A capture–recapture model of amphidromous fish dispersal

Adult movement scale was quantified for two tropical Caribbean diadromous fishes, bigmouth sleeper Gobiomorus dormitor and mountain mullet Agonostomus monticola, using passive integrated transponders (PITs) and radio‐telemetry. Large numbers of fishes were tagged in Río Mameyes, Puerto Rico, U.S.A., with PITs and monitored at three fixed locations over a 2·5 year period to estimate transition probabilities between upper and lower elevations and survival probabilities with a multistate Cormack–Jolly–Seber model. A sub‐set of fishes were tagged with radio‐transmitters and tracked at weekly intervals to estimate fine‐scale dispersal. Changes in spatial and temporal distributions of tagged fishes indicated that neither G. dormitor nor A. monticola moved into the lowest, estuarine reaches of Río Mameyes during two consecutive reproductive periods, thus demonstrating that both species follow an amphidromous, rather than catadromous, migratory strategy. Further, both species were relatively sedentary, with restricted linear ranges. While substantial dispersal of these species occurs at the larval stage during recruitment to fresh water, the results indicate minimal dispersal in spawning adults. Successful conservation of diadromous fauna on tropical islands requires management at both broad basin and localized spatial scales.     

Low interbasin connectivity in a facultatively diadromous fish: evidence from genetics and otolith chemistry

Southern smelts (Retropinna spp.) in coastal rivers of Australia are facultatively diadromous, with populations potentially containing individuals with diadromous or wholly freshwater life histories. The presence of diadromous individuals is expected to reduce genetic structuring between river basins due to larval dispersal via the sea. We use otolith chemistry to distinguish between diadromous and nondiadromous life histories and population genetics to examine interbasin connectivity resulting from diadromy. Otolith strontium isotope (⁸⁷Sr:⁸⁶Sr) transects identified three main life history patterns: amphidromy, freshwater residency and estuarine/marine residency. Despite the potential for interbasin connectivity via larval mixing in the marine environment, we found unprecedented levels of genetic structure for an amphidromous species. Strong hierarchical structure along putative taxonomic boundaries was detected, along with highly structured populations within groups using microsatellites (F_ST = 0.046–0.181), and mtDNA (Φ_ST = 0.498–0.816). The presence of strong genetic subdivision, despite the fact that many individuals reside in saline water during their early life history, appears incongruous. However, analysis of multielemental signatures in the otolith cores of diadromous fish revealed strong discrimination between river basins, suggesting that diadromous fish spend their early lives within chemically distinct estuaries rather than the more homogenous marine environment, thus avoiding dispersal and maintaining genetic structure.     

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.     

Reconstructing larval growth and habitat use in an amphidromous goby using otolith increments and microchemistry

High‐resolution analysis of growth increments, trace element chemistry and oxygen isotope ratios (δ¹⁸O) in otoliths were combined to assess larval and post‐larval habitat use and growth of Awaous stamineus, an amphidromous goby native to Hawai‘i. Otolith increment widths indicate that all individuals experience a brief period of rapid growth during early life as larvae and that the duration of this growth anomaly is negatively correlated with larval duration. A protracted high‐growth period early in larval life is associated with a lower ratio of Sr:Ca, which may reflect low salinity conditions in nearshore habitats. A distinct shift in δ¹⁸O (range: 4–5‰) is closely associated with the metamorphic mark in otoliths, indicating that larval metamorphosis occurs promptly upon return to fresh water. Strontium and other trace elements are not as tightly coupled to the metamorphosis mark, but confirm the marine‐to‐freshwater transition. Integration of microstructural and microchemical approaches reveals that larvae vary substantially in growth rate, possibly in association with habitat differences. Although time and financial costs make it difficult to achieve large sample sizes, present results show that examining even a small number of individuals can lead to novel inferences about early life history in diadromous fishes and illustrates the value of integrating analyses.     

Elevated river discharge enhances the immigration of juvenile catadromous and amphidromous fishes into temperate coastal rivers

