Determinants of tapeworm species richness in elasmobranch fishes: untangling environmental and phylogenetic influences

Parasite species richness is a fundamental characteristic of host species and varies substantially among host communities. Hypotheses aiming to explain observed patterns of richness are numerous, and none is universal. In this study, we use tapeworm parasites of elasmobranch fishes to examine the phylogenetic and environmental influences on the variation in species richness for this specific system. Tapeworms are the most diverse group of helminths to infect elasmobranchs. Elasmobranchs are cosmopolitan in distribution and their tapeworm parasites are remarkably host specific; therefore, making this an ideal system in which to examine global patterns in species diversity. Here, we 1) quantify the tapeworm richness in elasmobranch fishes, 2) identify the host features correlated with tapeworm richness, and 3) determine whether tapeworm richness follows a latitudinal gradient. The individual and combined effects of host size, factors associated with water temperatures (influenced by latitude and depth), host habitat, and type of elasmobranch (shark or batoid) on measures of species diversity were assessed using general linear models. These analyses included tapeworm host records for 317 different elasmobranch species (124 species were included in our analyses) and were conducted with and without taking into account phylogenetic relationships between host species. Since sharks and batoids differ substantially in body form, analyses were repeated for each host subset. On average, batoids harboured significantly more tapeworm species than shark hosts. Tapeworm richness in sharks was influenced by median depth, whereas no predictor variable included in our models could adequately account for interspecific variation in tapeworm richness in batoid hosts. The taxonomic diversity of tapeworm assemblages of sharks and batoids was influenced by median depth and median latitude, respectively. When the influence of host phylogeny is accounted for, larger hosts harbour a greater tapeworm richness, whereas hosts exploiting wider latitudinal ranges harbour more taxonomically distinct tapeworm assemblages. Species richness and taxonomic diversity of tapeworm assemblages in elasmobranch fishes are influenced by different evolutionary pressures, including host phylogenetic relationships, space constraints and geographical area. Our results suggest that ca 3600 tapeworm species have yet to be described from elasmobranch fishes.     

Molecular phylogeny of elasmobranchs inferred from mitochondrial and nuclear markers

The elasmobranchs (sharks, rays and skates) being the extant survivors of one of the earliest offshoots of the vertebrate evolutionary tree are good model organisms to study the primitive vertebrate conditions. They play a significant role in maintaining the ecological balance and have high economic value. Due to over-exploitation and illegal fishing worldwide, the elasmobranch stocks are being decimated at an alarming rate. Appropriate management measures are necessary for restoring depleted elasmobranch stocks. One approach for restoring stocks is implementation of conservation measures and these measures can be formulated effectively by knowing the evolutionary relationship among the elasmobranchs. In this study, a total of 30 species were chosen for molecular phylogeny studies using mitochondrial cytochrome c oxidase subunit I, 12S ribosomal RNA gene and nuclear Internal Transcribed Spacer 2. Among different genes, the combined dataset of COI and 12S rRNA resulted in a well resolved tree topology with significant bootstrap/posterior probabilities values. The results supported the reciprocal monophyly of sharks and batoids. Within Galeomorphii, Heterodontiformes (bullhead sharks) formed as a sister group to Lamniformes (mackerel sharks): Orectolobiformes (carpet sharks) and to Carcharhiniformes (ground sharks). Within batoids, the Myliobatiformes formed a monophyly group while Pristiformes (sawfishes) and Rhinobatiformes (guitar fishes) formed a sister group to all other batoids.     

Molecular phylogenetic evidence refuting the hypothesis of Batoidea (rays and skates) as derived sharks

