Migration and habitat use of the tropical eels Anguilla marmorata and A. bicolor pacifica in Vietnam

The patterns of use of marine and freshwater habitats by the tropical anguillid eels Anguilla marmorata and A. bicolor pacifica were examined by analysing the otolith strontium (Sr) and calcium (Ca) concentrations of yellow (immature) and silver (mature) stage eels collected in Vietnamese waters. In A. marmorata, the change in the Sr:Ca ratios outside the high Sr:Ca core was generally divided into three patterns: (1) typical catadromous life history pattern; (2) constant residence in brackish water; and (3) habitat shifting between sea and brackish waters with no freshwater life. In A. bicolor pacifica, no eels had a general life history as freshwater residents. The eels were also divided into three patterns: (1) constant residence in sea water; (2) constantly living in brackish water; and (3) habitat shifting from brackish to sea water with no freshwater residence. The mean Sr:Ca ratio value after recruitment to coastal waters ranged from 1.73 to 5.67 × 10^?3 (mean 3.2 × 10^?3) in A. marmorata and from 2.53 to 6.32 × 10^?3 (mean 4.3 × 10^?3) in A. bicolor pacifica. The wide range of otolith Sr:Ca ratios in both species indicated that the habitat use of these tropical eels was facultative among fresh, brackish, and marine waters during their growth phases after recruitment to coastal areas. Tropical eel species may have the same behavioural plasticity as temperate anguillid species regarding whether to enter freshwater or to remain in estuarine and marine environments.     

Otolith Sr:Ca and Ba:Ca may give inconsistent indications of estuarine habitat use for American eels (<Emphasis Type="Italic">Anguilla rostrata</Emphasis>)

Temporal patterns in otolith Sr:Ca and Ba:Ca ratio values of American eels Anguilla rostrata from two sites in western Newfoundland gave insight into the use of freshwater and saline habitats. Mean Sr:Ca and Ba:Ca values at the core zone did not differ between sites, indicative of a common oceanic origin. At the otolith edge, representing continental life, both Sr:Ca and Ba:Ca values varied between sites consistent with ambient element:Ca ratio values and salinity, with typically higher Sr:Ca and lower Ba:Ca values in saline than in fresh waters. Most eels (73%) from Muddy Hole, an estuarine site, were evaluated as estuarine residents while most (70%) eels from Castors River, a freshwater site, were evaluated as freshwater residents, with the remaining eels from each site evaluated as inter-habitat migrants. An otolith element:Ca critical value appropriate for distinguishing between fresh and saline water residence is fundamental for estimating the proportion of eel residence in freshwater and their subsequent classification into habitat residence groups. Such classification is moderately robust to the critical value selected. For inter-habitat migrants, moderate otolith Sr:Ca values between the elver check and otolith edge suggestive of estuarine residence may coincide with Ba:Ca values suggestive of freshwater residence. No general critical value for separating fresh and estuarine habitats was found for otolith Ba:Ca. Otolith Ba:Ca temporal patterns may assist the use of Sr:Ca in the evaluation of historical habitat residence and inter-habitat movement but the use of otolith Ba:Ca values should be applied cautiously for American eels and perhaps of other estuarine/freshwater migratory fishes.     

Incorporation of strontium into otoliths of an estuarine fish

Patterns of Sr/Ca variability in fish otoliths have been widely applied as tracers of movement between freshwater and marine habitats, with the assumption that low salinity habitats correspond to lower otolith levels of Sr/Ca. On the other hand, fluvial estuaries can contain steep gradients in Sr/Ca, and in some estuaries, freshwater values of Sr/Ca can exceed marine values, which are relatively constant across marine habitats. Therefore, to interpret Sr/Ca variability in otoliths of fish that move through estuaries, information is needed about both the incorporation of strontium into otoliths and the nature of the gradient of Sr/Ca in the water. We conducted four experiments to evaluate the incorporation of strontium into fish otoliths under estuarine conditions, using white perch (Morone americana) as a model estuarine fish. One laboratory and the two field experiments tested the relationship between Sr/Ca in the otolith and that in the water. A fourth experiment investigated the effect of salinity, independently of the water chemistry (Sr was manipulated while maintaining a constant salinity and Ca level). All four experiments supported a direct relationship between Sr/Ca in the otolith and the water, across a range of estuarine salinities. Results also indicated that the incorporation of strontium into otoliths of estuarine fishes should be constant across broad gradients of Sr/Ca in estuarine waters. While the experiments supported past applications of tracing estuarine and diadromous movements with otolith Sr/Ca chronologies, we emphasize the need to understand the underlying nature of Sr/Ca gradients in estuaries, which may limit or confound reconstructions of estuarine habitat use.     