Anthropogenic alterations to river flow regimes threaten freshwater biodiversity globally, with potentially disproportionate impacts on species that rely on flow cues to trigger critical life history processes, such as migration for diadromous fishes. This study investigates the influence of river discharge on the abundance of juvenile fish moving into rivers by four temperate catadromous or amphidromous species (common galaxias Galaxias maculatus, spotted galaxias Galaxias truttaceus, climbing galaxias Galaxias brevipinnis and the threatened Australian grayling Prototroctes maraena). Fyke netting or fishway trapping was used to catch juvenile fish moving from estuaries into freshwater in five coastal waterways in south-eastern Australia during the spring migratory period. There was a positive relationship between the probability of high catch rates and mean discharge in September. We also found a positive relationship between discharge and the number of recruits captured 22–30 days later in a flow stressed system. In addition, day-of-year had a strong influence on catch rates, with the peak abundance of juveniles for three species most likely to occur midway through the sampling period (spotted galaxias in October, climbing galaxias in late October and Australian grayling in late October and early November). Our study shows that higher magnitudes of river discharge were associated with increased catches of juvenile catadromous and amphidromous fishes. With a limited supply of environmental water, environmental flows used to enhance immigration of these fishes may be best targeted to maintain small amounts of immigration into freshwater populations in waterways or years when discharges are low and stable. When there are natural, large discharge volumes, relatively large numbers of juvenile fish can be expected to enter coastal waterways and during these times environmental flows may not be required to promote immigration.     

Otolith chemistry of prey fish consumed by a fish predator: does digestion hinder Russian doll techniques?

The effect of digestion by a predatory fish (largemouth bass Micropterus salmoides) on stable isotopic (δ¹³C and δ¹⁸O) and trace elemental (Sr:Ca and Ba:Ca) compositions of prey fish (bluegill Lepomis macrochirus) otoliths was investigated in a laboratory experiment. Trace element and stable‐isotopic signatures of L. macrochirus otoliths were not significantly altered for up to 16 h after L. macrochirus were consumed by M. salmoides. Prey fish otoliths recovered from predator digesta can retain environmental stable isotopic and trace elemental signatures, suggesting that determination of environmental history for prey fishes by stable‐isotope and trace‐element analysis of otoliths recovered from stomachs of piscivorous fishes will be feasible.     

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.     

Movements of diadromous fish in large unregulated tropical rivers inferred from geochemical tracers

Patterns of migration and habitat use in diadromous fishes can be highly variable among individuals. Most investigations into diadromous movement patterns have been restricted to populations in regulated rivers, and little information exists for those in unregulated catchments. We quantified movements of migratory barramundi Lates calcarifer (Bloch) in two large unregulated rivers in northern Australia using both elemental (Sr/Ba) and isotope ((87)Sr/(86)Sr) ratios in aragonitic ear stones, or otoliths. Chemical life history profiles indicated significant individual variation in habitat use, particularly among chemically distinct freshwater habitats within a catchment. A global zoning algorithm was used to quantify distinct changes in chemical signatures across profiles. This algorithm identified between 2 and 6 distinct chemical habitats in individual profiles, indicating variable movement among habitats. Profiles of (87)Sr/(86)Sr ratios were notably distinct among individuals, with highly radiogenic values recorded in some otoliths. This variation suggested that fish made full use of habitats across the entire catchment basin. Our results show that unrestricted movement among freshwater habitats is an important component of diadromous life histories for populations in unregulated systems.     

Habitat Uses of Freshwater Fishes on the Scale of Reach System Provided in Small Streams

Synopsis Before we can conserve freshwater fish communities, we must first elucidate the habitat uses of each species. We investigated the relationships between the habits of freshwater fishes and environmental factors in small streams located in western Toyama Prefecture, Japan. We found 30 species – 11 fluvial, 5 brackish water, 2 anadromous, 11 amphidromous, and 1 catadromous – by a semi-quantitative collecting method. Using binary data (occurrence/non-occurrence of dominant fish), we conducted logistic regression analyses to construct predictive models. Stepwise logistic regression analyses on the basis of eight biotic and abiotic factors showed that the occurrence of dominant fish was explained by environmental conditions. The flow variation, ratio of stream bed slope to the surrounding area, distance from sea, and drift plant richness were relevant. Freshwater minnow, Phoxinus oxycephalus jouyi, preferred the gentle slopes of upstream catchments. The freshwater goby, Rhinogobius flumineus, appeared to prefer steeper slopes relative to the surrounding area. In contrast, diadromous gobies preferred reaches near the sea. These suggest a correspondence between lifestyle and preferred habitat. Consequently, the maintenance of various habitat conditions is necessary for the conservation of freshwater fishes.     

