macropredators was closely linked to metabolic-level shifts promoted by various factors (i.e., endothermy, highly efficient respiratory systems, warm temperatures, and high oxygen levels) and suggested that several extinct aquatic macropredators, including Xiphactinus, could have been meso- or endotherms on the basis of their body size and life history. Later, Ferrón (2017) established a useful methodology to assess the swimming energetics of extinct aquatic organisms, which can be used to interpret their metabolic levels and thermoregulatory strategies. This study presents an equivalent methodology developed for assessing locomotion energetics in extinct osteichthyans and provides evidence of endothermy in X. audax

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 number of extant shark species whose ecology is well known. Multivariate analyses have defined a characteristic squamation pattern for each ecological group, thus establishing a comparative framework for inferring lifestyles in thelodonts. We then use this information to study the squamation of the currently described 147 species of thelodonts, known from both articulated and disarticulated remains. Discriminant analysis has allowed recognizing squamation patterns comparable to those of sharks and links them to specific ecological groups. Our results suggest a remarkable ecological diversity in thelodonts. A large number of them were probably demersal species inhabiting hard substrates, within caves and crevices in rocky environments or reefs, taking advantage of the flexibility provided by their micromeric squamations. Contrary to classical interpretations, only few thelodonts were placed among demersal species inhabiting sandy and muddy substrates. Schooling species with defensive scales against ectoparasites could be also abundant suggesting that social interactions and pressure of ectoparasites were present in vertebrates as early the Silurian. The presence of species showing scales suggestive of low to moderate speed and a lifestyle presumably associated with open water environments indicates adaptation of thelodonts to deep water habitats. Scale morphology suggests that some other thelodonts were strong-swimming pelagic species, most of them radiating during the Early Devonian in association with the Nekton Revolution

Máster en Paleontología Aplicada (UV). Paleoecología, paleoclimatología y paleobiogegrafía (Power Point)

1. PALEOECOLOGÍA: INTRODUCCIÓN 1.1. Conceptos básicos 1.2. Genética de paleopoblaciones 1.3. Dinámica de paleopoblaciones 1.4. Interacciones en el paleoecosistema 2. APROXIMACIONES PALEOECOLÓGICAS 2.1. Sedimentología, asociaciones fósiles y tafonomía 2.2. Icnofósiles 2.3. Analogía funcional y 'phylogenetic bracketing' 2.4. Análisis morfométrico 2.5. Análisis geoquímico y biomarcadores 2.6. Análisis biomecánico 2.7. Análisis de redes 2.8. Modelado de nicho ecológico 2.9. Métodos comparativos filogenéticos 2.10. Análisis de disparidad y filo(morfoespacio) 2.11. Mapas de rendimiento y eficiencia de Paret

Grouping behaviour impacts on the parasitic pressure and squamation of sharks

The evolution of grouping behaviour involves a complex trade-off of benefits and costs. Among the latter, an increase in the risk of parasitic transmission is a well-documented phenomenon that has likely promoted the evolution of defensive mechanisms in aquatic vertebrates. Here, we explore the relationship between grouping behaviour, parasitic richness (∼parasitic pressure), and the evolution of potential defensive traits in the squamation of sharks through phylogenetic, standard and zero-inflation regression models. Our results demonstrate that sharks that frequently aggregate show increased parasitic pressure, which may constitute an agent of selection. Accordingly, their squamation is characterized by large-scale crown insertion angles and low-scale coverage, which are interpreted as traits that compromise parasite attachment and survival. These traits are less evident in regions of the body and ecological groups that are subjected to high abrasive stress or increased drag. Thus, the squamation of sharks responds to a compromise between various functions, where protective and hydrodynamic roles prevail over the rest (e.g. ectoparasitic defence and bioluminescence aiding). This work establishes a quantitative framework for inferring parasitic pressure and social interaction from squamation traits and provides an empirical basis from which to explore these phenomena through early vertebrate and chondrichthyan evolution

A new Meckel’s cartilage from the Devonian Hangenberg black shale in Morocco and its position in chondrichthyan jaw morphospace

Fossil chondrichthyan remains are mostly known from their teeth, scales or fin spines only, whereas their cartilaginous endoskeletons require exceptional preservational conditions to become fossilized. While most cartilaginous remains of Famennian (Late Devonian) chondrichthyans were found in older layers of the eastern Anti-Atlas, such fossils were unknown from the Hangenberg black shale (HBS) and only a few chondrichthyan teeth had been found therein previously. Here, we describe a Meckel’s cartilage from the Hangenberg black shale in Morocco, which is the first fossil cartilage from these strata. Since no teeth or other skeletal elements have been found in articulation, we used elliptical Fourier (EFA), principal component (PCA), and hierarchical cluster (HCA) analyses to morphologically compare it with 41 chondrichthyan taxa of different size and age and to evaluate its possible systematic affiliation. PCA and HCA position the new specimen closest to some acanthodian and elasmobranch jaws. Accordingly, a holocephalan origin was excluded. The jaw shape as well as the presence of a polygonal pattern, typical for tessellated calcified cartilage, suggest a ctenacanth origin and we assigned the new HBS Meckel’s cartilage to the order Ctenacanthiformes with reservations

