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.     

Elasmobranchs in southern Indonesian fisheries: the fisheries, the status of the stocks and management options

The biology of elasmobranchs makes them very vulnerable to fishing pressure and there is increasing international concern over their exploitation. In northern Australia the stocks of some species may be shared with those in southern Indonesia. Indonesia has the highest landings of elasmobranchs worldwide (>100,000 t p.a.) and millions of Indonesian artisanal fishers rely heavily on elasmobranchs taken in target fisheries. They are also taken by industrial trawlers and as bycatch in pelagic tuna fisheries. This paper, resulting from a collaborative project between Australia and Indonesia, summarises the elasmobranch fisheries; the characteristics of the fisheries are outlined, the status of the stocks are assessed, and management options described and discussed. The project focussed on representative markets and fish landing sites in southern Indonesia from 2001 to 2005. Data were from market surveys, the records of the Indonesian Directorate General of Capture Fisheries, and from research cruises. Data from the ongoing tuna monitoring programme showed that shark bycatch from the tuna fleets forms about 11% of shark landings in Indonesia. Yield per recruit and related analyses were used to integrate biological information to indicate the productivity of each species to allow for management policy options and constraints. Research cruise data show that catch rates of elasmobranchs in the Java Sea declined by at least one order of magnitude between 1976 and 1997. The results indicate strongly that many of the shark and ray species in Indonesia are overfished and that the most effective management strategy may need to involve capacity control, such as licencing, gear restrictions and catch limits, together with controls on the fin trade.     

Morphological abnormalities in elasmobranchs

A total of 10 abnormal free‐swimming (i.e. post‐birth) elasmobranchs are reported from The (Persian–Arabian) Gulf, encompassing five species and including deformed heads, snouts, caudal fins and claspers. The complete absence of pelvic fins in a milk shark Rhizoprionodon acutus may be the first record in any elasmobranch. Possible causes, including the extreme environmental conditions and the high level of anthropogenic pollution particular to The Gulf, are briefly discussed.     

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.     

The distribution,diversity and abundance of elasmobranch fishes in a modified subtropical estuarine system in Kuwait

To address the paucity of information on the spatial and temporal composition of elasmobranch fish assemblages in the Arabian region, data from fisheries surveys around Kuwait's Boubyan Island, in the Tigris‐Euphrates‐Karun delta at the head of the Arabian (Persian) Gulf were examined. A total number of 461 elasmobranchs consisting of at least 13 taxa were captured by gillnets (300 m length, 4, 4.5, 5.5, and 5.75 inch stretch mesh) and otter trawls (5‐m chain line length, 34 and 10 mm stretch mesh in body and cod end, respectively) from February 2004 to March 2005. Seventy‐nine gillnet sets averaged 24 min soak time, while 431 5‐min trawl tows averaged 4.1 km h^?1 for a swept area of 1025 m² tow^?1 with a net spread of 3 m. Most taxa were infrequently recorded. Of the six species that accounted for over 90% of individual abundance, the longheaded eagle ray Aetobatus flagellum (170, 269.5 kg) and the scaly whipray Himantura imbricata (137, 17.3 kg) ranked first and second, respectively. Most sharks were Arabian carpetshark Chiloscyllium arabicum (22, 7.7 kg) and milk shark Rhizoprionodon acutus (21, 2.3 kg). Species richness for gillnets was greatest along the main Gulf coast (H' = 1.87), although H. imbricata and A. flagellum were widespread. Based on published birth sizes, neonates or young‐of‐year of several species were recorded. Results indicated that the water temperature strongly determined seasonal distribution: no elasmobranchs were recorded in the winter following a 14°C drop after October, and diversity, abundance and biomass generally increased in summer. While salinity varied from 28.4 to 39.8, major anthropogenic modifications in the upper Tigris‐Euphrates watersheds and development of Boubyan Port will result in higher salinities, which in turn will influence longer‐term patterns of elasmobranch distribution.     