Fish mediate high food web connectivity in the lower reaches of a tropical floodplain river

High levels of hydrological connectivity during seasonal flooding provide significant opportunities for movements of fish between rivers and their floodplains, estuaries and the sea, possibly mediating food web subsidies among habitats. To determine the degree of utilisation of food sources from different habitats in a tropical river with a short floodplain inundation duration (~2 months), stable isotope ratios in fishes and their available food were measured from three habitats (inundated floodplain, dry season freshwater, coastal marine) in the lower reaches of the Mitchell River, Queensland (Australia). Floodplain food sources constituted the majority of the diet of large-bodied fishes (barramundi Lates calcarifer, catfish Neoarius graeffei) captured on the floodplain in the wet season and for gonadal tissues of a common herbivorous fish (gizzard shad Nematalosa come), the latter suggesting that critical reproductive phases are fuelled by floodplain production. Floodplain food sources also subsidised barramundi from the recreational fishery in adjacent coastal and estuarine areas, and the broader fish community from a freshwater lagoon. These findings highlight the importance of the floodplain in supporting the production of large fishes in spite of the episodic nature and relatively short duration of inundation compared to large river floodplains of humid tropical regions. They also illustrate the high degree of food web connectivity mediated by mobile fish in this system in the absence of human modification, and point to the potential consequences of water resource development that may reduce or eliminate hydrological connectivity between the river and its floodplain.     

Life histories of closely related amphidromous and non‐migratory fish species: a trade‐off between egg size and fecundity

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Otolith trace element and stable isotopic compositions differentiate fishes from the Middle Mississippi River, its tributaries, and floodplain lakes

Naturally occurring stable isotope and trace elemental markers in otoliths have emerged as powerful tools for determining natal origins and environmental history of fishes in a variety of marine and freshwater environments. However, few studies have examined the applicability of this technique in large river-floodplain ecosystems. This study evaluated otolith microchemistry and stable isotopic composition as tools for determining environmental history of fishes in the Middle Mississippi River, its tributaries, and floodplain lakes in Illinois and Missouri, USA. Fishes were collected from 14 sites and water samples obtained from 16 sites during summer and fall 2006 and spring 2007. Otolith and water samples were analyzed for stable oxygen isotopic composition (δ¹⁸O) and concentrations of a suite of trace elements; otoliths were also analyzed for carbon isotopic composition (δ¹³C). Tributaries, floodplain lakes, and the Mississippi and Lower Missouri Rivers possessed distinct isotopic and elemental signatures that were reflected in fish otoliths. Fish from tributaries on the Missouri and Illinois sides of the middle Mississippi River could also be distinguished from one another by their elemental and isotopic fingerprints. Linear discriminant function analysis of otolith chemical signatures indicated that fish could be classified back to their environment of capture (Mississippi River, floodplain lake, tributary on the Illinois or Missouri side of the Mississippi River, or lower Missouri River) with 71–100% accuracy. This study demonstrates the potential applicability of otolith microchemistry and stable isotope analyses to determine natal origins and describe environmental history of fishes in the Middle Mississippi River, its tributaries, and floodplain lakes. The ability to reconstruct environmental history of individual fish using naturally occurring isotopic markers in otoliths may also facilitate efforts to quantify nutrient and energy subsidies to the Mississippi River provided by fishes that emigrate from floodplain lakes or tributaries.     

Life history of Coilia nasus from the Yellow Sea inferred from otolith Sr:Ca ratios

The habitat use and migratory patterns of the estuarine tapertail anchovy, Coilia nasus, from three sites near- or off-shore of the Yellow Sea, were studied by examining the environmental signatures of Sr and Ca in otoliths using electron probe microanalysis (EPMA). All the samples analyzed were offsprings of typical anadromous C. nasus. The fish grew in freshwater habitat nearly for 1?year, and then entered the brackish and sea water which could be as far as ca. 300?km from the Chinese coast line. The Sr:Ca ratios at the edge of otoliths appeared to correlated with water salinities in sampling sites.     

The use of otolith chemistry to characterize diadromous migrations

Chemical constituents in otoliths have become a valuable tool for fish ecologists seeking to reconstruct migratory patterns and life-history diversity in a wide range of species worldwide. This approach has proved particularly effective with fishes that move across substantial salinity gradients over the course of their life, including many diadromous species. Freshwater endmembers of several elemental and isotope ratios (e.g. Sr:Ca, Ba:Ca and (87)Sr:(86)Sr) are typically identifiably distinct from marine values, and often differ among freshwater tributaries at fine spatial scales. Because these chemical tags are generally incorporated in proportion to their ambient dissolved concentrations, they can be effective proxies for quantifying the presence, duration and frequency of movements between freshwater and marine habitats. The development of high precision probe-based analytical techniques, such as laser ablation inductively coupled plasma mass spectrometry (ICP-MS) and microbeam methods, has allowed researchers to glean increasingly detailed life-history profiles of these proxies across otoliths. Researchers are also combining multiple chemical proxies in an attempt to refine interpretations of habitat residence patterns. A thorough understanding of the spatial and temporal variation in water chemistry as well as environmental and physiological controls on incorporation of specific elements into otoliths is required for confident estimation of lifetime salinity experience. First some assumptions, methodological considerations and data processing options that are particularly relevant to diadromous otolith chemistry studies are discussed. Insights into diadromous migrations obtained from decades of otolith chemistry research, highlighting the increasingly recognized importance of contingent behaviour and partial migration are then discussed. Finally, areas for future research and the need to integrate otolith chemistry studies into comprehensive assessments of the effects of global environmental change are identified.     

