Allometric Growth and Scaling of Body Form of the Spadenose Shark (Scoliodon laticaudus).

The versatility of the shark body form is suggested to be one of the key factors underlying their evolutionary success and persistence. Nevertheless, sharks exhibit a huge diversity of body forms and morphological adaptations. More subtly, it is increasingly evident that in many species, morphology varies through ontogeny. Multiple competing hypotheses exist explaining both the function of specific morphological structures and the interspecific distribution of these ontogenetic morphological shifts. However, existing studies are restricted to a small number of mostly large-bodied species. In this study, we report allometric scaling relationships from functionally important morphological structures in the spadenose shark (Scoliodon laticaudus). We find that a mosaic of isometric and allometric growth underlies the scaling trends in this species and that cases of allometry are consistent with an ontogenetic shift in diet. Moreover, our results refute suggestions that small-bodied sharks grow isometrically. Given the small number of existing studies of ontogenetic morphometry in sharks and the life-history/ecological characteristics of S. laticaudus, this study is a valuable contribution to our understanding of the adaptive value of ontogenetic morphological shifts in elasmobranchs.

Cautionary tales on the use of proxies to estimate body size and form of extinct animals.

Body size is of fundamental importance to our understanding of extinct organisms. Physiology, ecology and life history are all strongly influenced by body size and shape, which ultimately determine how a species interacts with its environment. Reconstruction of body size and form in extinct animals provides insight into the dynamics underlying community composition and faunal turnover in past ecosystems and broad macroevolutionary trends. Many extinct animals are known only from incomplete remains, necessitating the use of anatomical proxies to reconstruct body size and form. Numerous limitations affecting the appropriateness of these proxies are often overlooked, leading to controversy and downstream inaccuracies in studies for which reconstructions represent key input data. In this perspective, we discuss four prominent case studies (Dunkleosteus, Helicoprion, Megalodon and Perucetus) in which proxy taxa have been used to estimate body size and shape from fragmentary remains. We synthesise the results of these and other studies to discuss nuances affecting the validity of taxon selection when reconstructing extinct organisms, as well as mitigation measures that can ensure the selection of the most appropriate proxy. We argue that these precautionary measures are necessary to maximise the robustness of reconstructions in extinct taxa for better evolutionary and ecological inferences.

No evidence for population-level benefits of polyandry in sharks and rays.

Mating system variation refers to the spectrum between genetic monogamy and polyandry, and has important consequences for sexual conflict, sexual selection and individual fitness in animals. Theoretically this variation could also have substantial population-level effects, influencing population viability and extinction risk. Evidence for these effects is mixed, in part due to the fact that substantial environmental change is thought to be required for them to have visible demographic consequences. In this study we test for the presence of relationships between polyandry and population status in Elasmobranchii (sharks and rays). Elasmobranchii is a large vertebrate clade that exhibits substantial interspecific variation in both genetic mating system and population status, as well as being subject to intense anthropogenically-mediated environmental change. We also predict past macroevolutionary shifts in genetic mating system through elasmobranch phylogeny. Our results show that both genetic monogamy and polyandry have evolved multiple times independently within Elasmobranchii, and we suggest that both of these extremes represent alternative adaptive strategies that are favoured under discrete ecological and biological conditions. Nevertheless, there is no evidence of population-level consequences of mating system variation in elasmobranchs. These results are significant as they suggest that mating system variation in this clade is unlikely to be a major determinant of extinction vulnerability. Ultimately additional work will be required, however this study improves our understanding of the evolutionary dynamics underlying mating system variation in elasmobranchs, and the potential for resultant population-level consequences.

First record of a neonate bluntnose sixgill shark (Hexanchus griseus) from Baja California Sur, Mexico.

The bluntnose sixgill shark (Hexanchus griseus) is a wide-ranged deep-water shark species found off continental and insular shelves. Despite its global distribution, little is known about the reproductive ecology of the species, particularly with regard to the location and timing of important phenological events such as mating and pupping. In this study, we report the landing of a neonate H. griseus individual from an artisanal fishing camp in Baja California Sur, Mexico. This represents only the ninth confirmed record of the species from the Mexican Pacific and the first to report a neonate specimen in Mexican waters. We discuss this specimen in the context of the environmental conditions in which it was found, ultimately suggesting that these shallow coastal waters may be an important pupping ground for H. griseus in the region. Furthermore, the specimen was found during the winter months (whereas all previous reports have suggested H. griseus pups during the summer), implying regional variation in reproductive periodicity, or the presence of multiple reproductive events per year. This study provides novel insight into the reproductive biology of H. griseus and the ecological characteristics of the species in the Northern Mexican Pacific.

Evolutionary trends in the elasmobranch neurocranium.