Early morphological studies regarding the evolutionary history of elasmobranchs suggested sharks and batoids (skates and rays) were respectively monophyletic. More modern morphological cladistic studies, however, have tended to suggest that batoids are derived sharks, closely related to sawsharks and angelsharks, a phylogenetic arrangement known as the Hypnosqualea hypothesis. Very few molecular studies addressing interordinal relationships of elasmobranchs have been published; the few that do exist, are very limited in terms of both taxon representation and/or aligned sequence positions, and are insufficient to answer the question of whether batoids are derived sharks. The purpose of this study was to address this issue with more complete taxon representation, concomitant with a reasonable number of aligned sequence positions. The data set included a 2.4-kb segment of the mitochondrial 12S rRNA-tRNA valine-16S rRNA locus, and in terms of taxa, representatives of two orders of Batoidea, at least one representative of all orders of sharks, and as an outgroup, the widely recognized sister group to elasmobranchs-Holocephali. The results provide the first convincing molecular evidence for shark monophyly and the rejection of the Hypnosqualea hypothesis. Our phylogenetic placement of batoids as a basal elasmobranch lineage means that much of the current thinking regarding the evolution of morphological and life history characteristics in elasmobranchs needs to be re-evaluated.     

The elasmobranchs of Malpelo Flora and Fauna Sanctuary,Colombia

Management and conservation actions in marine-protected areas require baselines for monitoring threatened marine fauna such as elasmobranchs. This article provides evidence of the occurrence of 34 species of elasmobranchs (21 sharks and 13 batoids) in the Malpelo Flora and Fauna Sanctuary, Colombia, including five new records of sharks and three of rays. From 1987 to 2021, new records were obtained by underwater visual census using SCUBA, manned submersibles and deep-ocean cameras to depths of up to 2211 m. Of the recorded species, 21 are considered as threatened taxa (64%) by the IUCN, making the Malpelo Flora and Fauna Sanctuary an essential conservation area for this highly threatened group of species.     

Phylogeny of elasmobranchs based on LSU and SSU ribosomal RNA genes

The dominant view of the phylogeny of living elasmobranchs, based on morphological characters, is that batoids (skates and rays) are derived sharks, joined with saw sharks, and angel sharks in the clade Hypnosqualea [S. Shirai, Squalean Phylogeny: A New Framework of 'Squaloid' Sharks and Related Taxa, Hokkaido University Press, Sapporo, 1992]. By contrast, a recent molecular-phylogenetic study based on mitochondrial genes for 12S and 16S rRNA and tRNA valine [C.J. Douady et al., Mol. Phylogenet. Evol., 26 (2003) 215-221] supported the older view that batoids and sharks are separate lineages. Here, we tested these two different views using combined, nuclear large-subunit and small-subunit rRNA gene sequences ( approximately 5.3kb) from 22 elasmobranchs, two chimeras, and two bony fishes. We used maximum likelihood, maximum parsimony, minimum evolution, and Bayesian inference for tree reconstruction, and found the large-subunit rRNA gene to contain far more signal than the small-subunit gene for resolving this mostly Mesozoic radiation. Our findings matched those of in separating batoids from sharks and in statistically rejecting Hypnosqualea. The angel shark (Squatina) was the sister group to squaliforms (dogfish sharks), and our findings are consistent with the idea that "orbitostylic" sharks form a monophyletic group (squaliforms+the hexanchiform Chlamydoselachus+Squatina+Pristiophorus). In the galeomorph sharks, however, lamniforms grouped with orectolobiforms, opposing the widely accepted 'lamniform+carcharhiniform' grouping. A tree based on the mitochondrial gene for cytochrome b also supported a separation of sharks and batoids, in contrast to Hypnosqualea. Among elasmobranchs, variation in the evolutionary rates of the nuclear rRNA genes was higher than that of cytochrome b genes, mainly due to the relatively rapid evolution of rRNA in some carcharhiniforms. In conclusion, several different molecular studies now refute the Hypnosqualea hypothesis of elasmobranch interrelationships.     