Deep-ocean origin of the freshwater eels

Of more than 800 species of eels of the order Anguilliformes, only freshwater eels (genus Anguilla with 16 species plus three subspecies) spend most of their lives in freshwater during their catadromous life cycle. Nevertheless, because their spawning areas are located offshore in the open ocean, they migrate back to their specific breeding places in the ocean, often located thousands of kilometres away. The evolutionary origin of such enigmatic behaviour, however, remains elusive because of the uncertain phylogenetic position of freshwater eels within the principally marine anguilliforms. Here, we show strong evidence for a deep oceanic origin of the freshwater eels, based on the phylogenetic analysis of whole mitochondrial genome sequences from 56 species representing all of the 19 anguilliform families. The freshwater eels occupy an apical position within the anguilliforms, forming a highly supported monophyletic group with various oceanic midwater eel species. Moreover, reconstruction of the growth habitats on the resulting tree unequivocally indicates an origination of the freshwater eels from the midwater of the deep ocean. This shows significant concordance with the recent collection of mature adults of the Japanese eel in the upper midwater of the Pacific, suggesting that they have retained their evolutionary origin as a behavioural trait in their spawning areas.     

Application of otolith microchemistry to estimate the migratory history of Japanese eel Anguilla japonica on the Sanriku Coast of Japan

The age and migratory history of the Japanese eel, Anguilla japonica Temminck & Schlegel, collected in Miyako Bay along the Sanriku coast of Japan, was examined using the otolith microstructure and analysis of strontium (Sr) and calcium (Ca) concentrations conducted with wavelength dispersive X‐ray spectrometry by an electron microprobe. The line analysis of Sr : Ca ratios along the life history transect of each otolith showed a peak (ca. 15–17 × 10^?3) which corresponded with the period of their leptocephalus and early glass eel stages in the ocean. The mean Sr : Ca ratios from the elver mark to the otolith edge indicated that there were eels with several general categories of migratory history, including sea eels that never entered freshwater (average Sr : Ca ratios, ≥6.0 × 10^?3), and others that entered freshwater for brief periods but returned to the estuary or bay. This evidence of the occurrence of sea eels in this northern area indicates that Japanese eels of the Sanriku coast do not necessarily migrate into freshwater rivers during recruitment as do glass eels at the beginning of their growth phase; even those that do enter freshwater may later return to the marine environment. Thus, anguillid eel migrations into freshwater are clearly not an obligatory migratory pathway, but rather a facultative catadromy with seawater or estuarine residents as an ecophenotype.     

Differences in the Trace Element Deposition in Otoliths Between Marine- and Freshwater-resident Japanese Eels, Anguilla japonica, as Determined by Laser Ablation ICPMS

Synopsis In order to determine whether the trace element composition in otolith of the Japanese eel Anguilla japonica could be used to determine its habitat use, we used laser ablation inductivity coupled plasma mass spectrometry (LA-ICPMS) to assay sectioned otoliths of both marine-resident (sea eels) and freshwater-resident (river eels) eels. A close linear relationship in the Sr:Ca ratios between EPMA (X-ray analysis with an electron microprobe) and LA-ICPMS analyses was found, suggesting that the latter technique could be used to separate the marine and freshwater life phases. Elemental signatures in the otolith outside the elver mark showed significant differences in Cr:Ca, Mn:Ca, and Ba:Ca ratios as well as Sr:Ca ratios between sea and river eels. These results indicate that the elemental compositions may reflect environmental variability between marine and fresh water masses. Thus, those elemental ratios determined by LA-ICPMS analysis seem to have the potential to help distinguish the habitat of the eel.     

