Spawning ofConger oceanicus andConger triporiceps (Congridae) in the Sargasso Sea and subsequent distribution of leptocephali

Synopsis Distribution of leptocephali ofConger in the Western North Atlantic Ocean was studied using specimens from our collections, specimens from other collections, and various existing collection records. The presence of leptocephali ofConger oceanicus andConger triporiceps < 30 mm long over deep water in the southwestern Sargasso Sea in autumn and winter implies a protracted spawning period there. The subtropical convergence zone, meandering east-west across the Sargasso Sea, is probably the northern limit of spawning of both species. Spawning may also occur close to the Bahamas and Antilles.C. triporiceps may spawn also in the Caribbean Sea judging by the capture of small leptocephali in the western Caribbean and of the more southerly continental distribution of its juveniles. The claim of Johannes Schmidt in 1931 that the EuropeanC. conger spawns across the North Atlantic into the western Sargasso Sea is probably incorrect, because leptocephali ofConger are rare in the eastern Sargasso Sea and becauseC. triporiceps, with myomere numbers overlapping those ofC. conger, was recently described in the western North Atlantic. With increasing size, leptocephali ofC. oceanicus and a portion ofC. triporiceps spread westward and northward in the Florida Current and Gulf Stream, but larger leptocephali especially ofC. triporiceps are found also in the Caribbean and Gulf of Mexico. Spawning ofC. oceanicus in the Sargasso Sea indicates that adults cross the Florida Current-Gulf Stream, and successful leptocephali cross the current in the opposite direction to colonize juvenile habitat on the continental shelf, a migratory pattern similar to that of the American eelAnguilla rostrata (Anguillidae).     

The Distribution and Ecology of Ariosoma Balearicum (Congridae) Leptocephali in the Western North Atlantic

A total of 4589 leptocephali of the congrid eel, Ariosoma balearicum, were examined from 17 cruises in the western North Atlantic Ocean. Myomere counts made on 915 of these indicated there were two ranges of number of myomeres that appear to be associated with separate spawning populations. Those with the higher range (high count: 128–137) were consistently 70–100mm in length in the Sargasso Sea from February to April and 20–80mm in length in the northern Sargasso Sea and Gulf Stream from September to October. Those with the lower range (low count: 120–130) were rare in the northern and eastern Sargasso Sea where they had consistently greater lengths than high count leptocephali and were most abundant in the Florida Current and Providence Channel. The geographic distributions of size and myomere ranges in relation to hydrography provide strong support for the hypothesis that high count eels found along the South Atlantic Bight (SAB) migrate across the Florida Current to spawn in the northwest Sargasso Sea. This migratory pattern is similar to those of Anguilla rostrata and Conger oceanicus, which use the southern Sargasso Sea for development as larvae. However, the distribution of high count leptocephali suggests that they use the entire Sargasso Sea gyre as a development area as larvae before crossing the Florida Current and recruiting to the SAB. The low count eels inhabiting the Bahamas appear to spawn near the banks and their abundance in the Providence Channel and southwest Sargasso Sea suggests most are retained close to the Bahamas. These two distinct styles of spawning, distribution and recruitment of larvae are hypothesized to be related to the different hydrographic regimes of the two juvenile habitats and the resulting constraints on growth and recruitment of larvae. Vertebral and myomere counts reported from other areas suggest there are distinct populations in other regions of the North Atlantic Ocean.     

Entry of migrating American eel leptocephali into the Gulf Stream system

Abundance, size and depth distributions of American eel(Anguilla rostrata) leptocephali collected in four transects across the Florida Current between the Straits of Florida and Cape Hatteras, North Carolina, were examined in order to assess the relative importance of two migration routes into the Gulf Stream system. A nine-fold increase in the abundance and an increase in the mean length of leptocephali collected in the Florida Current north of the Straits of Florida indicate that most leptocephali enter the Gulf Stream system directly from the Sargasso Sea rather than by a more southern route through the Bahama Islands. Leptocephali were concentrated in the upper 140 m at night and upper 350 m during the day. Wide vertical ranges in daytime collections precluded more refined estimation of vertical distribution. The collection of eight small leptocephali (11 to 17 mm total length) in the western Caribbean Sea is discussed in relation to the origin of leptocephali taken in the Straits of Florida.     