Direct and indirect fishery effects on small coastal elasmobranchs in the northern Gulf of Mexico

Globally, bycatch in tropical/subtropical shrimp trawl and longline fisheries is threatening many marine species. Here we examine the joint effects of increased mortality caused by shrimp trawling bycatch, and reduced predation caused by losses of large sharks because of longline fishing. Research surveys in the Gulf of Mexico (1972–2002) demonstrated precipitous declines in shallow water coastal elasmobranchs where shrimping effort was highest (bonnethead 96%, Bancroft's numbfish (lesser electric ray) 98%, smooth butterfly ray > 99%) and consistent increases in deeper water elasmobranchs (Atlantic angel shark, smooth dogfish). These increases are the first empirical support for predation release caused by the loss of large sharks, which have been theorized to structure tropical/subtropical marine ecosystems. Bycatch of elasmobranchs in shrimp trawls is a critical conservation concern which is not solved by present mitigation measures; similar loss of elasmobranchs is expected to be occurring in tropical/subtropical regions worldwide where ever intensive shrimp trawling occurs.     

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.     

A review of elasmobranch research in the Red Sea

Given the global concern about the status of elasmobranch fishes, the paucity of information on elasmobranchs in the Red Sea is worrisome. Management of elasmobranchs in areas other than the Red Sea has been helped by research on population ecology, reproductive biology and resource partitioning, subjects that are virtually absent in the Red Sea elasmobranch literature. This review provides the first comprehensive summary of elasmobranch biology in the Red Sea with the aim of facilitating research in a region that remains remarkably under-studied.     

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.     

Box-Jenkins analysis for shark landings in Costa Rica

Sharks are highly vulnerable to intense and prolonged fishery extraction. This article analyzes the data on shark landings from the artisan fishing fleet on Costa Rica's Pacific coast between 1988 and 1997. The data come from an invoicing system administered by the Costa Rican Fisheries Institute (Instituto Costarricense de la Pesca y Acuacultura, INCOPESCA). Pacific coast shark fishing during the period under study represented approximately 20% of the total national fisheries volume. According to data from the invoicing system, the Northern Pacific region was the most productive, reporting 58% of the shark catch nationwide. Within this region, shark fishing in Papagayo Gulf represented 91% and 53% of the landings by fishery region and nationwide, respectively. The mid-sized and advanced (length of boat > 10 meters) artisan fishing fleets reported 96% of the shark catches in the zone. The study of shark fisheries in the Papagayo Gulf zone is crucial for an understanding of fishery dynamics for this resource at the national level. A monthly chronological series was constructed with the landings in the Papagayo Gulf zone, and a Univariate Box-Jenkins (UBJ) Model was estimated for first-order moving averages MA(1) with a seasonal component of the Yt = lamda(t-1) + gammaS12 + a(t) type.     

The CALFISH database: A century of California's non-confidential fisheries landings and participation data

California's commercial and recreational fisheries support vibrant coastal economies and communities. Maintaining healthy fishing communities into the future requires a detailed understanding of their past. The California Department of Fish and Wildlife (CDFW) has been monitoring statewide fisheries landings and participation since 1916 and releases confidential versions of this data through authorized data requests and non-confidential summaries of this data in its quasi-annual landings reports. The non-confidential data published in the landings reports provide a rich history of California's fisheries but are scattered across 1000s of tables in 100 s of documents, limiting their accessibility to researchers, fishers, and other interested stakeholders. We reviewed the 58 landings reports published from 1929 to 2020 and extracted and carefully curated 13 datasets with long time series and wide public interest. These datasets include: (1) annual landings in pounds and value by port and species from 1941 to 2019; (2) annual number of commercial fishing vessels by length class from 1934 to 2020; (3) annual number of licensed commercial fishers by area of residence from 1916 to 2020; and (4) annual number of party boat (CPFV) vessels, anglers, and their total catch by species from 1936 to 2020. Notably, we harmonized port names, species common names, and species scientific names across all years and datasets. We make these curated datasets, collectively called the CALFISH database, publicly available to any interested stakeholder in the supplementary materials of this paper, on an open-access data-repository, and in the wcfish R package. These datasets can be used (1) to understand the historical context of California's fisheries; (2) for original research requiring only summaries of historical landings and participation data; and (3) to anticipate the likely characteristics of confidential data requested from the state. We conclude the paper by identifying key principles for increasing the accessibility and utility of historical fisheries landings and participation data.     