The neurocranium (braincase) is one of the defining vertebrate characters. Housing the brain and other key sensory organs, articulating with the jaws and contributing to the shape of the anteriormost portion of the body, the braincase is undoubtedly of great functional importance. Through studying relationships between braincase shape and ecology we can gain an improved understanding of form-function relationships in extant and fossil taxa. Elasmobranchii (sharks and rays) represent an important case study of vertebrate braincase diversity as their neurocranium is simplified and somewhat decoupled from other components of the cranium relative to other vertebrates. Little is known about the associations between ecology and braincase shape in this clade. In this study we report patterns of mosaic cranial evolution in Elasmobranchii that differ significantly from those present in other clades. The degree of evolutionary modularity also differs between Selachii and Batoidea. In both cases innovation in the jaw suspension appears to have driven shifts in patterns of integration and modularity, subsequently facilitating ecological diversification. Our results confirm the importance of water depth and biogeography as drivers of elasmobranch cranial diversity and indicate that skeletal articulation between the neurocranium and jaws represents a major constraint upon the evolution of braincase shape in vertebrates.

The multidimensional spectrum of eco-evolutionary relationships between sharks and remoras.

Remoras are a highly specialised group of fishes known to associate with a range of marine megafauna, including elasmobranchs, cetaceans and marine reptiles. Remoras appear to benefit from these interspecific interactions through consumption of host dermal parasites or reduced cost of transport. Shark-remora associations are widely documented, yet our understanding of the costs and benefits involved in these interactions is poor. Studies frequently make claims about mutualistic, commensalistic or parasitic relationships without providing the necessary quantitative information necessary to make these claims. Here I explain why this approach is problematic, and proceed to examine shark-remora interactions in a rigorous eco-evolutionary framework. These interactions cannot be properly classified without considering net evolutionary fitness and context dependence. In reality, shark-remora interactions are best defined by a multidimensional spectrum of fitness consequences, with net fitness outcomes shifting between mutualism and parasitism (and vice versa) through space and time.

The origins and drivers of sexual size dimorphism in sharks.

While sexual size dimorphism (SSD) is abundant in nature, there is huge variation in both the intensity and direction of SSD. SSD results from a combination of sexual selection for large male size, fecundity selection for large female size and ecological selection for either. In most vertebrates, it is variation in the intensity of male-male competition that primarily underlies variation in SSD. In this study, we test four hypotheses regarding the adaptive value of SSD in sharks-considering the potential for each of fecundity, sexual, ecological selection and reproductive mode as the primary driver of variation in SSD between species. We also estimate past macroevolutionary shifts in SSD direction/intensity through shark phylogeny. We were unable to find evidence of significant SSD in early sharks and hypothesise that SSD is a derived state in this clade, that has evolved independently of SSD observed in other vertebrates. Moreover, there is no significant relationship between SSD and fecundity, testes mass or oceanic depth in sharks. However, there is evidence to support previous speculation that reproductive mode is an important determinant of interspecific variation in SSD in sharks. This is significant as in most vertebrates sexual selection is thought to be the primary driver of SSD trends, with evidence for the role of fecundity selection in other clades being inconsistent at best. While the phylogenetic distribution of SSD among sharks is superficially similar to that observed in other vertebrate clades, the relative importance of selective pressures underlying its evolution appears to differ.

Ontogenetic shifts in body form in the bull shark Carcharhinus leucas.

Recent studies have uncovered mosaic patterns of allometric and isometric growth underlying ontogenetic shifts in the body form of elasmobranch species (shark and rays). It is thought that shifts in trophic and spatial ecology through ontogeny drive these morphological changes; however, additional hypotheses relating to developmental constraints have also been posed. The bull shark (Carcharhinus leucas) is a large-bodied coastal shark that exhibits strong ontogenetic shifts in trophic and spatial ecology. In this study, we utilise a large data set covering a large number of morphological structures to reveal ontogenetic shifts in the body form of C. leucas, stratifying analyses by sex and size classes to provide fine-scale, more ecomorphologically relevant results. Our results indicate shifts in functional demands across the body through ontogeny, driven by selective pressures relating to trophic and spatial ecology driving the evolution of allometry. We also find significant differences in scaling trends between life stages, and between the sexes, highlighting the importance of utilising large, diverse datasets that can be stratified in this way to improve our understanding of elasmobranch morphological evolution. Ultimately, we discuss the implications of these results for existing ecomorphological hypotheses regarding the evolution of specific morphological structures, and pose novel hypotheses where relevant.

Heavy metal and trace element concentrations in the blood of scalloped hammerhead sharks (Sphyrna lewini) from La Paz Bay, México.