Migratory patterns and contribution of stocking to the population of European eel in Lithuanian waters as indicated by otolith Sr:Ca ratios

Otolith Sr:Ca ratios were examined to evaluate the contribution of the stocked eel Anguilla anguilla elvers, which have been stocked in Lithuanian waters and mixed with naturally recruited eels for several decades, to the native eel population. Stocked eels were identified by the freshwater signature (Sr:Ca ratios <2·24 × 10⁻³) on the otolith after the glass eel stage. Naturally recruited eels, that had migrated through the North and Baltic Seas, were characterized by an extended seawater and brackish-water signature (Sr:Ca ratios >3·23 × 10⁻³) after the glass eel stage. Of 108 eels analysed, 21 eels had otolith Sr:Ca ratio profiles consistent with stocking while 87 showed patterns of natural recruitment. The ages of naturally recruited eels arriving in Lithuanian fresh waters varied from 1 to 10 years, with a mean ±  s.d . age of 5·2 ± 2·1 years. Eels from the inland Lake Baluošai were all freshwater residents of stocked origin. Stocked eels, however, accounted for only 20% of the eels from the Curonian Lagoon and 2% of eels sampled in Baltic coastal waters. This finding does not support the hypothesis that the eel fishery in the Curonian Lagoon depends mostly on stocking.     

Late-stage metamorphosing Conger myriaster leptocephali collected in a river estuary of Ariake Bay, Japan

Little is known about where anguilliform fishes complete metamorphosis from the leptocephalus to juvenile stage. Conger myriaster leptocephali in the final stages of metamorphosing were collected from the estuarine portions of the lower Rokkaku River and in the northwest region of Ariake Bay. Three metamorphosing leptocephali with PAM/TM ratios of 0.27–0.34 were collected in the lower river, and five (PAM/TM, 0.30–0.46) were collected at sea stations in the bay. The collection of metamorphosing larvae of this species in a river estuary with lower salinity and high turbidity raises the question of how much these habitats are used by the metamorphosing larvae of this marine eel species.     

Effects of body condition and water temperature on Anguilla anguilla glass eel migratory behavior

Glass eels arriving from the sea use alternative migratory tactics, leading either to the colonization of rivers or to an early settlement in marine or estuarine habitats. In the field, the migration may be environmentally affected by water temperature and the migratory behavior could be physiologically dependent on the body condition (energetic status). To investigate how these environmental and physiological effects on the migration are behaviorally mediated, we experimentally tested the effects of changes in water temperature and body condition on locomotor activity (upstream swimming) and salinity preference of Anguilla anguilla glass eels. Low water temperature reduced significantly both locomotor activity and preference for freshwater, in accordance with field data showing that low water temperatures hinder both the estuarine migration and river recruitment. Glass eels switched from a freshwater- towards a saltwater-preference as their body condition decreased, confirming that the energetic status may affect the migratory behavior. We suggest that, in the wild, this condition-dependent change in salinity preference of low body condition glass eels induces an early settlement in marine or estuarine habitats. Such a behavioral shift, stopping the energy expenditure linked to river-oriented migratory behavior, may be adaptive by limiting the probability of death due to exhaustion. Our results show that the glass eel migratory behavior, through locomotor activity and salinity preference, may be controlled by interacting physiological and environmental factors.     

Unravelling the life-history patterns and habitat preferences of the Japanese eel (Anguilla japonica) in the Pearl River,China

Eels have fascinated biologists for centuries due to their amazing long-distance migrations between freshwater habitats and very distant ocean spawning areas. The migratory life histories of the Japanese eel, Anguilla japonica, in the waters of south China are not very clear despite its ecological importance, and the need for fishery regulation and management. In this study, strontium (Sr) and calcium (Ca) microchemical profiles of the otoliths of silver eels were measured by X-ray electron probe microanalysis based on data collected from different habitats (including freshwater and brackish habitats), in the large subtropical Pearl River. The corresponding habitat preference characteristics were further analysed using redundancy analysis (RDA). A total of 195 Japanese eels were collected over 6 years. The collected individuals ranged from 180 to 771 mm in total length and from 8 to 612 g in body weight. Two-dimensional pictures of the Sr:Ca concentrations in otoliths revealed that the A. japonica in the Pearl River are almost entirely river eels, spending the majority of their lives in fresh water without exposure to salt water, while the catadromous migration time has delayed about 1 month in the Pearl River estuary in the past 20 years. RDA analysis further indicated that juveniles and adults preferred water with high salinity and high tide levels. Youth preferred habitats with high river fractals. Our findings contribute to a growing body of evidence showing that the eels are extremely scarce currently and conservation measures against them are imminent, including the protection of brackish and freshwater areas where they live in south China.     

