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1.
Declines of large sharks and subsequent release of elasmobranch mesopredators (smaller sharks and rays) may pose problems for marine fisheries management as some mesopredators consume exploitable shellfish species. The spotted eagle ray (Aetobatus narinari) is the most abundant inshore elasmobranch in subtropical Bermuda, but its predatory role remains unexamined despite suspected abundance increases and its hypothesized specialization for mollusks. We utilized a combination of acoustic telemetry, benthic invertebrate sampling, gut content analysis and manipulative experiments to assess the impact of spotted eagle rays on Bermudian shellfish resources. Residency and distribution of adult spotted eagle rays was monitored over two consecutive summers in Harrington Sound (HS), an enclosed inshore lagoon that has historically supported multiple recreational and commercial shellfish species. Telemetered rays exhibited variable fidelity (depending on sex) to HS, though generally selected regions that supported relatively high densities of potential mollusk prey. Gut content analysis from rays collected in HS revealed a diet of mainly bivalves and a few gastropods, with calico clam (Macrocallista maculata) representing the most important prey item. Manipulative field and mesocosm experiments with calico clams suggested that rays selected prey patches based on density, though there was no evidence of rays depleting clam patches to extirpation. Overall, spotted eagle rays had modest impacts on local shellfish populations at current population levels, suggesting a reduced role in transmitting cascading effects from apex predator loss. However, due to the strong degree of coupling between rays and multiple protected mollusks in HS, ecosystem-based management that accounts for ray predation should be adopted.  相似文献   

2.
As environmental change persists, understanding resource use patterns is of value to predict the consequences of shifting trophic structures. While many sharks are opportunistic predators, some exhibit prey selectivity, putting them at higher risk compared to species with greater trophic plasticity. In the Gulf of Mexico (GOM), Clupeids and Sciaenids comprise 69% of blacktip shark (Carcharhinus limbatus) diets, which is consequential considering potential responses of these prey groups to disturbance and over harvesting. We assessed if blacktips exhibit selectivity for Clupeids and Sciaenids in the western GOM based on stomach contents from sharks in coastal Texas. Clupeids comprised <2% of diets, while striped mullet (Mugil cephalus) and red drum (Sciaenops ocellatus) comprised >70% of identifiable prey. Ontogenetic shifts from smaller (Clupeids, small Sciaenids) to larger, higher trophic level (Ariidae, Elasmobranchii) prey fits our understanding of foraging among coastal sharks, and suggests our regional understanding of blacktip trophic ecology may be limited by the sizes of sampled sharks. Observed increases in blacktip densities coupled with declines in prey (Mugilids, Sciaenids) is concerning if blacktips have limited diet plasticity. Yet GOM blacktips may be more generalized than previously thought, which is promising for conservation and management.  相似文献   

3.
Synopsis We determined age and growth, size at maturity, and fecundity for cownose rays, Rhinoptera bonasus, collected from the northern Gulf of Mexico. Vertebral age estimates ranged from 0+ to 18+ years for females and 0+ to 16+ years for males. Annual deposition of growth increments was verified with marginal increment analysis. Likelihood ratio tests indicated that the growth of the cownose ray was best described by a combined sexes Gompertz model. Median size at 50% maturity was determined to be 642 mm DW for males and 653 mm DW for females, or 4–5 years of age. Median pup size-at-birth was estimated to be 350 mm DW, with a gestation period of 11–12 months. In all cases, gravid females contained only one pup. Statistically significant differences were detected between growth curves for the Gulf of Mexico and the western Atlantic Ocean. Cownose rays in the Gulf of Mexico had lower estimates of DW and K, and a higher theoretical longevity than their conspecifics in the western Atlantic Ocean. Cownose rays in the Gulf of Mexico also attain maturity at a smaller size and earlier age than their counterparts in the western Atlantic Ocean.  相似文献   