A century of research on the larval distributions of the Atlantic eels: a re‐examination of the data

The spawning areas of the Atlantic freshwater eels were discovered about a century ago by the Danish scientist Johannes Schmidt who after years of searching found newly hatched larvae of the European eel, Anguilla anguilla, and the American eel, Anguilla rostrata, in the southern Sargasso Sea. The discovery showed that anguillid eels migrate thousands of kilometers to offshore spawning areas for reproduction, and that their larvae, called leptocephali, are transported equally long distances by ocean currents to their continental recruitment areas. The spawning sites were found to be related to oceanographic conditions several decades later by German and American surveys from 1979 to 1989 and by a Danish survey in 2007 and a German survey in 2011. All these later surveys showed that spawning occurred within a restricted latitudinal range, between temperature fronts within the Subtropical Convergence Zone of the Sargasso Sea. New data and re‐examinations of Schmidt's data confirmed his original conclusions about the two species having some overlap in spawning areas. Although there have been additional collections of leptocephali in various parts of the North Atlantic, and both otolith research and transport modelling studies have subsequently been carried out, there is still a range of unresolved questions about the routes of larval transport and durations of migration. This paper reviews the history and basic findings of surveys for anguillid leptocephali in the North Atlantic and analyses a new comprehensive database that includes 22612 A. anguilla and 9634 A. rostrata leptocephali, which provides a detailed view of the spatial and temporal distributions and size of the larvae across the Atlantic basin and in the Mediterranean Sea. The differences in distributions, maximum sizes, and growth rates of the two species of larvae are likely linked to the contrasting migration distances to their recruitment areas on each side of the basin. Anguilla rostrata leptocephali originate from a more western spawning area, grow faster, and metamorphose at smaller sizes of <70 mm than the larvae of A. anguilla, which mostly are spawned further east and can reach sizes of almost 90 mm. The larvae of A. rostrata spread west and northwest from the spawning area as they grow larger, with some being present in the western Caribbean and eastern Gulf of Mexico. Larvae of A. anguilla appear to be able to reach Europe by entering the Gulf Stream system or by being entrained into frontal countercurrents that transport them directly northeastward. The larval duration of A. anguilla is suggested to be quite variable, but gaps in sampling effort prevent firm conclusions. Although knowledge about larval behaviour is lacking, some influences of directional swimming are implicated by the temporal distributions of the largest larvae. Ocean–atmosphere changes have been hypothesized to affect the survival of the larvae and cause reduced recruitment, so even after about a century following the discovery of their spawning areas, mysteries still remain about the marine life histories of the Atlantic eels.     

Taxonomic revision,ecology and endangerment categorization of the Andean catfish Astroblepus ubidiai (Teleostei: Astroblepidae)

The European eel (Anguilla anguilla Linnaeus 1758) is a species typical for waters of Western Europe. Thanks to early expeditions on the Atlantic Ocean by the Danish biologist Johannes Schmidt who found small (<10mm) leptocephali larvae in the Sargasso Sea about 100 years ago, we have now a strong indication where the spawning site for this species is located. The American eel (Anguilla rostrata, LeSueur) also spawns in the Sargasso Sea. The spawning time and location of both species have been supported and refined in recent analyses of the available historical data. Subsequent ichthyoplankton surveys conducted by McCleave (USA) and Tesch (Germany) in the 1980s indicated an increase in the number of leptocephali <10 mm , confirming and refining the Sargasso Sea theory of Johannes Schmidt. Distinctions between the European and American eel are based on morphological characteristics (number of vertebrae) as well as molecular markers (allozymes, mitochondrial DNA and anonymous genomic-DNA. Although recognised as two distinct species, it remains unclear which mechanisms play a role in species separation during larval drift, and what orientation mechanism eels use during migration in the open sea. The current status of knowledge on these issues will be presented. The hypothesis that all European eel migrate to the Sargasso Sea for reproduction and comprise a single randomly mating population, the so called panmixia theory, was until recently broadly accepted. However, based on field observations, morphological parameters and molecular studies there are some indications that Schmidt’s claim of complete homogeneity of the European eel population and a unique spawning location may be an overstatement. Recent molecular work on European eel indicated a genetic mosaic consisting of several isolated groups, leading to a rejection of the panmixia theory. Nevertheless, the latest extensive genetic survey indicated that the geographical component of genetic structure lacked temporal stability, emphasising the need for temporal replication in the study of highly vagile marine species. Induced spawning of hormone treated eels in the aquarium was collective and simultaneous. In this work for the first time group spawning behaviour has ever been observed and recorded in eels. Studies in swim-tunnels indicate that eels can swim four to six times more efficiently than non-anguilliform fish such as trout. After a laboratory swim trial of eels over 5,500 km, the body composition did not change and fat, protein and carbohydrate were used in the same proportion. This study demonstrated for the first time that European eel are physiologically able of reaching the Sargasso Sea without feeding. Based on catches of newly hatched larvae, temperature preference tests and telemetry tracking of mature hormone treated animals, it can be hypothesised that spawning in the Sargasso Sea is collective and simultaneous, while presumably taking place in the upper 200 m of the ocean. Successful satellite tracking of longfin female eels in New Zealand has been performed to monitor migration pathways. Implementation of this new technology is possible in this species because it is three times larger than the European eel. In the future, miniaturisation of tagging technology may allow European eels to be tracked in time by satellite. The most interesting potential contribution of telemetry tracking of silver eels is additional knowledge about migration routes, rates, and depths. In combination with catches of larvae in the Sargasso Sea, it may elucidate the precise spawning locations of different eel species or groups. Only then, we will be able to define sustainable management issues by integrating this novel knowledge into spawners escapement and juvenile fishing quota.     