Facultative amphidromy involving estuaries in an annual amphidromous fish from a subtropical marginal range

Variation in the life cycle of diadromous fishes can be explained by differential food availability between marine and freshwater habitats, since migration is often interpreted as a mechanism for exploiting food resources. Theoretically, a migration pattern of mainly remaining in freshwater occurs in tropical and subtropical habitats where fluvial productivity possibly exceeds marine productivity. However, in Yakugachi River, Amami-Oshima Island, southern Japan, low nutrient concentrations in the river suggest that food availability is limited for the subtropical Ryukyu-ayu Plecoglossus altivelis ryukyuensis. Since Ryukyu-ayu is an amphidromous fish that mainly grows in rivers after spending 2 months in the sea, limited food availability in rivers would force this species to migrate to other habitats with better food availability. Otolith increment and Sr:Ca analyses of 48 adult Ryukyu-ayu collected from the Yakugachi River revealed that all individuals visited estuaries more than three times after moving upstream. Although the specific growth rates of this species in the river had no correlation with the salinity profile in the fluvial period, this movement may be an adaptive choice because the salinity profile significantly affected the body size at maturity. Our results highlighted individual-based variations in amphidromous migration for utilizing estuaries, which could be explained by relatively higher productivity in estuarine than in freshwater and marine habitats.     

Determinants of dietary specialization: a comparison of two sympatric species of sea snakes

Why do some predator species specialize on only a single type of prey whereas others take a broad range? One critical determinant may be the ontogenetic range of body sizes of the predator compared to that of its prey. If any single prey taxon spans only part of the range of prey sizes ingestible by the predator, then the predator will be more likely to take multiple prey taxa. We exploit a model system that provides a robust opportunity to test this hypothesis. We studied two sympatric species of predatory sea snakes, similar in size and general ecology that feed on anguilliform fishes from different habitats in the Great Lagoon of New Caledonia. Eel species from soft‐bottom habitats must construct their own burrows, and thus tend to be more slender‐bodied and less variable in body size than eel species that inhabit variable‐sized crevices among hard coral. As a result, a laticaudine sea snake species (Laticauda saintgironsi) that feeds on hard‐coral‐dwelling eels relies primarily on a single prey species: juveniles take young eels whereas adults consume adult eels of the same species. In contrast, a laticaudine species (L. laticaudata) that forages on soft‐bottom eels switches its prey ontogenetically: juveniles take small eel species whereas adults consume large eel species. Thus, habitat‐imposed constraints on the range of body sizes within each prey taxon generate a striking difference in the degree of dietary specialization of two closely related, sympatric predator species.     

Variations in the migratory history of the tropical catadromous eels Anguilla bicolor bicolor and A. bicolor pacifica in south-east Asian waters

Fish movements between aquatic habitats of different salinity ranges (fresh, estuarine, marine) by the tropical catadromous eels Anguilla bicolor bicolor and A. bicolor pacifica were examined by analysing the otolith strontium and calcium concentrations of yellow (immature) and silver (mature) stage eels collected in south-east Asian (Indonesia, Malaysia and Vietnam) waters. The ratios suggest that all migratory-type eels, including freshwater, brackish water and marine residents, pass the river mouth. However, the habitat preference was different among the sites (countries). In Indonesia and Vietnam, most A. bicolor bicolor and A. bicolor pacifica were either marine or brackish water residents in this study. Alternatively, most A. bicolor bicolor were freshwater residents in Malaysia; such a typical catadromous migration pattern in these eels has not been found in previous studies. The wide range of otolith Sr:Ca in both subspecies indicates that the habitat use of these tropical eels was opportunistic among fresh, brackish and marine waters during their growth phases following recruitment to coastal areas. The geographical variability of migratory histories suggests that habitat use might be determined by the inter and intraspecific competition and environmental conditions at each site.     