Sharks are particularly susceptible to bioaccumulation due to their life history characteristics and trophic position within marine ecosystems. Despite this, studies of bioaccumulation cover only a small proportion of extant species. In this study we report concentrations of trace elements and heavy metals in blood samples of Sphyrna lewini for the first time. We report high concentrations of several trace elements and heavy metals, with concentrations of some elements exceeding the limit determined safe for human consumption. High elemental concentrations may reflect biochemical differences between blood plasma and other tissues; however, they may also be symptomatic of high levels of exposure triggered by anthropogenic activities. We also provide evidence of elemental accumulation through ontogeny, the nature of which differs from that previously reported. Ultimately, this baseline study increases our understanding of interspecific and intraspecific variation in bioaccumulation and ecotoxicology in elasmobranchs which may prove important in ensuring adequate management.

The biology and ecology of the Pacific sharpnose shark Rhizoprionodon longurio.

Amidst global declines in elasmobranch populations resulting predominantly from overfishing, the need to gather data regarding shark ecology is greater than ever. Many species remain data deficient or at risk of going extinct before sufficient conservation measures can be applied. In this review, we summarise existing knowledge regarding the biology and ecology of the Pacific sharpnose shark Rhizoprionodon longurio (Jordan & Hilbert, 1882), a small-bodied carcharhinid shark found in coastal waters of the Eastern Tropical Pacific Ocean that is of both commercial and ecological importance. We compare ecological parameters of this species with its closest extant relatives and identify major knowledge gaps and avenues for future research. In particular, additional studies investigating the behavioural and sensory ecology, as well as potential migratory patterns of the species are needed. Such studies will not only improve our understanding of R. longurio, but provide insight into the extent to which the numerous studies performed on a close relative-Rhizoprionodon terraenovae-provide an accurate representation of the biology and ecology of Rhizoprionodon and carcharhinids more generally.

Quantifying the behavioural consequences of shark ecotourism.

Shark populations globally are facing catastrophic declines. Ecotourism has been posited as a potential solution to many of the issues facing shark conservation, yet increasingly studies suggest that such activity may negatively influence aspects of shark ecology and so further pressure declining populations. Here we combine UAV videography with deep learning algorithms, multivariate statistics and hidden Markov models (HMM) to quantitatively investigate the behavioural consequences of ecotourism in the whale shark (Rhincodon typus). We find that ecotourism increases the probability of sharks being in a disturbed behavioural state, likely increasing energetic expenditure and potentially leading to downstream ecological effects. These results are only recovered when fitting models that account for individual variation in behavioural responses and past behavioural history. Our results demonstrate that behavioural responses to ecotourism are context dependent, as the initial behavioural state is important in determining responses to human activity. We argue that models incorporating individuality and context-dependence should, wherever possible, be incorporated into future studies investigating the ecological impacts of shark ecotourism, which are only likely to increase in importance given the expansion of the industry and the dire conservation status of many shark species.

Ontogenetic morphometry of the brown smoothhound shark Mustelus henlei with implications for ecology and evolution.

The central tenet of ecomorphology links ecological and morphological variation through the process of selection. Traditionally used to rationalise morphological differences between taxa, an ecomorphological approach is increasingly being utilised to study morphological differences expressed through ontogeny. Elasmobranchii (sharks, rays and skates) is one clade in which such ontogenetic shifts in body form have been reported. Such studies are limited to a relatively small proportion of total elasmobranch ecological and morphological diversity, and questions remain regarding the extent to which ecological selection are driving observed morphometric trends. In this study, we report ontogenetic growth trajectories obtained via traditional linear morphometrics from a large data set of the brown smoothhound shark (Mustelus henlei). We consider various morphological structures including the caudal, dorsal and pectoral fins, as well as several girth measurements. We use an ecomorphological approach to infer the broad ecological characteristics of this population and refine understanding of the selective forces underlying the evolution of specific morphological structures. We suggest that observed scaling trends in M. henlei are inconsistent with migratory behaviour, but do not contradict a putative trophic niche shift. We also highlight the role of predation pressure and sex-based ecological differences in driving observed trends in morphometry, a factor which has previously been neglected when considering the evolution of body form in sharks.

The genetics-morphology-behavior trifecta: Unraveling the single greatest limitation affecting our understanding of chondrichthyan evolution.

Sharks, rays, and chimaera form the clade Chondrichthyes, an ancient group of morphologically and ecologically diverse vertebrates that has played an important role in our understanding of gnathostome evolution. Increasingly, studies seek to investigate evolutionary processes operating within the chondrichthyan crown group, with the broad aim of understanding the driving forces behind the vast phenotypic diversity observed among its constituent taxa. Genetic, morphological, and behavioral studies have all contributed to our understanding of phenotypic evolution yet are typically considered in isolation in the context of Chondrichthyes. In this viewpoint, I discuss why such isolation is prevalent in the literature, how it constrains our understanding of evolution, and how it might be overcome. I argue that integrating these core fields of organismal biology is vital if we are to understand the evolutionary processes operating in contemporary chondrichthyan taxa and how such processes have contributed to past phenotypic evolution. Despite this, the necessary tools to overcome this major limitation already exist and have been applied to other taxa.