Status of the Nemertea as prey in marine ecosystems

Nemerteans are predators of a wide variety of animals, but little is known of their role as prey for other animals. The presence of toxins in the tissues and secretions of these worms has led to the assumption usually suggested that they are ingested only rarely. However, analysis of a Food Habits Data Base from the United States National Marine Fisheries Service, compiled for fishes collected in the Atlantic Ocean from the Canadian border to Cape Hatteras, North Carolina (1973–1990), showed that nemerteans were recovered from the stomachs of 27 species of fishes in 14 families. They were found in 223 of 26 642 (0.84%) fish stomachs examined in the laboratory, but only in 0.09% of 58 812 fish examined in the field. Among species in the former category, for which 1000 were examined, the winter flounder, Pseudopleuronectes americanus(Walbaum), and the yellowtail flounder, Limanda ferruginea (Storer), had the greatest frequency of nemerteans, 71 of 1545 (4.6%) and 33 of 1045 (3.2%), respectively. These nemerteans were identified as Nemertea, Cerebratulus sp. or Micrura sp., but it is likely that they were all of a Cerebratulus-like type. Nemerteans have also been recorded from the guts of eight additional species (including four additional families) of fishes collected from the western Atlantic Ocean, Gulf of Mexico, eastern Pacific Ocean (Washington, Alaska), North Sea and Indian Ocean off South Africa. The black-bellied plover, Pluvialis squatarola(L.), semipalmated plover, Charadrius semipalmatus Bonapart and the herring gull, Larus argentatus Pontoppidan are the only three species known to feed on nemerteans (Cerebratulus lacteus (Leidy) and Paranemertes peregrina Coe by the black-bellied plover and C. lacteus by the other species). Several species of nemerteans are known to ingest other nemerteans, and several arthropods, a squid, and a few other invertebrates also feed on these worms. On the other hand, careful laboratory studies have shown that some members of the Palaeonemertea, Heteronemertea and Hoplonemertea, when fed to various species of fishes, crustaceans (Astacidea, Anomura, Brachyura and Amphipoda) and a polychaete, were rejected, usually violently. There is a history of using large nemerteans, e.g. Cerebratulus lacteus and Polybrachiorhynchus dayi, as bait by sport fishermen in the United States and South Africa. The incongruity of successfuly using these toxic animals to catch fish is discussed in relation to conclusions on the importance of nemerteans as prey in the marine environment.     

Morphology and taxonomic significance of the otoliths of some bathypelagic Anguilloidei and Saccopharyngoidei from the Sargasso Sea

The sagittal otoliths of seven anguilliform species belonging to the families Nemichthyidae, Serrivomeridae and Eurypharyngidae are described and illustrated, and a key for their identification is provided. Although the leptocephali of the various fish species treated here have similar otoliths, some species develop specific otolith characteristics during the adult stage. The shape as well as the ratio of fish length to otolith length ratio in adults may serve as a taxonomic aid. Since the sagittae of these fishes can be used for specific identification, the digested remains of prey in the stomachs and guts of predators can be identified.     