Stable isotope analyses provide new insights into ecological plasticity in a mixohaline population of European eel

Recent studies have shown that anguillid eel populations in habitats spanning the marine–freshwater ecotone can display extreme plasticity in the range of catadromy expressed by individual fishes. Carbon and nitrogen stable isotope analysis was used to differentiate between European eels (Anguilla anguilla) collected along a short (2 km) salinity gradient ranging from <1‰ to ~30‰ in Lough Ahalia, a tidal Atlantic lake system. Significant differences were recorded in mean δ¹³C, δ¹⁵N and C:N values from eels collected from fresh, brackish and marine-dominated basins. A discriminant analysis using these predictor variables correctly classified ca. 85% of eels to salinity zone, allowing eels to be classified as freshwater (FW), brackish (BW) or marine (MW) residents. The results of the discriminant analysis also suggested that a significant proportion of eels moved between habitats (especially between FW and BW). Comparisons of several key population parameters showed significant variation between eels resident in different salinity zones. Mean condition and estimated age was significantly lower in MW eels, whilst observed length at age (a correlate of growth) was significantly higher in MW eels, intermediate in BW and lowest in FW eels. This study has demonstrated that the ecology of eels found along a short salinity gradient can be extremely plastic and that stable isotope analysis has considerable utility in demonstrating intra-population variation in diadromous fishes.     

Changes in otolith microchemistry of the Japanese eel, Anguitta japonica, during its migration from the ocean to the rivers of Taiwan

The newly recruited Japanese eel, Anguilla japonica , elvers and 1-year-old eels collected in estuaries and in rivers, respectively, were studied. The microstructure and chemical composition of the sagittal otolith of these eels were examined by SEM and wavelength-dispersive spectrometer (WDS), A transition zone or'elver mark'was observed in the otolith of the young eels. A comparison of the otoliths of elvers with those from the 1-year-old eels suggests that this transition zone was deposited during upstream migration, a change from a marine to freshwater environment. Strontium (Sr) content in the primordium of the otolith of both elvers and young eels was low, probably due to the maternal or freshwater origin of the oocyte. The concentration of Sr in the otolith increased gradually during marine life and reached a peak approximately 1 month before upstream migration. As the elvers entered the estuary, the Src concentration dramatically decreased and remained at a low level thereafter. These findings indicate that the history of the migratory environment of the eel can be reconstructed from a combined study of otolith microstructure and microchemistry analysis.     

Biogeography,habitat transitions and hybridization in a radiation of South American silverside fishes revealed by mitochondrial and genomic RAD data

Rivers and lake systems in the southern cone of South America have been widely influenced by historical glaciations, carrying important implications for the evolution of aquatic organisms, including prompting transitions between marine and freshwater habitats and by triggering hybridization among incipient species via waterway connectivity and stream capture events. Silverside fishes (Odontesthes) in the region comprise a radiation of 19 marine and freshwater species that have been hypothesized on the basis of morphological or mitochondrial DNA data to have either transitioned repeatedly into continental waters from the sea or colonized marine habitats following freshwater diversification. New double digest restriction‐site associated DNA data presented here provide a robust framework to investigate the biogeographical history of and habitat transitions in Odontesthes. We show that Odontesthes silversides originally diversified in the Pacific but independently colonized the Atlantic three times, producing three independent marine‐to‐freshwater transitions. Our results also indicate recent introgression of marine mitochondrial haplotypes into two freshwater clades, with more recurring instances of hybridization among Atlantic‐ versus Pacific‐slope species. In Pacific freshwater drainages, hybridization with a marine species appears to be geographically isolated and may be related to glaciation events. Substantial structural differences of estuarine gradients between these two geographical areas may have influenced the frequency, intensity and evolutionary effects of hybridization events.     

Tropical aquatic fungi

This paper reports on the distribution of fungal communities in aquatic habitats in tropical regions and highlights differences in the taxa observed in freshwater and marine habitats. Ascomycetes are dominant on substrata in marine environments, with few basidiomycetes and discomycetes. Equally, few freshwater basidiomycetes and discomycetes have been reported from the tropics. In marine habitats, Dothideomycetes dominate on mangrove substrata, and halosphaeriaceous species are most numerous on submerged woody substrata in coastal waters, while yeasts are common in seawater and estuarine habitats. In freshwater, Ingoldian anamorphic fungi are most numerous on decaying leaves, while ascomycetes (Dothideomycetes, Sordariomycetes) are prevalent on submerged/exposed woody substrata. Unique fungi are found in tropical waters and differ from those in temperate locations.     

