Pelagic Sharks Research Articles

Superoxide production, oxidative damage and enzymatic antioxidant defenses in shark skeletal muscle

Pelagic sharks, unlike teleost fish, require constant active swimming to obtain a suitable oxygen (O 2) supply. An increase in O 2 consumption during exercise enhances production of reactive oxygen species (ROS). We hypothesized that shark species that display vigorous exercise, such as Isurus oxyrinchus and Carcharhinus falciformis, have higher ROS production and, in consequence, higher antioxidant enzyme activities in muscle in comparison with species with less active swimming, like Sphyrna zygaena. Superoxide radical (O 2 •−) production, lipid peroxidation levels (TBARS) and the activity of antioxidant enzymes: superoxide dismutase (total, t-SOD; manganese-dependent, Mn-SOD, and copper and zinc-dependent, Cu, Zn-SOD), catalase (CAT), glutathione peroxidase (GPx), glutathione-S-transferase (GST) and glutathione reductase (GR), were measured by spectrophotometric assays in skeletal muscle extracts of three shark species ( C. falciformis, I. oxyrinchus and S. zygaena). Higher O 2 •− production and GPx and GST activities ( p < 0.05) were found in C. falciformis and I. oxyrinchus than in S. zygaena. These results suggest that in antioxidant enzymes (GPx, GST) activity suffices to balance the production of ROS and to maintain lower TBARS levels ( p < 0.05) than in C. falciformis or S. zygaena, contributing to the capacity of I. oxyrinchus to maintain high muscular activity.

Satellite tagging of blue sharks (Prionace glauca) and other pelagic sharks off eastern Australia: depth behaviour, temperature experience and movements

Satellite telemetry was used to study the movements and behaviour of ten blue sharks and one individual each of shortfin mako, thresher and bigeye thresher off eastern Australia. The tracks showed latitudinal movements of up to 1,900 km, but none of the sharks travelled away from the eastern Australian region. Tracking periods did not exceed 177 days. All species showed oscillatory dive behaviour between the surface layers to as deep as 560–1,000 m. Blue sharks spent 35–58% of their time in 300 m. Of these four species, the bigeye thresher spent the least time in the surface layers and the most time at >300 m depth. All four species showed clear diel behaviour generally occupying shallower depths at night than during the day. Blue sharks were mainly in 17.5–20.0°C water, while the thresher sharks showed a more bimodal temperature distribution.

Evaluating ecosystem structure and functioning of the East China Sea Shelf ecosystem, China

As China’s second-largest large marine ecosystem, the East China Sea Shelf has suffered from overfishing, eutrophication, and physical disturbance over the last several decades. A trophic mass-balance model of this ecosystem was developed in order to characterize the structure and functioning of its food web, to identify its keystone species, and to quantify the ecological impacts of fishing that it sustained during the early 2000s. Using a multivariate statistical analysis, we identified 38 functional groups for the trophic model, including fish and invertebrate groups targeted and not targeted by fisheries. Pelagic sharks and rays were identified as the keystone species in the ecosystem. Strong benthic–pelagic coupling was indicated in this ecosystem. In particular, this study highlighted the interdependent relationships that exist among plankton, benthic invertebrates, and detritus. Recent fishing activities were characterized by high exploitation rates for various commercially targeted and non-targeted species, leading to the removal of much of the ecosystem’s fishable production. Overall, our findings give a preliminary explanation of the current problems of eutrophication and fishery depletion and other changes in the East China Sea Shelf, and highlight the need for developing ecosystem-based fisheries management.

Ecological risk assessment of pelagic sharks caught in Atlantic pelagic longline fisheries