Limited contribution of predation by zooplanktivorous cyprinids to 0+ fish mortality

Predation by zooplanktivorous cyprinids was hypothesized as substantially contributing to larval fish mortality. To test this assumption, potential predators were sampled by electric fishing at sites and times of high fish larvae densities in the lower Oder River, Germany, in 2002 and 2003. At ten sites with high prey fish densities, all fishes ≥1+ and 0+ pike – all considered potentially predating on fish larvae – were captured, anaesthetised, and fixed. In the laboratory, 982 guts, respectively, stomachs were dissected and checked for fish larvae and larvae remains. Predation on 0+ fish was detected in the primary piscivorous fishes Esox lucius L. and Aspius aspius (L.), and facultative piscivorous Perca fluviatilis L., whilst none of the other nine species examined consumed fish. These findings implicate that predation by planktivorous cyprinids did not explain the typically strong decline in larval fish densities in the first weeks after hatching.     

Vertical distributions, diel and ontogenetic vertical migrations and net avoidance of leptocephali of Anguilla and other common species in the Sargasso Sea

Day- and night-time vertical distributions and their ontogeneticchanges in Anguilla leptocephali and other common species ofleptocephali were determined and compared during five cruisesin the Sargasso Sea using an opening - closing 2-m ring netto sample discrete depth strata between 0 m and 350 m deep.No difference in vertical distribution was ever found betweenAnguilla rostrata (American eel) and A. anguilla (European eel).Anguilla leptocephali <5 mm long did not exhibit a diel verticalmigration, as they were distributed between 50 m and 300 m bothby day and by night. The vertical distribution of these smalllepto-cephali is probably roughly representative of the depthdistribution of adult spawning. Anguilla     

Feeding habits of anglerfish,Lophius budegassa (Spinola, 1807) in the Adriatic Sea,north‐eastern Mediterranean

Stomach contents of 283 anglerfish (Lophius budegassa), collected in the Adriatic Sea (north‐eastern Mediterranean) from experimental trawl surveys in 2005 and 2006, were examined to assess their diet as influenced by fish size, sex and water depth. Changes in food quantities in the stomach were observed, the percentage of empty stomachs decreasing with increasing body size and depth. No differences were found between male and female diets. The anglerfish is essentially an ‘opportunistic feeder’ that consumes mainly fishes, crustaceans and molluscs. Fishes were the primary food consumed by all size classes; the favourite prey was Merluccius merluccius (TL >150 mm for medium and large anglerfish) and Gaidropsarus biscayensis (TL <150 mm for smaller anglerfish). However, the proportion of fish species and molluscs changed with the body size of the anglerfish. The trophic level of L. budegassa estimated in the study area reached a mean value of 4.38, confirming that the species was a carnivore with a preference for large decapodes, cephalopods and fish.     

A non‐destructive sampling protocol for field studies of seed dispersal by fishes

This paper presents a standardized protocol for the non‐lethal capture of fishes, sampling of stomach contents and quantification of seed dispersal efficiency by frugivorous fishes. Neotropical pacu Piaractus mesopotamicus individuals were collected with fruit‐baited hooks. The diets of 110 fish were sampled using a lavage method, which retrieved >90% of stomach contents of both juveniles and adults and allowed individuals to recover within 5 min of treatment. The proportional volume of six food categories was similar for stomachs and whole digestive tracts retrieved by dissection. Fruit pulp was proportionally lower in the stomach. The abundance and species richness of intact seeds increased with fish size independent of whether only stomachs or whole digestive tracts were analysed. The analysis of stomach contents accounted for 62·5% of the total species richness of seeds dispersed by P. mesopotamicus and 96% of common seeds (seed species retrieved from more than one fish). Germination trials revealed that seed viability was similar for seeds collected from the stomach via lavage and seeds that passed through the entire digestive tract. Therefore, stomach contents provide an unbiased representation of the dietary patterns and seed dispersal of frugivorous fishes.     