The neural control of feeding in elasmobranchs: A review and working model

A working model of the neural control of feeding in elasmobranchs is presented and summarized in graphic form. The model is based on a review of studies in sharks and batoids augmented by suggestions and comparisons from research in mammals and teleosts. The focal point of the model is a proposed Hypothalamic Feeding Area (HFA) that encompasses the medial periventricular zone in the inferior lobe and a small area immediately dorsal to it. Electrical stimulation in the HFA has evoked feeding in nurse sharks and neuropeptides and neurotransmitters known to influence feeding in mammals and teleosts have been localized immunocytochemically in the region in several elasmobranchs. The HFA of elasmobranchs appears to be analogous to and possibly homologous with ??hypothalamic feeding centers?? in bony fishes and tetrapods. Such ??centers?? are thought to integrate external and internal stimuli and control feeding in relation to available energy stores. The HFA??s strong olfactory connections in elasmobranchs are consistent with smell-induced feeding activities. In elasmobranchs, the HFA has reciprocal connections with the central pallium of the telencephalon, a region that processes visual, acoustic, mechanoreceptive and electroreceptive lateral line and possibly somatosensory information. These pathways may provide multisensory control in feeding. HFA connections with the cerebellum, brainstem and spinal cord most likely mediate hypothalamic co-ordination of the sensorimotor components of elasmobranch feeding. The review and model help to identify areas for suggested research.     

Morphological and histochemical characterization of the pectoral fin muscle of batoids

Batoids differ from other elasmobranch fishes in that they possess dorsoventrally flattened bodies with enlarged muscled pectoral fins. Most batoids also swim using either of two modes of locomotion: undulation or oscillation of the pectoral fins. In other elasmobranchs (e.g., sharks), the main locomotory muscle is located in the axial myotome; in contrast, the main locomotory muscle in batoids is found in the enlarged pectoral fins. The pectoral fin muscles of sharks have a simple structure, confined to the base of the fin; however, little to no data are available on the more complex musculature within the pectoral fins of batoids. Understanding the types of fibers and their arrangement within the pectoral fins may elucidate how batoid fishes are able to utilize such unique swimming modes. In the present study, histochemical methods including succinate dehydrogenase (SDH) and immunofluoresence were used to determine the different fiber types comprising these muscles in three batoid species: Atlantic stingray (Dasyatis sabina), ocellate river stingray (Potamotrygon motoro) and cownose ray (Rhinoptera bonasus). All three species had muscles comprised of two muscle fiber types (slow-red and fast-white). The undulatory species, D. sabina and P. motoro, had a larger proportion of fast-white muscle fibers compared to the oscillatory species, R. bonasus. The muscle fiber sizes were similar between each species, though generally smaller compared to the axial musculature in other elasmobranch fishes. These results suggest that batoid locomotion can be distinguished using muscle fiber type proportions. Undulatory species are more benthic with fast-white fibers allowing them to contract their muscles quickly, as a possible means of escape from potential predators. Oscillatory species are pelagic and are known to migrate long distances with muscles using slow-red fibers to aid in sustained swimming.     

Elasmobranch captures in shrimps trammel net fishery off the Gulf of Gabès (Southern Tunisia,Mediterranean Sea)

Small‐scale fisheries are generally promoted as a sustainable alternative to large‐scale industrial fisheries. However, there is recent growing evidence that small‐scale fisheries may be the largest threat to marine species of conservation concern. The objective of this study was to evaluate the potential impact of the trammel net fishery on elasmobranchs in the Gulf of Gabès, Southern Tunisia. Data are based on 191 shrimp trammel net set (40 mm stretched mesh size) surveys conducted aboard commercial fishing vessels from May to July 2009. Five species of the small coastal elasmobranchs (Mustelus mustelus (Linnaeus, 1758), Mustelus punctulatus Risso 1827, Dasyatis pastinaca (Linnaeus, 1758), Dasyatis marmorata (Steindachner, 1892) and Torpedo torpedo (Linnaeus, 1758)) and two species from the large coastal shark (Carcharhinus plumbeus (Nardo, 1827) and Carcharhinus brevipinna (Müller & Henle, 1839)) were recognized as by‐catch in this fishery. Elasmobranch by‐catch was dominated by sharks (90.3%), smoothhound sharks Mustelus sp. being by far the most important (88.9%) and reflecting their abundance in the area; 58% of the sets caught at least one specimen, with 4.8 ± 1.3 caught per set. Captures were composed essentially of neonate and juvenile sharks, while the batoids were dominated by mature individuals. This study shows that shrimp trammel nets represent a considerable source of mortality for early life stages of elasmobranch species in the Gulf of Gabès. Additionally, there was a high density of neonates and small juvenile M. mustelus in the Sfax zone, suggesting that these nearshore waters are a nursery grounds for smoothhound sharks. Further research should focus on the incidents of by‐catch and evaluate the potential solutions to allow trammel net fisheries to coexist alongside the elasmobranch species.     