4.
Cownose rays are implicated in the consumption of commercially important shellfish on the U.S. East Coast. We tested this assumption by developing a molecular technique for species identification from cownose ray gut contents. Digestive tracts sampled from 33 rays in Pamlico Sound, NC and Chesapeake Bay, VA contained pieces of partially-digested tissue, well-digested tissue, fluid, and minute shell fragments which made visual identification to the species level nearly impossible. We sequenced the cytochrome oxidase subunit I (COI) for seven locally acquired bivalve species, chosen for their commercial and ecological importance in NC and VA. Sequences were used to design species-specific primers for each bivalve species to amplify polymerase chain reaction (PCR) products. We designed primers such that PCR products were sufficiently different in size to be distinguishable from one another when resolved on an agarose gel, and multiplexing of several species in one reaction was possible. Digestive tract sample testing revealed that cownose rays in Chesapeake Bay ate stout tagelus and soft shell clams. There was no evidence of the rays in the study consuming commercially important oysters, hard clams, and bay scallops. Further sampling over an extended period of time and additional locations is required to confirm these results. Our diagnostic tests could easily be expanded to elucidate the impact of cownose ray predation on prey populations.  相似文献   

5.
Cownose ray, Rhinoptera bonasus, is a common elasmobranch species along the southeast United States coast that recently has received negative attention. These rays can consume considerable amounts of commercial shellfish raising concerns regarding their control and need for effective management. However, limited information is known regarding their population abundance and migration patterns. We addressed the latter by reviewing 25 tagged cownose rays in Chesapeake Bay with pop-up satellite archival tags (PSATs) to study their movement patterns during summer and fall and identify wintering grounds. Eleven tags provided useful data on temperature, depth, light level and/or end locations. The migration tracks were deciphered through geolocation based on light levels, sea surface temperatures and depth constraints. PSAT end locations indicated southern wintering grounds in the coastal waters of central Florida. Female rays migrated out of Chesapeake Bay at the end of September to early October and continued their southerly migration to Florida. Male rays exited the bay in July and migrated northward based on their estimated movement tracks. The male rays appeared to have a second summer feeding ground off the coast of southern New England. In the fall, males migrated south from New England to the same wintering grounds as the females. No diel differences in habitat use were detected; however, males tended to occupy a wider depth and temperature range compared to females. Information on the movement patterns and habitat use for cownose rays will assist in determining more effective recreational and commercial management plans.  相似文献   

6.
The cownose ray, Rhinoptera bonasus, a K-selected species in Chesapeake Bay is subject to an unregulated fishery with no management plan in place. Understanding the reproductive potential for cownose ray is important for successful conservation and management. This study marks the first observation of multiple embryos and right uterus functionality. Eight instances of multiple embryos in cownose rays are reported; two sets of twin live births from captive rays and six separate in utero multiple embryos discovered during necropsy of fishery-dependent and -independent samples. All multiple embryos were in the left uterus. Live births in captivity were confirmed as two sets of twins through direct sequencing of a portion of the mitochondrial DNA and analysis of nuclear microsatellite loci from the newborn pups and putative mothers. In addition, first- and third- quarter term embryos were removed from the right uterus of two female rays during necropsy, marking the first reports of gestation in the right oviduct of cownose rays. The recovery of a three-quarter term albinistic cownose ray embryo through necropsy is also reported.  相似文献   

7.
Assessing the realized effect of dispersal in the genetic makeup of a species has significant evolutionary, ecological, and economical consequences. Here, we investigate the genetic diversity and population differentiation in the aquilopelagic golden cownose ray Rhinoptera steindachneri from the Gulf of California (GC) and the Pacific coast of Baja California (PCBC) using the mitochondrial NADH2 gene. Low levels of genetic diversity were found with only 4 polymerase chain reaction-restriction fragment length polymorphism haplotypes among 76 specimens. Pacific coast organisms were fixed for a unique haplotype not shared with rays from the gulf; 92% of GC rays possessed a single NADH2 haplotype not found in the Pacific. This produced significant differentiation between the GC and the PCBC (Φ(CT) = 0.972, P < 0.001). A pronounced phylogeographic pattern was found in which GC haplotypes were reciprocally monophyletic relative to a very divergent Pacific lineage (d = 10%). Our results indicate that despite high dispersal potential, GC and PCBC golden cownose ray populations are characterized by highly divergent mitochondrial lineages. Although more evidence is needed to corroborate the genetic isolation and systematic status of PCBC and GC golden cownose rays, our results suggest a possible cryptic species in the region.  相似文献   