A review of the application of molecular genetics for fisheries management and conservation of sharks and rays

Since the first investigation 25 years ago, the application of genetic tools to address ecological and evolutionary questions in elasmobranch studies has greatly expanded. Major developments in genetic theory as well as in the availability, cost effectiveness and resolution of genetic markers were instrumental for particularly rapid progress over the last 10 years. Genetic studies of elasmobranchs are of direct importance and have application to fisheries management and conservation issues such as the definition of management units and identification of species from fins. In the future, increased application of the most recent and emerging technologies will enable accelerated genetic data production and the development of new markers at reduced costs, paving the way for a paradigm shift from gene to genome-scale research, and more focus on adaptive rather than just neutral variation. Current literature is reviewed in six fields of elasmobranch molecular genetics relevant to fisheries and conservation management (species identification, phylogeography, philopatry, genetic effective population size, molecular evolutionary rate and emerging methods). Where possible, examples from the Indo-Pacific region, which has been underrepresented in previous reviews, are emphasized within a global perspective.     

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.     

Life-history,exploitation and extinction risk of the data-poor Baraka's whipray (Maculabatis ambigua) in small-scale tropical fisheries

The Baraka's whipray (Maculabatis ambigua) is a major constituent of small-scale fisheries catch in the south-western Indian Ocean. Despite this, little is known of its life-history or exploitation status. We provide the first estimates of crucial life-history parameters and the maximum intrinsic population growth rate r_max , using specimens collected from small-scale fisheries landings in Kenya, Zanzibar and Madagascar (with northern Madagascar representing a range extension for this species). We assess the relative risk of overexploitation by combining r_max with estimates of total Z , fishing F , and natural M mortality, and an estimate of the exploitation ratio E . The data indicate that Baraka's whipray is a medium-sized, fast-growing, early maturing species, with a relatively long lifespan. This results in a high r_max relative to many other elasmobranchs, which when combined with estimates of F suggests that the species is not at imminent risk of extinction. Yet, estimates of exploitation ratio E indicate likely overfishing for the species, with full recruitment to the fishery being post-maturation and exploitation occurring across a broad range of age and size classes. Thus, Baraka's whipray is unlikely to be biologically sustainable in the face of current fisheries pressures. This paper makes an important contribution to filling the gap in available data and is a step towards developing evidence-based fisheries management for this species. Further, it demonstrates a simple and widely applicable framework for assessment of data-poor elasmobranch exploitation status and extinction risk.     

Urea based osmoregulation and endocrine control in elasmobranch fish with special reference to euryhalinity

Since the landmark contributions of Homer Smith and co-workers in the 1930s there has been a considerable advance in our knowledge regarding the osmoregulatory strategy of elasmobranch fish. Smith recognised that urea was retained in the body fluids as part of the ‘osmoregulatory ballast’ of elasmobranch fish so that body fluid osmolality is raised to a level that is iso- or slightly hyper-osmotic to that of the surrounding medium. From studies at that time he also postulated that many marine dwelling elasmobranchs were not capable of adaptation to dilute environments. However, more recent investigations have demonstrated that, at least in some species, this may not be the case. Gradual acclimation of marine dwelling elasmobranchs to varying environmental salinities under laboratory conditions has demonstrated that these fish do have the capacity to acclimate to changes in salinity through independent regulation of Na⁺, Cl⁻ and urea levels. This suggests that many of the presumed stenohaline marine elasmobranchs could in fact be described as partially euryhaline. The contributions of Thomas Thorson in the 1970s demonstrated the osmoregulatory strategy of a fully euryhaline elasmobranch, the bull shark, Carcharhinus leucas, and more recent investigations have examined the mechanisms behind this strategy in the euryhaline elasmobranch, Dasyatis sabina. Both partially euryhaline and fully euryhaline species utilise the same physiological processes to control urea, Na⁺ and Cl⁻ levels within the body fluids. The role of the gills, kidney, liver, rectal gland and drinking process is discussed in relation to the endocrine control of urea, Na⁺ and Cl⁻ levels as elasmobranchs acclimate to different environmental salinities.     