Review of Elasmobranch Behavioral Studies Using Ultrasonic Telemetry with Special Reference to the Lemon Shark, Negaprion Brevirostris, Around Bimini Islands, Bahamas

A review of past behavioral ultrasonic telemetry studies of sharks and rays is presented together with previously unpublished material on the behavior of the lemon shark, Negaprion brevirostris, around the Bimini Islands, Bahamas. The review, focusing on movement behaviors of 20 shark and three ray species, reveals that elasmobranchs exhibit a variety of temporal and spatial patterns in terms of rates-of-movement and vertical as well as horizontal migrations. The lack of an apparent pattern in a few species is probably attributable to the scarcity of tracking data. Movements are probably governed by several factors, some still not studied, but data show that food, water temperature, bottom type, and magnetic gradient play major roles in a shark's decision of where and when to swim. A few species exhibit differences in behavior between groups of sharks within the same geographical area. This interesting finding warrants further research to evaluate the causes of these apparent differences and whether these groups constitute different subpopulations of the same species. The lack of telemetry data on batoids and some orders of sharks must be addressed before we can gain a more comprehensive understanding of the behavior of elasmobranch fishes. Previously unpublished data from 47 smaller and 38 larger juvenile lemon sharks, collected over the decade 1988–1998, provide new results on movement patterns, habitat selection, activity rhythms, swimming speed, rate-of-movement, and homing behavior. From these results we conclude that the lemon shark is an active predator with a strong, apparently innate homing mechanism. This species shows ontogenetic differences in habitat selection and behavior, as well as differences in movements between groups of individuals within the same area. We suggest three hypotheses for future research on related topics that will help to understand the enigmatic behavior of sharks.     

Diet,trophic interactions and possible ecological role of commercial sharks and batoids in northern Peruvian waters

The Peruvian sea represents one of the most productive ocean ecosystems and possesses one of the largest elasmobranch fisheries in the Pacific Ocean. Ecosystem-based management of these fisheries will require information on the trophic ecology of elasmobranchs. This study aimed to understand the diet, trophic interactions and the role of nine commercial elasmobranch species in northern Peru through the analysis of stomach contents. A total of 865 non-empty stomachs were analysed. Off northern Peru, elasmobranchs function as upper-trophic-level species consuming 78 prey items, predominantly teleosts and cephalopods. Two distinctive trophic assemblages were identified: (a) sharks (smooth hammerhead shark Sphyrna zygaena, thresher shark Alopias spp. and blue shark Prionace glauca) that feed mainly on cephalopods in the pelagic ecosystem; and (b) sharks and batoids (Chilean eagle ray Myliobatis chilensis, humpback smooth-hound Mustelus whitneyi, spotted houndshark Triakis maculata, Pacific guitarfish Pseudobatos planiceps, copper shark Carcharhinus brachyurus and school shark Galeorhinus galeus) that feed mainly on teleosts and invertebrates in the benthonic and pelagic coastal ecosystem. This study reveals for the first time the diet of T. maculata and the importance of elasmobranchs as predators of abundant and commercial species (i.e., jumbo squid Dosidicus gigas and Peruvian anchovy Engraulis ringens). The results of this study can assist in the design of an ecosystem-based management for the northern Peruvian sea and the conservation of these highly exploited, threatened or poorly understood group of predators in one of the most productive marine ecosystems.     

The physiological response to anthropogenic stressors in marine elasmobranch fishes: a review with a focus on the secondary response