An ecological risk assessment (ERA; also known as productivity and susceptibility analysis, PSA) was conducted on eleven species of pelagic elasmobranchs (10 sharks and 1 ray) to assess their vulnerability to pelagic longline fisheries in the Atlantic Ocean. This was a level-3 quantitative assessment consisting of a risk analysis to evaluate the biological productivity of these species and a susceptibility analysis to assess their propensity to capture and mortality in pelagic longline fisheries. The risk analysis estimated productivity (intrinsic rate of increase, r) using a stochastic Leslie matrix approach that incorporated uncertainty in age at maturity, lifespan, age-specific natural mortality and fecundity. Susceptibility to the fishery was calculated as the product of four components, which were also calculated quantitatively: availability of the species to the fleet, encounterability of the gear given the species vertical distribution, gear selectivity and post-capture mortality. Information from observer programs by several ICCAT nations was used to derive fleet-specific susceptibility values. Results indicated that most species of pelagic sharks have low productivities and varying levels of susceptibility to pelagic longline gear. A number of species were grouped near the high-risk area of the productivity-susceptibility plot, particularly the silky (Carcharhinus falciformis), shortfin mako (Isurus oxyrinchus), and bigeye thresher (Alopias superciliosus) sharks. Other species, such as the oceanic whitetip (Carcharhinus longimanus) and longfin mako (Isurus paucus) sharks, are also highly vulnerable. The blue shark (Prionace glauca) has intermediate vulnerability, whereas the smooth hammerhead (Sphyrna zygaena), scalloped hammerhead (Sphyrna lewini), and porbeagle (Lamna nasus) sharks are less vulnerable, and the pelagic stingray (Pteroplatytrygon violacea) and common thresher (Alopias vulpinus) sharks have the lowest vulnerabilities. As a group, pelagic sharks are particularly vulnerable to pelagic longline fisheries mostly as a result of their limited productivity.

Melanomacrophages in three species of free‐ranging sharks from the northwestern Atlantic, the blue shark Prionacae glauca (L.), the shortfin mako, Isurus oxyrhinchus Rafinesque, and the thresher, Alopias vulpinus (Bonnaterre)

The melanomacrophage aggregates or cells (MMC) are commonly used as biomarkers of exposure to pollution in fish, albeit their numbers and morphological characteristics can be influenced not only by environmental toxins but also by a range of physiological parameters and pathological conditions. Accordingly, before we can use MMC as biomarkers in any fish species, their normal, 'background' characteristics have to be established in apparently healthy fish. The knowledge of MMC in sharks is minimal. The aim of this study was to characterize MMC from 51 free-ranging, large pelagic sharks from the northwestern Atlantic, including shortfin mako, Isurus oxyrhinchus, thresher, Alopias vulpinus and blue shark, Prionacae glauca. The rationale of this study was twofold. First, because, sharks have life histories predisposing them to the accumulation of environmental toxins they constitute good sentinel species of the health of the global marine ecosystem. Second, because many shark populations are in decline or threatened by extinction, we need to expand our understanding of their health status in order to develop appropriate protective measures. All sharks were collected by sports fishing gear between June and August in 2007. Their health condition was assessed by necropsy, morphometrics, and by microscopic examination of gonads, livers, spleens and kidneys. Routine, haematoxylin and eosin and/or Pearl's reagent-stained paraffin embedded sections were studied by light microscopy. Our results provide the first data on the morphometric and morphological characteristics of MMC in viscera of apparently healthy free-ranging sharks from the northwestern Atlantic.

Pelagic shark bycatch in the tuna- and swordfish-directed longline fishery off southern Africa

The capture of pelagic sharks as bycatch of the South African pelagic longline fleet targeting tuna Thunnus spp. and swordfish Xiphias gladius was investigated during the period 1998-2005. In all, 26 species were caught, of which six are listed by the International Union for Conservation of Nature (lUCN) as Vulnerable and one (scalloped hammerhead Sphyrna lewini) as Endangered. Blue shark Prionace glauca and short-finned mako Isurus oxyrinchus were the most commonly caught species (69.2% and 17.2% respectively). Generalised linear models explained 70.4% of blue shark and 22.2% of short-finned mako bycatch patterns and showed that vessel name was the most important explanatory variable. Other significant explanatory variables included month, year, area, bathymetry, bait type, moon phase and time of set. South African-flagged, swordfish-directed vessels caught more sharks (11.7 blue sharks and 1.4 short-finned mako sharks per 1 000 hooks) than Asian-flagged, tuna-directed vessels (1.8 blue sharks and 0.9 short-finned mako per 1 000 hooks). The catch per unit effort of blue sharks and short-finned mako sharks started to decrease from 2001 and 2000 respectively. This was accompanied by a decrease in average length for both species over the period 2002-2007. Three techniques for extrapolating total shark mortality were investigated. Simple extrapolation yielded an estimate of 73 500 sharks per year; if vessel flag was taken into account, the estimate decreased to 39 200 sharks per year and to 43 100 sharks if vessel flag and area (5° grid cells) were taken into account.