Regional variation in otolith Sr:Ca ratios of African longfinned eel Anguilla mossambica and mottled eel Anguilla marmorata: a challenge to the classic tool for reconstructing migratory histories of fishes

Otolith Sr:Ca ratios of the African longfinned eel Anguilla mossambica and giant mottled eel Anguilla marmorata from nine freshwater sites in four rivers of South Africa were analysed to reconstruct their migratory life histories between freshwater and saltwater habitats. For A. mossambica, the Sr:Ca ratios in the otolith edge differed significantly among rivers and had large effect sizes, but did not differ among sites within a river. Otolith Sr:Ca ratios did not differ among rivers for A. marmorata. When rivers were pooled, the edge Sr:Ca ratios of A. mossambica were not significantly different from those of A. marmorata. According to the river-specific critical Sr:Ca ratio distinguishing freshwater from saltwater residence, most A. mossambica and A. marmorata had saltwater habitat experience after settlement in fresh water. This was primarily during their elver stage or early in the yellow eel stage. During the middle and late yellow eel stage, freshwater residency was preferred and only sporadic visits were made to saltwater habitats. The data also suggest that regional variations in otolith Sr:Ca ratios affect the critical Sr:Ca value and are a challenge for the reconstruction of migratory life histories that should be explicitly considered to avoid bias and uncertainty.     

The icthyofauna of the Mhlathuze coastal lakes: some preliminary results

The fish faunas of the four Mhlathuze coastal lakes and the lower river comprise a diverse assortment of over fifty marine, estuarine and freshwater species. Three freshwater species are endemic to KwaZulu-Natal and nine estuarine species are endemic to southern Africa. Five species are of conservation significance. Species numbers in Lakes Mzingazi and Cubhu are similar historically and both lakes served as secondary nursery habitats for estuarine associated fishes. This role has been impacted by the construction of weirs at their outlets which prevent successful recruitment of estuarine species, especially during drought years when lake water levels are low. The fish faunas of Lakes Nsezi and Mangeza are depauperate and lack marine or estuarine components. In order that these systems fulfil their potential function as secondary nursery habitats to many estuarine fish species, minimum lake water levels must be set to ensure sufficient outflows at proposed fish ladders during critical spawning and migration times.     

Interspecific and sexual differences in riverine distribution of tropical eels Anguilla spp.

A total of 261 individuals of the four tropical eel species, Anguilla celebesensis, Anguilla marmorata, Anguilla bicolor pacifica and Anguilla interioris, were collected from 12 locations around Sulawesi Island, Indonesia, to gain knowledge about the riverine distribution of tropical eels. Anguilla marmorata was predominant in the lower reaches of Poso River (94·4% of total eel catch in the sampling area), Poso Lake (93·3%), three small inlet rivers of Tomini Bay (100%) and Laa River (92·3%). Anguilla celebesensis occurred frequently in the inlet rivers of Poso Lake (63·5%). Anguilla bicolor pacifica and Anguilla interioris were rare (1.5 and 0.4%, respectively). Otolith Sr:Ca ratio electron‐probe micro analysis (EPMA) for individual migratory histories revealed that 15 A. celebesensis caught in Poso Lake and its inlet rivers were categorized into 14 river eels (Sr:Ca < 2·5) showing upstream migration seemingly at their elver stage and only one sea eel (Sr:Ca ≥ 6·0) that stayed in the marine habitat for the majority of its life after recruiting to Sulawesi Island before its late upstream migration. In A. marmorata, 19 examined eels from Poso Lake and its inlet rivers were all river eels, while 17 eels from the lower reaches of Poso River were two river eels, six sea eels and nine estuarine eels (2·5 ≤ Sr:Ca < 6·0) that mostly lived in the brackish water. The sex ratio of A. celebesensis was highly skewed towards a dominance of females (99%). In A. marmorata, females were predominant in Poso Lake (95·2%), its inlet rivers (94·7%) and Laa River (100%), while males were more frequent in the lower reaches of Poso River (76·5%) and small inlet rivers of Tomini Bay (94·1%). These results indicate that the riverine distribution pattern of tropical eels differs among species and between sexes.