Elasmobranchs (sharks, rays, and skates) are currently facing substantial anthropogenic threats, which expose them to acute and chronic stressors that may exceed in severity and/or duration those typically imposed by natural events. To date, the number of directed studies on the response of elasmobranch fishes to acute and chronic stress are greatly exceeded by those related to teleosts. Of the limited number of studies conducted to date, most have centered on sharks; batoids are poorly represented. Like teleosts, sharks exhibit primary and secondary responses to stress that are manifested in their blood biochemistry. The former is characterized by immediate and profound increases in circulating catecholamines and corticosteroids, which are thought to mobilize energy reserves and maintain oxygen supply and osmotic balance. Mediated by these primary responses, the secondary effects of stress in elasmobranchs include hyperglycemia, acidemia resulting from metabolic and respiratory acidoses, and profound disturbances to ionic, osmotic, and fluid volume homeostasis. The nature and magnitude of these secondary effects are species-specific and may be tightly linked to metabolic scope and thermal physiology as well as the type and duration of the stressor. In fishes, acute and chronic stressors can incite a tertiary response, which involves physiological changes at the organismal level, thereby impacting growth rates, reproductive outputs or investments, and disease resistance. Virtually no studies to date have been conducted on the tertiary stress response in elasmobranchs. Given the diversity of elasmobranchs, additional studies that characterize the nature, magnitude, and consequences of physiological stress over a broad spectrum of stressors are essential for the development of conservation measures. Additional studies on the primary, secondary, and tertiary stress response in elasmobranchs are warranted, with particular emphasis on expanding the range of species and stressors examined. Future studies should move beyond simply studying the effects of known stressors and focus on the underlying physiological mechanisms. Such studies should include the coupling of stress indicators with quantifiable aspects of the stressor, which will allow researchers to test hypotheses on survivorship and, ultimately, derive models that effectively link physiology to mortality. Studies of this nature are essential for decision-making that will result in the effective management and conservation of these species.     

How can the feeding habits of the sand tiger shark influence the success of conservation programs?

The feeding habits of the sand tiger shark Carcharias taurus , one of the most threatened sharks of the world, are poorly known. Sand tiger sharks are critically endangered in the South-west Atlantic. Since 2007, the law requires that all individuals caught in recreational fisheries off Argentina must be released. Using data from a north Patagonian recreational fishery ( n =164 stomachs with contents), we analyzed the diet of sand tiger sharks in relation with size, sex, maturity stage and season; assessed prey consumption patterns and hooking location; and estimated diet overlap with fishery landings. Sand tiger sharks consumed mainly teleosts (55.4% of the total prey number, N ) and elasmobranchs (41.84% N ), and ate more benthic elasmobranchs (batoids and angel sharks) as they become larger. Sharks swallowed prey mostly in one piece (93.7%) and were hooked mainly in internal organs (87.4%, n =175), causing occlusion and perforation of the esophagus and stomach, and lacerations to the pericardium, heart and liver. Sand tiger sharks fed on the most heavily landed species, overlapping almost completely (>90%) with fishery landings. Conservation plans should take into account that releasing hooked sharks could be insufficient to minimize fishing mortality and that competition for food with fisheries is likely to occur.     

Carbon and nitrogen discrimination factors for elasmobranch soft tissues based on a long-term controlled feeding study

The foraging ecology of elasmobranchs (sharks, skates, and rays) is difficult to study because species have spatially and temporally diverse diets. Many diet and habitat preference studies for mammals, birds, and teleosts use stable isotope analysis, but interpretations are limited for elasmobranch studies because taxon-specific isotope discrimination factors from a controlled experiment are unavailable. Trophic discrimination factors for plasma, red blood cells, and muscle were determined from an experiment with leopard sharks (Triakis semifasciata) fed a constant diet of squid over 1000?days. The ??¹³C values for shark tissues at equilibrium with the squid diet did not vary significantly among individuals, but plasma and red blood cell ??¹⁵N values differed significantly among individuals and sampling day. Individual variation of muscle ??¹⁵N averages was observed and likely related to growth. Overall, carbon and nitrogen discrimination factors corresponded to previous studies featuring high-protein diets and carnivorous taxa. The muscle-to-diet discrimination factors from the controlled feeding study were applied to blue sharks (Prionace glauca) and smooth hammerhead sharks (Sphyrna zygaena) caught offshore from Baja California, Mexico. This case study demonstrates the potential of stable isotope analysis to illuminate differences in foraging patterns between elasmobranch species.     