Sexual segregation of pelagic sharks and the potential threat from fisheries

Large pelagic sharks are declining in abundance in many oceans owing to fisheries exploitation. What is not known however is whether within-species geographical segregation of the sexes exacerbates this as a consequence of differential exploitation by spatially focused fisheries. Here we show striking sexual segregation in the fastest swimming shark, the shortfin mako Isurus oxyrinchus, across the South Pacific Ocean. The novel finding of a sexual 'line in the sea' spans a historical longline-fishing intensity gradient, suggesting that differential exploitation of the sexes is possible, a phenomenon which may underlie changes in the shark populations observed elsewhere.

Age and growth of the shortfin mako (Isurus oxyrinchus) in the south-eastern Pacific off Chile

The shortfin mako, Isurus oxyrinchus, is a large pelagic shark with a widespread global distribution. However, very little is known about most aspects of this species for the south-eastern Pacific. In the present paper, the age and growth parameters of the shortfin mako, caught by Chilean swordfish longline fisheries from 2004 to 2005, are reported. Ages were estimated by counting band-pairs from sections of vertebral centra from 547 individuals, ranging from 76 to 330 cm in total length (TL). Trends in the proportion of opaque edges for all ages combined and grouped into ages 0–6 and 7–26 years indicated that they are formed during summer and showed that annually, one band-pair is formed in the vertebrae of shortfin makos. Modal-progression analysis was used to verify the first three age classes (ages 0–2 years). For both sexes, the oldest estimated age was 25+ years. Von Bertalanffy growth parameters were estimated at L∞ = 325.29 cm TL, K = 0.076 year–1 and t0 = –3.18 years for females and L∞ = 296.60 cm TL, K = 0.087 year–1 and t0 = –3.58 years for males. The results indicated that this species is highly vulnerable to exploitation and, thus, urgent conservation measures are required.

Characterization of the pelagic shark‐fin trade in north‐central Chile by genetic identification and trader surveys

Shark fins have become a highly valued commodity with the major Asian fin‐trade centres supplied from global sources, including Chile. With growing concerns about the resilience of shark populations to heavy fishing pressure, there is a need for better information on shark landings to aid management efforts. In the widespread absence of shark landing records especially by species, monitoring the fin trade has been proposed as a way to assess species exploitation levels. Here, the first species assessment of the Chilean shark‐fin trade was provided. The goals of this study were to (1) determine the species composition and relative species proportion of sharks utilized in the fin trade, (2) determine the relationship between fin trader market names and species and (3) assess trader accuracy in identifying shark fin species based on fin photographs. Fins were analysed from two different fin drying facilities (n = 654) (secaderos) and two fin‐storage warehouses (n = 251). In contrast to official government landing records that only document four species in the landings, molecular species identification of the fins demonstrated that at least 10 pelagic shark species are present in the north‐central Chilean shark fin trade: Alopias superciliosus, Alopias vulpinus, Carcharhinus obscurus, Galeorhinus galeus, Isurus oxyrinchus, Isurus paucus, Lamna nasus, Prionace glauca, Sphyrna lewini, Sphyrna zygaena. The species composition of the fins from the secaderos was P. glauca (83·9%), I. oxyrinchus (13·6%), L. nasus (1·7%) and A. superciliosus (0·2%). There was generally good agreement between market names and single shark species for the trade categories ‘Azulejo’, ‘Tiburon’, ‘Tintorera’, ‘Cola de zorro’ and ‘Martillo’. In contrast, the market category ‘Carcharhinus’ consisted of a mixture of at least five species. The molecular results also identified two species (S. lewini and I. paucus) not previously recorded in Chilean waters. The fin identification survey given to nine regional traders demonstrated that they were highly accurate in recognizing pictures of fins from P. glauca and I. oxyrinchus. The overall strong concordance between market categories and fins from single species and the trader accuracy in survey fin identification suggests that monitoring the Chilean fin trade by market names will provide a reasonably accurate picture of the volume of sharks landed by species.

Functional Dorsoventral Symmetry in Relation to Lift-Based Swimming in the Ocean Sunfish Mola mola