Cephalopod and Groundfish Landings: Evidence for Ecological Change in Global Fisheries?

Cephalopod fisheries are among the few still with some local potential for expansion; in fact, as groundfish landings have declined globally, cephalopod landings have increased. We propose the hypothesis that, although increased cephalopod landings may partly reflect increased market demand, overfishing groundfish stocks has positively affected cephalopod populations. Data from 15 key FAO areas reveal that, with the exception of the north- east Atlantic, cephalopod landings have increased significantly over the last 25 years while groundfish have risen more slowly, remained stable, or declined. In terms of volume, cephalopods have not replaced groundfish. This is hypothesized as owing to the shorter life cycle of cephalopods, and rapid turnover and lower standing stocks than for longer-lived finfish species. Under high fishing pressure, groundfish are probably poor competitors, having less opportunity for spawning and replacement. In West Africa, the Gulf of Thailand and Adriatic there is strong circumstantial evidence that fishing pressure has changed ecological conditions and cephalopod stocks have increased as predatory fish have declined. We recommend that this hypothesis be tested thoroughly in other areas where suitable data exist. Most coastal and shelf cephalopod fisheries are likely to be fully exploited or overexploited, and current annual fluctuations in cephalopod landings are probably largely environmentally-driven.     

Recovery potential and conservation options for elasmobranchs

Many elasmobranchs have experienced strong population declines, which have been largely attributed to the direct and indirect effects of exploitation. Recently, however, live elasmobranchs are being increasingly valued for their role in marine ecosystems, dive tourism and intrinsic worth. Thus, management plans have been implemented to slow and ultimately reverse negative trends, including shark-specific (e.g. anti-finning laws) to ecosystem-based (e.g. no-take marine reserves) strategies. Yet it is unclear how successful these measures are, or will be, given the degree of depletion and slow recovery potential of most elasmobranchs. Here, current understanding of elasmobranch population recoveries is reviewed. The potential and realized extent of population increases, including rates of increase, timelines and drivers are evaluated. Across 40 increasing populations, only 25% were attributed to decreased anthropogenic mortality, while the majority was attributed to predation release. It is also shown that even low exploitation rates (2-6% per year) can halt or reverse positive population trends in six populations currently managed under recovery plans. Management measures that help restore elasmobranch populations include enforcement or near-zero fishing mortality, protection of critical habitats, monitoring and education. These measures are highlighted in a case study from the south-eastern U.S.A., where some evidence of recovery is seen in Pristis pectinata, Galeocerdo cuvier and Sphyrna lewini populations. It is concluded that recovery of elasmobranchs is certainly possible but requires time and a combination of strong and dedicated management actions to be successful.     

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.     

Effects of biological traits on capture-induced parturition in a freshwater stingray and perspectives for species management

Elasmobranchs are particularly vulnerable to overexploitation and population depletion, especially due to their life-history traits, such as low reproductive output and slow growth. Given that capture-induced parturition (abortion or premature birth) is a common consequence of fisheries in elasmobranchs, but still little studied, we investigated how the abortion/premature birth process varies in response to reproductive traits in a freshwater stingray, Potamotrygon amandae. Our results revealed that capture-induced parturition was affected by reproductive traits, such as litter size (one to seven) and gestation stage. The event occurred faster in pregnant females with high litter size during late pregnancy. Also, as found in other elasmobranchs, litter size was positively correlated with maternal size. These findings indicate that larger pregnant females in late pregnancy are more vulnerable to capture-induced parturition. This study improves our understanding of the capture-induced parturition process in stingrays, and provides useful information for management strategies and future recommendations for elasmobranch conservation.