8.
Abstract: I studied behavior and range patterns of individual bottlenose dolphins during 1984 in the mid-eastern Gulf of California, Mexico. Dolphin sighting rate was significantly higher in areas close to estuary mouths, 0.306 sightings per hour compared with 0.155 sightings per hour in areas distant from estuary mouths. Dolphins used these estuarine areas to feed; 61% of all behavior observed near estuaries was feeding as compared with 23% elsewhere. Traveling comprised 61% of all behavior observed in areas distant from estuary mouths. Estuaries are sites of large concentrations of nutrients which support great numbers of filter-feeding zooplankton and fish. Bottlenose dolphins may specialize on esmarine prey, or they may feed in estuarine areas simply because of the abundance of potential prey that these systems support. In either case, data on relative numbers, distribution patterns, behavior and diet indicate that this is a general trend in habitat use for many coastal populations of this species in the Pacific and Atlantic.
Ranges of a few individuals spanned a minimum of 65 km of coastline, and animals were not permanent residents of a monitored bay. In contrast, dolphins off the coast of Sarasota, Florida, have been reported to be year-round residents with smaller ranges. This difference in degree of site fidelity may be related to habitat differences. The west coast of Florida is dotted with numerous and large estuarine systems which may host permanent prey populations and support resident groups of dolphins. The Gulf of California coastline contains few estuaries; most are small and perhaps support prey resources which are ephemeral, requiring dolphins to range over larger distances in search of food.  相似文献   

9.
10.
A critical issue in understanding trophic connectivity in ecological systems is the lack in quality and quantity of information about feeding habits. In this work, we present a method for integrating a diversity of feeding habits data from published studies to evaluate the impact on indices that describe characteristics of individual taxa and their connectivity. We focus our study on feeding habits of the fishes of the northern Gulf of Mexico and seek to understand the importance of the forage fish Gulf Menhaden (Brevoortia patronus) in predator diets. We created a database of diet studies from the northern Gulf of Mexico that included six diet metrics: frequency of occurrence, wet weight, dry weight, number, volume, index of relative importance, and index of caloric importance. We then used this information to construct a set of traditional networks (all prey and predators were from a single taxonomic level and trophic connections were parameterized with a single diet metric). We also constructed a “robust” network where all taxa were identified to the lowest taxonomic level and trophic connections were parameterized using a resampling approach that included all available information. Linear regression and resampling methods were used to convert data reported in other diet metrics into the frequency of occurrence diet metric. For both traditional and robust networks, we used network indices to describe topological properties. With the robust network, we conducted removal simulations where the forage fish species Gulf Menhaden, and associated Clupeidae representatives, were removed from the network and the feeding effort of the predators was reallocated among their other prey items. We found that network and node-specific indices were sensitive to the choice of taxonomy and diet metric level. In the robust network, predator species with the greatest number of identified prey had the lowest precision in their connections and prey from the Arthropoda phyla had the lowest precision for connections. From the removal and reallocation simulations, we found that Actinopterygii and Arthropoda were the most impacted prey taxa with 1.2% to 4.3% increase in predation and approximately 23 taxa would receive 50% of the reallocated predation. Overall, the resampling methods we present provide a potential means for combining disparate diet data and enables a comprehensive understanding of trophic interactions within an ecosystem.  相似文献   

11.
The flatfish Citharichthys spilopterus Günther 1862 is the most common bothid in the coastal lagoons of the Gulf of Mexico. The objective of the present study was to describe the trophic biology of this species in the tropical coastal lagoon of Tampamachoco, Mexico. For the diet analysis, we used multivariate discriminant analysis and trophic niche breadth. The morphological analysis showed that the features of the bucco-pharyngeal cavity, the large stomach and the short intestine of this flatfish relate to a carnivorous habit. The overall diet showed that this species is a third-order consumer, feeding mainly on fish (52%, mainly gobies) and crustaceans (36%, mostly decapods). Diet and trophic niche breadth showed no significant differences between sexes (P > 0.1). By contrast, there were significant ontogenetic differences in the diet and trophic niche breadth, where larger C. spilopterus were almost entirely piscivorous and showed the narrowest trophic niche. As the flatfish grew in size there was a trend toward the consumption of larger prey. The importance of copepods and peracarids correlated inversely with flatfish size (P < 0.001), but fish prey correlated directly to flatfish size (P < 0.001). Likewise, there was an inverse significant correlation between niche trophic breadth and flatfish size (P < 0.005). There were also significant seasonal differences in the diet (P < 0.02), related to the availability and vulnerability of prey in the lagoon during the dry and rainy seasons. However, the trophic niche showed no significant differences between seasons. Finally, we discuss the advantages of discriminant analysis applied to evaluate differences among diets of fish groups when compared with other bivariate and multivariate techniques.  相似文献   