Spatial Heterogeneity in Fishing Creates de facto Refugia for Endangered Celtic Sea Elasmobranchs

The life history characteristics of some elasmobranchs make them particularly vulnerable to fishing mortality; about a third of all species are listed by the IUCN as Threatened or Near Threatened. Marine Protected Areas (MPAs) have been suggested as a tool for conservation of elasmobranchs, but they are likely to be effective only if such populations respond to fishing impacts at spatial-scales corresponding to MPA size. Using the example of the Celtic Sea, we modelled elasmobranch biomass (kg h⁻¹) in fisheries-independent survey hauls as a function of environmental variables and ‘local’ (within 20 km radius) fishing effort (h y⁻¹) recorded from Vessel Monitoring Systems data. Model selection using AIC suggested strongest support for linear mixed effects models in which the variables (i) fishing effort, (ii) geographic location and (iii) demersal fish assemblage had approximately equal importance in explaining elasmobranch biomass. In the eastern Celtic Sea, sampling sites that occurred in the lowest 10% of the observed fishing effort range recorded 10 species of elasmobranch including the critically endangered Dipturus spp. The most intensely fished 10% of sites had only three elasmobranch species, with two IUCN listed as Least Concern. Our results suggest that stable spatial heterogeneity in fishing effort creates de facto refugia for elasmobranchs in the Celtic Sea. However, changes in the present fisheries management regime could impair the refuge effect by changing fisher''s behaviour and displacing effort into these areas.     

A comparison of the external morphology of the membranous inner ear in elasmobranchs

Studies on the elasmobranch inner ear have focused predominantly on a small group of sharks, particularly, carcharhinids. As a result, subsequent studies in other species have subdivided species into two main groups: those typical and those atypical of carcharhinid sharks. This study proposes a different set of inner‐ear morphology groupings to those previously suggested. The inner ears from 17 species of elasmobranchs (representing both sharks and rays) are examined in this study and based on morphometric data some groups include both rays and sharks. Four groups are now proposed based predominantly on the shape and dimensions of the membranous otoconial organs, and characteristics of the semicircular canals. Evident morphological differences between the ear types belonging to the new groups include the membranes of the semicircular canals being bound to the otoconial organs in some species, while only being connected via the canal ducts in others, as well as clear variation present in saccular organ size. Previous studies examining variation in the inner ear have attributed differences to either phylogeny or functional significance. Results from this study suggest that neither phylogeny nor feeding strategy solely accounts for the morphological diversity present in the external morphology of the elasmobranch inner ear. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Species, sex, size and male maturity composition of previously unreported elasmobranch landings in Kuwait, Qatar and Abu Dhabi Emirate

This paper presents data from the first major survey of the diversity, biology and fisheries of elasmobranchs in the Persian (Arabian) Gulf. Substantial landings of elasmobranchs, usually as gillnet by-catch, were recorded in Kuwait, Qatar and the Emirate of Abu Dhabi (part of the United Arab Emirates), although larger elasmobranchs from targeted line fisheries were landed in Abu Dhabi. The elasmobranch fauna recorded was distinctive and included species that are undescribed, rare and have a highly restricted known distribution. Numerical abundance was dominated by sharks (c. 80%), of which carcharhinids were by far the most important. The milk shark Rhizoprionodon acutus and whitecheek shark Carcharhinus dussumieri together comprised just under half of all recorded individuals. Around 90% of recorded sharks were small (50-90 cm total length, L(T) ) individuals, most of which were mature individuals of species with a small maximum size (<100 cm L(T) ), although immature individuals of larger species (e.g. Carcharhinus sorrah and other Carcharhinus spp.) were also important. The most frequently recorded batoid taxa were cownose rays Rhinoptera spp., an undescribed whipray, and the granulated guitarfish Rhinobatos granulatus. The first size, sex and maturity data for a wide range of Gulf elasmobranch species are presented (including L(T) at 50% maturity for males of four shark species) and include some notable differences from other locations in the Indo-West Pacific Ocean. A number of concerns regarding the sustainability of the fishery were highlighted by this study, notably that most of the batoid species recorded are classed by the IUCN Red List as vulnerable, endangered, data deficient or not evaluated. Despite their considerable elasmobranch landings, none of the three countries sampled have developed a 'Shark Plan' as encouraged to do so under the FAO International Plan of Action: Sharks. Furthermore, Kuwait and Qatar currently report zero or no elasmobranch landings to the FAO.     