The largest (up to 2 tons) and a globally distributed teleost—the ocean sunfish Mola mola—is commonly regarded as a planktonic fish because of its unusual shape including absence of caudal fin. This common view was recently questioned because the horizontal movements of the ocean sunfish tracked by acoustic telemetry were independent of ocean currents. However, direct information regarding their locomotor performance under natural conditions is still lacking. By using multi-sensor tags, we show that sunfish indeed swam continuously with frequent vertical movements at speeds of 0.4–0.7 m s–1, which is similar to the records of other large fishes such as salmons, marlins, and pelagic sharks. The acceleration data revealed that they stroked their dorsal and anal fins synchronously (dominant frequency, 0.3–0.6 Hz) to generate a lift-based thrust, as penguins do using two symmetrical flippers. Morphological studies of sunfish (mass, 2–959 kg) showed that the dorsal and anal fins had similar external (symmetrical shape and identical area) and internal (identical locomotory muscle mass) features; however, the muscle shape differed markedly. We conclude that ocean sunfish have functional dorsoventral symmetry with regards to the non-homologous dorsal and anal fins that act as a pair of vertical hydrofoils. Although sunfish lack a swimbladder, we found that they are neutrally buoyant independent of depth because of their subcutaneous gelatinous tissue that has low density and is incompressible. Efficient lift-based swimming in conjunction with neutral buoyancy enables sunfish to travel long distances both horizontally and vertically.

Use of shark fin trade data to estimate historic total shark removals in the Atlantic Ocean

In order to address critical gaps arising from limited available data on historic shark catches in the Atlantic Ocean, a method was developed to estimate shark removals using shark fin trade data. A characterization of the global fin trade as of 2000, including number and biomass by shark species, was used as the basis of the methodology. A first step involved scaling Hong Kong trade-derived estimates for 2000 to annual global values for 1980-2006 based on the observed quantity of imports to Hong Kong and an approximation of Hong Kong's share of the global trade in each year. The resulting global fin trade figures for each year were then scaled to Atlantic-specific values using three different factors: (1) area of the Atlantic range relative to the global range of pelagic sharks; (2) Atlantic catches of tunas and billfishes relative to global catches of tunas and billfishes; and (3) Atlantic longline effort relative to global longline effort. The strengths and weaknesses of each scaling factor and the assumptions inherent in the methodology are discussed. These estimates are not intended to replace reliable fisheries dependent catch data compiled by ICCAT from submissions of members and cooperators, but can serve as one of a variety of useful cross-validation tools when historic catch data are missing or uncertain.

You can swim but you can't hide: the global status and conservation of oceanic pelagic sharks and rays

1.Fishing spans all oceans and the impact on ocean predators such as sharks and rays is largely unknown. A lack of data and complicated jurisdictional issues present particular challenges for assessing and conserving high seas biodiversity. It is clear, however, that pelagic sharks and rays of the open ocean are subject to high and often unrestricted levels of mortality from bycatch and targeted fisheries for their meat and valuable fins. 2.These species exhibit a wide range of life-history characteristics, but many have relatively low productivity and consequently relatively high intrinsic vulnerability to over-exploitation. The IUCN — World Conservation Union Red List criteria were used to assess the global status of 21 oceanic pelagic shark and ray species. 3.Three-quarters (16) of these species are classified as Threatened or Near Threatened. Eleven species are globally threatened with higher risk of extinction: the giant devilray is Endangered, ten sharks are Vulnerable and a further five species are Near Threatened. Threat status depends on the interaction between the demographic resilience of the species and intensity of fisheries exploitation. 4.4. Most threatened species, like the shortfin mako shark, have low population increase rates and suffer high fishing mortality throughout their range. Species with a lower risk of extinction have either fast, resilient life histories (e.g. pelagic stingray) or are species with slow, less resilient life histories but subject to fisheries management (e.g. salmon shark). 5.5. Recommendations, including implementing and enforcing finning bans and catch limits, are made to guide effective conservation and management of these sharks and rays. Copyright © 2008 John Wiley & Sons, Ltd.

Sea turtles: A review of some key recent discoveries and remaining questions

This review highlights some recent important developments in our understanding of the physiological, behavioural and molecular ecology of sea turtles as well as their conservation biology. Key commonalities are identified between the biological questions being addressed with respect to sea turtles compared to those with marine mammals (e.g. pinnipeds and whales), birds (e.g. albatrosses) and fish (e.g. tuna, swordfish and pelagic sharks). Key areas highlighted include (1) objectively identifying oceanic hotspots; (2) measuring what animals are actually doing in different parts of their range (e.g. foraging) rather than simply inferring their state from their position and movement; (3) completing energy budgets for species over ecologically relevant timescales (e.g. the period between breeding seasons); (4) using biological information to help drive conservation management; and (5) assessing the impact of climate change.

International fisheries agreement: Review of the International Commission for the Conservation of Atlantic Tunas

To properly manage world fisheries, especially highly migratory species, there are numerous treaties that have been implemented by specialized international fishery commissions. One of the oldest organizations is the International Commission for the Conservation of Atlantic Tunas (ICCAT). The main focus of this review was to summarize the history, roles and responsibilities of ICCAT, and to evaluate ICCAT's management of sharks in general and specifically for shortfin mako (Isurus oxyrinchus) shark. This review found that the Convention was established 40 years ago, but it was not until 2004 that the commission conducted its first population assessment on pelagic sharks.