12.
Benthic bivalves are important links between primary production and consumers, and are essential intermediates in the flow of energy through estuarine systems. However, information on the diet of filter feeding bivalves in estuarine ecosystems is uncertain, as estuarine waters contain particulate matter from a range of sources and as bivalves are opportunistic feeders. We surveyed bivalves at different distances from the creek mouth at the Yangtze estuarine marsh in winter and summer, and analyzed trophic dynamics using stable isotope (SI) and fatty acid (FA) techniques. Different bivalve species had different spatial distributions in the estuary. Glauconome chinensis mainly occurred in marshes near the creek mouth, while Sinonovacula constricta preferred the creek. Differences were found in the diets of different species. S. constricta consumed more diatoms and bacteria than G. chinensis, while G. chinensis assimilated more macrophyte material. FA markers showed that plants contributed the most (38.86 ± 4.25%) to particular organic matter (POM) in summer, while diatoms contributed the most (12.68 ± 1.17%) during winter. Diatoms made the largest contribution to the diet of S. constricta in both summer (24.73 ± 0.44%) and winter (25.51 ± 0.59%), and plants contributed no more than 4%. This inconsistency indicates seasonal changes in food availability and the active feeding habits of the bivalve. Similar FA profiles for S. constricta indicated that the bivalve had a similar diet composition at different sites, while different δ13C results suggested the diet was derived from different carbon sources (C4 plant Spartina alterniflora and C3 plant Phragmites australis and Scirpus mariqueter) at different sites. Species-specific and temporal and/or spatial variability in bivalve feeding may affect their ecological functions in intertidal marshes, which should be considered in the study of food webs and material flows in estuarine ecosystems.  相似文献   

13.
The structurally reinforced jaws of the cownose ray, Rhinoptera bonasus testify to this species' durophagous diet of mollusks, but seem ill-suited to the behaviors necessary for excavating such prey. This study explores this discordance by investigating the prey excavation and capture kinematics of R. bonasus. Based on the basal suction feeding mechanism in this group of fishes, we hypothesized a hydraulic method of excavation. As expected, prey capture kinematics of R. bonasus show marked differences relative to other elasmobranchs, relating to prey excavation and use of the cephalic lobes (modified anterior pectoral fin extensions unique to derived myliobatiform rays). Prey are excavated by repeated opening and closing of the jaws to fluidize surrounding sand. The food item is then enclosed laterally by the depressed cephalic lobes, which transport it toward the mouth for ingestion by inertial suction. Unlike in most sharks, upper jaw protrusion and mandibular depression are simultaneous. During food capture, the ray's spiracle, mouth, and gill slit movements are timed such that water enters only the mouth (e.g., the spiracle closes prior to prey capture and reopens immediately following). Indigestible parts are then hydraulically winnowed from edible prey portions, by mouth movements similar to those used in excavation, and ejected through the mouth. The unique sensory/manipulatory capabilities of the cephalic lobes, as well as the cownose ray's hydraulic excavation/winnowing behaviors and suction feeding, make this species an effective benthic predator, despite its epibenthic lifestyle.  相似文献   

14.
While foraging theory predicts that predatory responses should be determined by the energy content and size of prey, it is becoming increasingly clear that carnivores regulate their intake of specific nutrients. We tested the hypothesis that prey nutrient composition and predator nutritional history affects foraging intensity, consumption, and prey selection by the wolf spider, Pardosa milvina. By altering the rearing environment for fruit flies, Drosophila melanogaster, we produced high quality flies containing more nitrogen and protein and less lipid than low quality fruit flies. In one experiment, we quantified the proportion of flies taken and consumption across a range of densities of either high or low quality flies and, in a second experiment, we determined the prey capture and consumption of spiders that had been maintained on contrasting diets prior to testing. In both cases, the proportion of prey captured declined with increasing prey density, which characterizes the Type II functional response that is typical of wolf spiders. Spiders with similar nutritional histories killed similar numbers of each prey type but consumed more of the low quality prey. Spiders provided high quality prey in the weeks prior to testing killed more prey than those on the low quality diet but there was no effect of prior diet on consumption. In the third experiment, spiders were maintained on contrasting diets for three weeks and then allowed to select from a mixture of high and low quality prey. Interestingly, feeding history affected prey preferences: spiders that had been on a low quality diet showed no preference but those on the high quality diet selected high quality flies from the mixture. Our results suggest that, even when prey size and species identity are controlled, the nutritional experience of the predator as well as the specific content of the prey shapes predator-prey interactions.  相似文献   