Evolution of Upper Jaw Protrusion Mechanisms in Elasmobranchs

Upper jaw protrusion is a prominent component of the feedingmechanism in most elasmobranchs and has received considerableattention over the years. In this paper, we review what is knownof muscle activity during prey capture in elasmobranchs, particularlythat of upper jaw protrusion, and evaluate the extent to whichfunctional modifications have evolved through changes in anatomyor patterns of muscle activity. To date, motor activity duringfeeding has been documented in only four species of elasmobranchs,although they represent the three major elasmobranch groups:Galea (typical sharks); Squalea (dogfish sharks); and Batoidea(skates and rays). Our efforts show that while muscles involvedin cranial elevation and lower jaw depression and elevationshow a conserved pattern of motor activity and function acrossspecies, other muscles show a more variable history. Our observationsof elasmobranch upper jaw protrusion mechanisms suggest a mosaicof character changes over the course of evolution that involveanatomical changes in all cases and modifications of muscleactivation patterns in some cases. During the evolution of feedingmechanisms of elasmobranchs, there have been two structuralchanges incorporating a pre-existing motor pattern to yieldan unmodified kinematic profile, the original preorbitalis andthe descendent preorbitalis. One additional instance of structuralmodification is accompanied by an alteration in the motor patternleading to a change in movement pattern, the levator palatoquadrati.     

Influence of environmental factors on shark and ray movement,behaviour and habitat use: a review

Any attempt to describe the spatial ecology of sharks and rays should consider the drivers responsible for movement. Research has shown fluctuations in the environment (abiotic factors) can trigger movement and changes in behaviour and habitat use for many elasmobranch species. Most studies to date have selectively focused on a small number of abiotic factors (i.e. temperature, salinity); however, other factors such as dissolved oxygen, tide, photoperiod, barometric pressure and pH have also been documented to act as drivers of movement in shark and ray species. Although usually examined individually, abiotic factors rarely act in isolation and often differ in their level of influence between species, sex, ontogenetic stage, season and geographic location. This paper reviews the role of abiotic factors as a driver of movement and changes in behaviour and habitat use in elasmobranchs. In the context of a changing climate, insight into how sharks and rays may respond to fluctuating environmental conditions projected under future scenarios is required.     

An investigation of the co-evolutionary relationships between onchobothriid tapeworms and their elasmobranch hosts

There is general consensus that the living elasmobranchs comprise a monophyletic taxon. There is evidence that, among tetraphyllidean tapeworms, the approximately 201 hooked species (Onchobothriidae) may also comprise a monophyletic group. Determinations of host specificity are contingent upon correct specific identifications. Since 1960, over 200 new elasmobranch species and over 100 new onchobothriid species have been described. Some confidence can be placed in host and parasite identifications of recent studies, but specific identifications provided in older literature in many cases are suspect. There is some consensus among published works on the phylogenetic relationships among elasmobranchs. Phylogenetic relationships among onchobothriids remain largely unresolved. Elasmobranchs have been poorly sampled for onchobothriids; records exist for approximately 20% of the 911 species and approximately 44% of the 170 elasmobranch genera. Onchobothriids are remarkably host specific, exhibiting essentially oioxenous specificity for their definitive hosts. Multiple onchobothriid species commonly parasitise the same host species; in some cases these are congeners, in other cases these are members of two different onchobothriid genera. There is substantial incongruence between available host and parasite phylogenies. For example, Acanthobothrium is by far the most ubiquitous onchobothriid genus, parasitising almost all orders of elasmobranchs known to host onchobothriids, yet, there is no evidence of major clades of Acanthobothrium corresponding to postulated major subgroupings of elasmobranchs (e.g. Galea and Squalea or sharks and rays). Potamotrygonocestus appears to be among the most basal onchobothriid groups, yet it parasitises one of the most derived elasmobranch groups (the freshwater stingray genus Potamotrygon). It appears that congeners parasitising the same host species are not necessarily each other's closest relatives. At this point the preliminary and limited available data suggest that, at least in this system, strict host specificity is not necessarily indicative of strict co-evolution. This study was extremely limited by the lack of available robust phylogenies for onchobothriids and elasmobranchs.