Diet and feeding strategy of blackmouth catshark Galeus melastomus

Diet and feeding strategy of the blackmouth catshark Galeus melastomus in the deep waters of the eastern Ionian Sea were investigated. Sampling was carried out using experimental bottom longline fishing at depths ranging from 300 to 855 m in summer and autumn 2010. Diet variability with fish size, season, area, sex and depth zone was tested and only season was found to significantly affect the diet of the species. Of the 870 stomachs examined, only 1·4% were empty. Cumulative prey curves showed that the sample sizes were adequate to describe the main prey items of the diet for both seasons. Prey identified belonged primarily to three major groups: fishes, cephalopods and dendrobranchiatan and caridean shrimps. In autumn, the above three major groups were found as principal prey, whereas in summer cephalopods followed by fishes were the principal prey and shrimps were found as secondary prey. SIMPER analysis indicated high dissimilarity between seasons and highlighted that Sepiolidae, Myctophidae, fishes, cephalopods, shrimps and other crustaceans contributed to seasonal differences in the diet of G. melastomus. The prey diversity index was higher in autumn than in summer. A high dietary overlap was observed between the two seasons. Galeus melastomus behaved as an opportunistic predator with a variety of unimportant prey in its diet. Feeding strategy indicated that G. melastomus holds a generalist niche with a considerable specialization at the individual level. This strategy seems to be an adaptation to a food‐scarce environment, as typified in deep‐water habitats.     

STOMACH CONTENTS OF FALSE KILLER WHALES (PSEUDORCA CRASSIDENS) STRANDED ON THE COASTS OF THE STRAIT OF MAGELLAN,TIERRA DEL FUEGO

We examined the stomachs of 25 false killer whales collected from a mass stranding of 181 animals along both coasts of the Strait of Magellan, Chile, in March 1989. The 21 stomachs (nine males and 12 females), with food remains contained 11 prey species (nine cephalopods and two fishes) with a total of 442 individuals. Except for one case, food remains were meager, indicating that the animals had not eaten for some time or through stress had vomited on the beach. Eleven of the 21 animals had mud (often with squid beaks) in the esophagus and first stomach. The prey were identified employing squid beaks, fish otoliths and bones, and their wet weights were estimated using regressions between hard parts and known weights of species. The most important prey were the oceanic and neritic-oceanic squids, Martialia byadesi and Illex argentinus, followed by the neritic fish, Macruronus magellanicus. Of less importance were the oceanic squid, Todaroes fillipovae, the oceanic and epipelagic octopus, Ocytboe sp., and the oceanic squid, Moroteuthis ingens. The rest of the prey were poorly represented and included four oceanic squids and one neritic fish. The prey species of these animals were subantarctic, with two antarctic species, abundant over the Patagonian shelf and adjacent oceanic waters around Tierra del Fuego.     

Pleuragramma antarcticum (Pisces,Nototheniidae) as food for other fishes in McMurdo Sound,Antarctica

Summary Nine spcies of notothenioid fishes were captured near the southern limit of their range in ice covered McMurdo Sound, Antarctica. Stomach contents were examined using an occurrence method. Fishes were present in the diets of 8 of 9 species;Dissostichus mawsoni andGymnodraco acuticeps were predominantly piscivorous.Pleuragramma antarcticum was the most common prey fish consumed, being present in stomachs of 4 of 8 species.Pleuragramma was also partially piscivorous (22%) as well as cannibalistic (13%). As evidenced by their presence in the stomach contents of other fishes, all life history stages from 20 mm SL postlarvae to 160 mm SL adultPleuragramma were represented under the ice of the Sound.Pleuragramma was also a major food item for Weddell seals, birds and possibly invertebrates. As a widely distributed species in the pelagic zone,Pleuragramma may be an ecological substitute for euphausiids in the food web of the Sound.     

Biometric studies on premetamorphic eel larvae ofAnguilla anguilla (Anguilliformes: Anguillidae) in comparison with younger developmental stages of the species

387 premetamorphic leptocephali of the familyAnguillidae caught off the west coast of Europe were examined taxonomically and compared with the youngest developmental stages of larvae ofAnguilla anguilla caught in the Sargasso Sea 1979 (Schoth, 1982). The total number of myomeres and the number of myomeres up to the third, opistonephritic blood vessel present features of this species which are significantly different from those of the larvae ofAnguilla rostrata and do not change during the whole larval phase. A combination of these two biometric features enables an infallible species identification of the AtlanticAnguilla larvae at all developmental stages. The number of predorsal and preanal myomeres, the preanal length and features of the head cannot be used for a distinction ofAnguilla larvae. One larva, 68.7 mm long, with 107 myomeres, and 44 myomeres to the opistonephritic blood vessel represents the hitherto southernmost record of anA. rostrata larva in the eastern North Atlantic.     