15.
We evaluated the impact of marine materials on the diet and abundance of arthropodivorous lizards inhabiting islands and the coast of the Gulf of California. Here, marine materials are brought onto land by seabirds and by tidal action, and both subsidy pathways cause arthropod abundance to increase. We evaluated Uta stansburiana (side-blotched lizard) diets in three habitats defined by having: (1) no marine subsidies available, (2) only seabird-derived subsidies, and (3) only tidal-derived subsidies. Stable isotope data indicated that lizard diets are subsidized indirectly by seabird and tidal activity. For example, in coastal areas we determined that 40% of a lizard's diet contains arthropods that have consumed algae. Such subsidies may explain why we found that lizards in coastal areas occupy a significantly higher trophic position than lizards in unsubsidized areas. We analyzed eight years of survey data on all arthropodivorous lizards to determine if diet subsidies result in increased lizard abundance. We found that lizards were more abundant in coastal areas than they were in inland habitats, and that they were more abundant on islands with seabirds than on islands without seabird populations. This study provides insight into the importance and effect of marine-derived nutrients from multiple sources on vertebrate consumers inhabiting islands and coastal areas.  相似文献   

16.
Ontogenetic variations in shark diet are often qualitatively inferred from dietary analysis and hindered by high levels of unidentified prey or small sample sizes. This study focused on nursery bound lemon sharks (Negaprion brevirostris, n?=?396), enabling some control over the confounding variables of prey choice associated with ontogeny. Nursery bound lemon sharks exhibited weak ontogenetic variation in dietary composition with high levels of dietary overlap. Variation in prey preference of lemon sharks with ontogeny was complex, but revealed a continuous shift from predominantly opportunistic benthic foraging as neonates to more selective piscivory with increasing shark size while in the nursery. Lemon sharks demonstrated a discrete ontogenetic shift in the number of prey consumed and stomach content weight (Kruskal-Wallis tests p?<?0.01), as well as prey size (ANOVA, p?<?0.001). All sizes of sharks exhibited positive size selection of prey (Mann?CWhitney U tests, p?<?0.01). However, the lack of size preference by all but the largest lemon sharks for their major prey (yellowfin mojarra, Gerres cinereus), suggests neonate sharks, while capable of occasionally foraging on large prey, are relatively inept opportunistic foragers. This was evident in high diet breadth, low diversity of consumed prey and lower trophic level than larger sharks. This study represents the first quantitative analysis of ontogenetic variation in prey preference and size selection in sharks, indicating a flexible foraging tactic in lemon sharks and the importance of hunting ability and predator size in prey choice.  相似文献   

17.
《Zoology (Jena, Germany)》2014,117(2):95-103
Electrosensory pore number, distribution, and sensitivity to prey-simulating electric fields have been described for many shark species. Electrosensory systems in batoids have received much less attention. Pore number and distribution have yet to be correlated to differences in sensitivity. However, pore number, pore distribution and sensitivity have been linked to behavior, diet, and morphology and follow species-specific trends. We report here that cownose rays have a greater number of pores than the yellow stingray, most of which are concentrated on the anterior ventral surface for both species. However, yellow stingrays have a broader arrangement of pores on both their dorsal and ventral surfaces than the cownose rays. Yellow stingrays demonstrated a median behavioral sensitivity to weak electric fields of 22 nV cm−1 and are among the most highly sensitive batoids studied to date. Cownose rays are less sensitive than all other elasmobranch species with a median sensitivity of 107 nV cm−1. As reported in previous studies, a higher pore number did not result in greater sensitivity. Cownose rays are benthopelagic schooling rays and may benefit from reduced sensitivity to bioelectric fields when they are surrounded by the bioelectric fields of conspecifics. Yellow stingrays, on the other hand, are typically solitary and bury in the substrate. A greater number of pores on their dorsal surface might improve detection of predators above them. Also, increased sensitivity and a broader distribution of pores may be beneficial as small prey items move past a buried ray.  相似文献   