Dieta de tiburones juveniles Prionace glauca (Linnaeus, 1758) (Carcharhiniformes: Carcharhinidae) en la zona litoral centro-sur de Chile

Feeding studies of pelagic sharks in Chilean waters are rare because of the difficulty in accessing and collecting stomachs. This paper is the first contribution to the knowledge of the diet of juvenile P. glauca (Linnaeus, 1758), caught in the coastal zone of central-southern Chile. We described the diet of 13 juvenile P. glauca using frequency of occurrence, percent by number and weight quantitative indexes. The relationship between prey size and number and predator size was quantified with correlation analysis. Only 77% of stomachs had food contents. One hundred percent of stomachs contained Engraulis ringens and 60% had Dosidicus gigas. The relationship found between predator and prey size reinforces the hypothesis of positive relationships between predator and prey body size

Circle Hooks for Pacific Longliners: Not a Panacea for Marlin and Shark Bycatch, but Part of the Solution

AbstractBlue marlin Makaira nigricans, striped marlin Tetrapturus audax, and pelagic sharks (e.g., blue shark Prionace glauca) are commonly caught as bycatch by longline fisheries in the central North Pacific Ocean. Recently, concern has increased about depletion of these species. Modifications in longline gear may offer one solution. Here, we test the use of circle hooks, rather than the conventional tuna‐style hooks, on longlines using an ecosystem model of the central North Pacific Ocean. The simulations considered span a range of reasonable circle hook catchability and survival rates for released fish. The results suggest that if circle hooks have higher catchability than the currently used tuna‐style hooks, switching to circle hooks depletes marlin biomass by 25–40% and shark biomass by 15–35% over 30 years. However, these depletions do not occur if circle hook catchability is equal to or lower than that of tuna‐style hooks. When the effects of catch‐and‐release requirements for marlins and sharks were also considered, we found that regardless of assumptions about circle hook catchability and survival rates, a combined policy of using circle hooks and releasing sharks and marlins leads to net increases in marlin and shark abundance. The simulations show a trade‐off between the abundance of marlins and sharks and their prey items, yellowfin tuna Thunnus albacares and small blue sharks. There is also evidence of trophic trade‐offs between yellowfin tuna and small blue sharks and their prey, small scombrids (Auxis spp.) and mahi mahi Coryphaena hippurus. The results illustrate the importance of understanding catchability and survival rates for circle hooks compared with those for tuna‐style hooks and encourage further research in this area.

Relative Abundance of Pelagic Sharks in the Western North Atlantic Ocean, Including the Gulf of Mexico and Caribbean Sea

Little information exists on the status of pelagic shark populations in the Atlantic Ocean, especially in the Gulf of Mexico and Caribbean Sea. We derived indices of relative abundance for pelagic sharks based on mandatory logbooks and observer reports from a scientific observer program of the United States (US) pelagic longline fleet. Time series data from the pelagic longline logbook program (1986-2005) and the pelagic longline observer program (1992-2005) were standardized with Generalized Linear Model (GLM) procedures. Declines in relative abundance for the 6 pelagic shark species or genera examined in the logbook data analysis ranged from 43% for mako sharks, Isurus spp., to 88% for blue sharks, Prionace glauca, whereas declines in relative abundance obtained from the observer data analysis were less accentuated than those in the logbook data analysis, with the trend being positive for night sharks, Carcharhinus signatus, and thresher sharks, Alopias spp. There was no significant change in the fork length at capture over the time period considered for blue sharks, shortfin makos Isurus oxyrinchus, or night sharks. The trends obtained must be viewed cautiously given recognized shortcomings, especially of the logbook dataset, and the highly migratory nature of pelagic sharks, which requires a more comprehensive evaluation of trends throughout their range.