A new Meckel's cartilage from the Devonian Hangenberg black shale in Morocco and its position in chondrichthyan jaw morphospace

Fossil chondrichthyan remains are mostly known from their teeth, scales or fin spines only, whereas their cartilaginous endoskeletons require exceptional preservational conditions to become fossilized. While most cartilaginous remains of Famennian (Late Devonian) chondrichthyans were found in older layers of the eastern Anti-Atlas, such fossils were unknown from the Hangenberg black shale (HBS) and only a few chondrichthyan teeth had been found therein previously. Here, we describe a Meckel's cartilage from the Hangenberg black shale in Morocco, which is the first fossil cartilage from these strata. Since no teeth or other skeletal elements have been found in articulation, we used elliptical Fourier (EFA), principal component (PCA), and hierarchical cluster (HCA) analyses to morphologically compare it with 41 chondrichthyan taxa of different size and age and to evaluate its possible systematic affiliation. PCA and HCA position the new specimen closest to some acanthodian and elasmobranch jaws. Accordingly, a holocephalan origin was excluded. The jaw shape as well as the presence of a polygonal pattern, typical for tessellated calcified cartilage, suggest a ctenacanth origin and we assigned the new HBS Meckel's cartilage to the order Ctenacanthiformes with reservations

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 non-closely related groups with similar lifestyles. Based on this well-known ecomorphological correlation, here we propose a useful comparative framework established on extant taxa for predicting some anatomical aspects in extinct aquatic vertebrates from palaeoecological data and vice versa. For this, we have assessed the relationship between the locomotory patterns and the morphological variability of the caudal region in extant sharks by means of geometric morphometrics and allometric regression analysis. Multivariate analyses reveal a strong morphological convergence in non-closely related shark species that share similar modes of life, enabling the characterization of the caudal fin morphology of different ecological subgroups. In addition, interspecific positive allometry, affecting mainly the caudal fin span, has been detected. This phenomenon seems to be stronger in sharks with more pelagic habits, supporting its role as a compensation mechanism for the loss of hydrodynamic lift associated with the increase in body size, as previously suggested for many other living and extinct aquatic vertebrates. The quantification of shape change per unit size in each ecological subgroup has allowed us to establish a basis for inferring not only qualitative aspects of the caudal fin morphology of extinct early vertebrates but also to predict absolute values of other variables such as the fin span or the hypocercal and heterocercal angles. The application of this ecomorphological approach to the specific case of Dunkleosteus terrelli has led to a new reconstruction of this emblematic placoderm. Our proposal suggests a caudal fin with a well-developed ventral lobe, narrow peduncle and wide span, in contrast to classical reconstructions founded on the phylogenetic proximity with much smaller placoderms known from complete specimens. Interestingly, this prediction gains support with the recent discovery of fin distal elements (ceratotrichia) in a well preserved D. terrelli, which suggests a possible greater morphological variability in placoderm caudal fins than previously thought

A new selenosteid placoderm from the Late Devonian of the eastern Anti-Atlas (Morocco) with preserved body outline and its ecomorphology

Placoderms are an extinct group of early jawed vertebrates that play a key role in understanding the evolution of the gnathostome body plan, including the origin of novelties such as jaws, teeth, and pelvic fins. As placoderms have a poorly ossified axial skeleton, preservation of the mainly cartilaginous axial and fin elements is extremely rare, contrary to the heavily mineralized bones of the skull and thoracic armor. Therefore, the gross anatomy of the animals and body shape is only known from a few taxa, and reconstructions of the swimming function and ecology are speculative. Here, we describe articulated specimens preserving skull roofs, shoulder girdles, most fins, and body outlines of a newly derived arthrodire. Specimens of the selenosteid Amazichthys trinajsticae gen. et sp. nov. display a skull roof with reticular ornamentation and raised sensory lines like Driscollaspis, a median dorsal plate with a unique sharp posterior depression, the pelvic girdle, the proportions and shape of the pectoral, dorsal, and caudal fins as well as a laterally enlarged region resembling the lateral keel of a few modern sharks and bony fishes. Our new phylogenetic analyses support the monophyly of the selenosteid family and place the new genus in a clade with Melanosteus, Enseosteus, Walterosteus, and Draconichthys. The shape of its body and heterocercal caudal fin in combination with the pronounced 'lateral keel' suggest Amazichthys trinajsticae was an active macropelagic swimmer capable of reaching high swimming speeds

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 explanation for the evolution of gigantism in active predators, including a vast quantity of fossil taxa. Therefore, assessing the metabolic ceilings of living aquatic vertebrates and the thermoregulatory strategies of certain key extinct groups is now crucial to define the energetic limits of predation and provide quantitative support for this model