Comparison of electrophoretic and meristic characters of 0-group eel larvae from the Sargasso Sea

Marked differences between continental samples of American and European eels have been detected electrophoretically in allele frequencies at the MDH-2 locus. Starch gel electrophoresis carried out on board F. R. V. Anton Dohrn during the eel expedition to the Sargasso Sea in 1979 revealed a similar clear-cut genetic difference in a sample of 0-groupAnguilla leptocephali, thus confirming the classical theory of Schmidt (1932). The MDH-2 genotypes provide an additional diagnostic character for the distinction between youngA. anguilla andA. rostrata leptocephali. Species identification by biochemical genetic characters did not correspond with that by meristic characters (myomere numbers) in ca. 13 % of the specimens studied; this discrepancy mainly concerns leptocephali of theA. anguilla genotype. The results obtained are critically discussed.     

Diet of Chelidonichthys obscurus and Chelidonichthys lastoviza (Pisces: Triglidae) from the Gulf of Gabes (Tunisia)

The most common gurnard species in the Gulf of Gabes (Tunisia) are Chelidonichthys lastoviza and C. obscurus. These two species were collected by trawl nets between February 2000 and July 2002 at depths ranging from 80 to 150 meters. In the laboratory the stomach contents were examined for diet assessment and to determine diet variation in relation to fish size, season and sex. The importance of different prey items was analysed utilising several feeding indexes, while diet overlap was tested using Schoener's index and the null model test. The two species showed similar diets, with the same qualitative and quantitative composition. Basic food consisted of crustaceans, mostly mysids and decapods. For Chelidonichthys lastoviza:Gastrosaccus sp. (F = 48.57%), Anchialina agilis (F = 30.57%) and Sicyonia carinata (F = 21.71%); for Chelidonichthys obscurus:Gastrosaccus sp. (F = 80.95%), A. agilis (F = 55.41%) and Pontocaris lacazei (F = 15.15%). A change in food composition based on fish size was observed in both species: mysids more frequently in the diet of small specimens, and decapods in the largest fishes. Seasonal variation in their diet was also recorded. The two fishes showed a significantly higher number of empty stomachs in late autumn and winter. Regardless of species and size class, a biologically significant diet overlap was observed between C. lastoviza and C. obscurus. This result may indicate a lack of competition among triglid species size‐class combinations or strong competition that has not yet led to resource portioning.     

Hybridization between Atlantic Salmon Salmo salar L. and Brown Trout S. trutta L. upon Artificial Propagation

Samples of Salmo salar and S. trutta were examined in 12 Russian fish hatcheries. With protein markers, hybrids of the two species were found in three hatcheries of the Baltic Sea basin. Some fishes had a phenotype intermediate between theS. salar and S. trutta phenotypes by morphological traits, but did not differ genetically from one of the parental species. Possible consequences of hybridization and ways to prevent it are discussed.     

Piscivory by the mummichog (Fundulus heteroclitus): Evidence from the laboratory and salt marshes

While piscivory is common in many fishes, there are few accounts for the fundulid, Fundulus heteroclitus (mummichog). We suspected that this species might be involved in several forms of piscivory including predation, cannibalism and scavenging. To evaluate these possibilities we conducted several laboratory experiments and field observations in its primary habitat, salt marshes. We found that digestion of larvae and small juveniles of F. heteroclitus was fast (< 1 h) and this makes detection of small fish prey difficult for any form of piscivory. In addition, laboratory experiments (one-on-one encounters, feeding on dead prey) indicated that both cannibalism and scavenging of conspecific prey were possible. Field observations (n = 2449 stomachs from fish 12-106 mm TL over four years at two salt marshes) also suggested that piscivory occurred frequently (4.3-24.7% of stomachs examined) based on the detection of numerous fish hard parts, especially pharyngeal jaws. These structures allowed us to determine that most prey were F. heteroclitus and to back-calculate the size of the prey and thus the piscivore/prey size relationship. As a result, it appears that both cannibalism (0.2-9.1% of stomachs) and scavenging (0.5-9.9%) are common feeding modes of F. heteroclitus in salt marshes and, subsequently, modes of energy transfer for salt marshes in general. We suggest that future studies of fish feeding consider that cannibalism and scavenging may be frequent possibilities when fish remains are detected in stomach contents.