Plasma catecholamine levels as indicators of the post-release survivorship of juvenile pelagic sharks caught on experimental drift longlines in the Southern California Bight

Between 1983 and 2004, nearly 12 000 shortf in mako (Isurus oxyrinchus), common thresher (Alopias vulpinus) and blue (Prionace glauca) sharks were tagged in the Southern California Bight; however, only 1.97% of these have been returned. One possible reason for this low return rate could be post-release mortality caused by capture stress from the experimental longline. Plasma catecholamine levels were analysed to evaluate stress levels in longline-captured, rod-and-reel-captured and unstressed docile sharks. The mean catecholamine values determined for the three tag–release species ranged from 6539 to 22 079 pg mL–1. The level of adrenaline found in moribund I. oxyrinchus (94 807 pg mL–1) was much higher than in either P. glauca (46 845 pg mL–1) or A. vulpinus (36 890 pg mL–1). In contrast, blood obtained from sharks that were landed within minutes had lower catecholamine values (P. glauca, 889 and 1347 pg mL–1; I. oxyrinchus, 2960 and 3946 pg mL–1, adrenaline and noradrenaline respectively). Among the nine I. oxyrinchus specimens that were recaptured long after their longline capture and release, the highest adrenaline level measured just before release was 33 352 pg mL–1. Because these mako sharks survived sufficiently long to be recaptured, their time-of-release catecholamine levels provide a conservative estimate of ~80% viability on the longline-captured and released population.

Diel and tidal rhythms in diving behaviour of pelagic sharks identified by signal processing of archival tagging data

Patterns of vertical movement in pelagic predators can be highly complex, reflecting behaviours such as foraging, thermoregulatory excursions and spawning. Here we used fast Fourier analysis to identify periodicity in the vertical movements of 6 basking sharks Cetorhinus maximus from archival tagging data that totalled 595 d. We analysed quantitatively fine-scale vertical movements of basking sharks over seasonal scales (May to February) and detected predominant periodicities related to the vertical movements of the sharks' zooplankton prey. Normal and reverse diel vertical migration (DVM) represented the main periodic dive behaviour, occurring for 11 to 72% of individual track times. A tidal pattern of vertical movement, previously unreported for sharks, was also identified. A possible mechanism for this behaviour appears related to the shark exploiting tidally-induced aggregations of zooplankton prey at depth. The youngest shark tagged showed a markedly different pattern of vertical behaviour. Long-term data sets of swimming depth are becom- ing increasingly available for pelagic predators from pressure-sensitive data loggers. This study demonstrates the utility of signal processing techniques in objectively identifying both expected and unexpected periodicity in these continuous, high-resolution tracks.

Insight into the role of cetaceans in the life cycle of the tetraphyllideans (Platyhelminthes: Cestoda)

Four types of tetraphyllidean larvae infect cetaceans worldwide: two plerocercoids differing in size, ‘small’ (SP) and ‘large’ (LP), and two merocercoids referred to as Phyllobothrium delphini and Monorygma grimaldii. The latter merocercoid larvae parasitize marine mammals exclusively and exhibit a specialised cystic structure. Adult stages are unknown for any of the larvae and thus the role of cetaceans in the life cycle of these species has been a long-standing problem. The SP and LP forms are thought to be earlier stages of P. delphini and M. grimaldii that are presumed to infect large pelagic sharks that feed on cetaceans. A molecular analysis of the D2 variable region of the large subunit ribosomal DNA gene based on several individuals of each larval type collected from three Mediterranean species of cetaceans showed consistent and unique molecular signatures for each type regardless of host species or site of infection. The degree of divergence suggested that LP, P. delphini and M. grimaldii larvae may represent separate species, whereas SP may be conspecific with M. grimaldii. In all host species, individuals of SP accumulated in the gut areas in which the lymphoid tissue was especially developed. We suggest therefore that these larvae use the lymphatic system to migrate to the abdominal peritoneum and mesenteries where they develop into forms recognizable as M. grimaldii. The plerocercoid stage of P. delphini remains unknown. In a partial phylogenetic tree of the Tetraphyllidea, all larvae formed a clade that included a representative of the genus Clistobothrium, some species of which parasitize sharks such as the great white which is known to feed on cetaceans. A bibliographic examination of tetraphyllidean infections in marine mammals indicated that these larvae are acquired mostly offshore. In summary, the evidence suggests that cetaceans play a significant role in the life cycle of these larvae. In addition, it seems clear that cetaceans act as natural intermediate hosts for P. delphini and M. grimaldii, as within these hosts they undergo development from the plerocercoid stage to the merocercoid stage. Because tetraphyllidean species use fish, cephalopods and other marine invertebrates as intermediate hosts, the inclusion of cetaceans in the life cycle would have facilitated their transmission to apex predators such as the large, lamnid sharks. The biological significance of infections of LP in cetaceans is unclear, but infections do not seem to be accidental as such larvae show high prevalence and abundance as well as a high degree of site specificity, particularly in the anal crypts and bile ducts.