18.
Diet analysis allows exploring how coastal dolphins interact with the environment and their role in the marine food webs. We studied the diet and feeding ecology of the Guiana dolphin, Sotalia guianensis, through analysis of stomach content from 42 animals stranded on the eastern coast of Brazil. A total of 1,336 semidigested prey items (fish, otoliths, cephalopod beaks, and crustaceans) were identified. Teleost fish comprised the most frequent food item (92% of the total), followed by cephalopods, and crustaceans. Prey belonged to 34 taxa and richness in individual stomachs varied from 1 to 15 prey taxa. Prey were generally small, but showed a significant trend to increase in size with dolphin length. The main prey of Guiana dolphins were demersal, estuarine, and sound-making fish such as catfish and sciaenids. No sex-related differences in diet were found. Diet composition varied seasonally and occurrence of prey items was coherent with breeding or high abundance periods of some fish species and squids. Our study shows the importance of demersal prey from estuarine and soft-sediment habitats to Guiana dolphin in the Abrolhos Bank and reveals that feeding habits are generalist and opportunistic, with diet reflecting the seasonal abundance and availability of prey.  相似文献   

19.
Optimal foraging theory predicts less diverse predator diets with a greater availability of preferred prey. This narrow diet niche should then be dominated by preferred prey, with implications for predator–prey dynamics and prey population ecology. We investigated lion (Panthera leo) diets in Hluhluwe–iMfolozi Park (HiP), South Africa, to assess whether lions in a site with a high density of preferred prey (prey species weighing 92–632 kg as estimated from a published meta-analysis) have a narrow diet, consisting primarily of preferred prey. HiP is a useful study site to investigate this prediction because it is a productive landscape (with a high density of prey) where lion-preferred prey constitutes up to 33% of the prey available to lions. Furthermore, to investigate whether lions in HiP exhibit sex-specific diets as documented in other southern African populations, we estimated male and female lion diets separately. We were specifically interested in testing whether traditional approaches of estimating lion diets at the population level mask sex-specific predation patterns, with possible implications for management of lions in small to medium-sized fenced reserves. Lions in HiP preferred larger prey species (63–684 kg) and had diets with a larger proportion of preferred prey than reported in an African-wide meta-analysis. However, despite the high density of preferred prey species, 36% of lion diets still consisted of typically non-preferred species such as nyala (Tragelaphus angasii). This finding suggests that lions in HiP maintain a degree of opportunism even when preferred prey are abundant. Therefore, abundant, non-preferred prey are likely to be an important resource for lion populations. Sex-specific differences in lion diets were evident in HiP, suggesting that estimation of lion resource use and carrying capacity should consider opportunistic hunting and sex-specific differences in lion diets.  相似文献   

20.
Quantifying the trophic role of sharks in coastal ecosystems is crucial for the construction of accurate ecosystem models. This is particularly important for wide-ranging species like the Atlantic sharpnose shark (Rhizoprionodon terraenovae), ubiquitous across the northern Gulf of Mexico. We used gut content and stable isotope analyses to determine if differences in abundance of Atlantic sharpnose sharks in the waters around Mobile Bay, Alabama translated into differences in dietary sources or trophic position among sharks sampled east and west relative to the mouth of the bay. Gut content analysis suggested that Atlantic sharpnose sharks eat primarily teleost fishes (%IRI?>?90% across size classes), and both stomach content and stable isotope analyses highlighted an ontogenetic shift in diet. Nitrogen stable isotope data from liver and muscle tissues indicated regional shifts in trophic position for Atlantic sharpnose sharks. The mixing model SIAR (stable isotope analysis in R) v.4.0.2 was used to suggest possible contributions from likely prey items for Atlantic sharpnose sharks sampled east and west of Mobile Bay. Portunid crabs and shrimp made higher contributions to the diet of Atlantic sharpnose sharks in the western region, compared to higher and more variable contributions from fish like croaker (Micropogonias undulatus) and hardhead catfish (Arius felis) in the eastern region. Our results suggest trophic plasticity in Atlantic sharpnose sharks, findings that emphasize the importance of examining regional variation in trophic position when constructing coastal foodweb models.  相似文献   

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