Overcoming urban stream syndrome: Trophic flexibility confers resilience in a Hawaiian stream fish

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The effects of diadromy and its loss on genomic divergence: The case of amphidromous Galaxias maculatus populations

Understanding the evolutionary mechanisms that affect the genetic divergence between diadromous and resident populations across heterogeneous environments is a challenging task. While diadromy may promote gene flow leading to a lack of genetic differentiation among populations, resident populations tend to be affected by local adaptation and/or plasticity. Studies on these effects on genomic divergence in nonmodel amphidromous species are scarce. Galaxias maculatus, one of the most widespread fish species in the Southern Hemisphere, exhibits two life histories, an ancestral diadromous, specifically, amphidromous form, and a derived freshwater resident form. We examined the genetic diversity and divergence among 20 estuarine and resident populations across the Chilean distribution of G. maculatus and assessed the extent to which selection is involved in the differentiation among resident populations. We obtained nearly 4,400 SNP markers using a RADcap approach for 224 individuals. As expected, collections from estuarine locations typically consist of diadromous individuals. Diadromous populations are highly differentiated from their resident counterparts by both neutral and putative adaptive markers. While diadromous populations exhibit high gene flow and lack site fidelity, resident populations appear to be the product of different colonization events with relatively low genetic diversity and varying levels of gene flow. In particular, the northernmost resident populations were clearly genetically distinct and reproductively isolated from each other suggesting local adaptation. Our study provides insights into the role of life history differences in the maintenance of genetic diversity and the importance of genetic divergence in species evolution.     

Extrinsic factors affecting otolith chemistry – implications for interpreting migration patterns in a diadromous fish

Much has been revealed about fish migration, including diadromous behaviour, through the use of otolith chemistry. Manipulative experiments assist with unravelling information on otolith chemical composition and incorporation thereby answering specific questions on diadromous movements. In this study, a laboratory-based experiment was used to determine the relative and interactive effects of salinity and water temperature on the composition of three key elements (Sr, Ba and Mg) within the otolith of a catadromous fish, Percalates novemaculeata, endemic to south-eastern Australia. Otolith incorporation of Sr and Ba was positively related to ambient water concentration, whereas Mg incorporation was not. Sr and Ba increased and decreased significantly across salinity gradients, respectively, with minor positive effects of temperature also being detected. Salinity and temperature interacted to significantly affect the elemental concentration ratios for Ba: Ca in otoliths. Discrimination between fresh water and marine environments shows promise for interpreting P. novemaculeata residency based on these elements alone. However, deciphering finer scale movements within estuarine environments may be difficult. Our data highlights the importance of multifactorial validation experiments and suggests complementary use of multiple approaches for unravelling species-specific patterns of fish movement and habitat use.     

Flexible migration of Japanese freshwater gobies Rhinogobius spp. as revealed by otolith Sr:Ca ratios

The migratory histories of six Rhinogobius spp., the cross‐band type (R. sp. CB), the large‐dark type (R. sp. LD), the dark type (R. sp. DA), the cobalt type (R. sp. CO), the orange type (R. sp. OR) and R. flumineus, were studied by examining strontium (Sr) and calcium (Ca) concentrations in their otoliths using wavelength dispersive X‐ray spectrometry on an electron microprobe. The Sr:Ca ratios in the otoliths changed both with the ontogenetic development and with the salinity level of the habitat. Most fishes had high Sr:Ca ratios around the otolith core in spite of the fact that those fishes live most of their lives in a freshwater environment. The high ratios in the otoliths were thought to be a physiological effect in those fishes. Thereafter, the Sr:Ca ratios changed remarkably along the life‐history transect, showing intraspecies and interspecies variations. The otolith Sr:Ca ratios of Rhinogobius sp. CB, R. sp. LD and R. sp. CO collected from three rivers connected to the sea were low around the core, subsequently increased sharply to the points 180–345 μm from the core and then decreased again towards the edge. They were thought to reflect the typical amphidromous life history. The R. sp. CO, however, remain in a brackish‐water environment after migration from the sea, while the other species showed typical amphidromous lives with complete freshwater residence after migration from the sea. The five species (R. sp. CB, R. sp. LD, R. sp. CO, R. sp. DA and R. sp. OR) collected above dams had never migrated to the sea, spending their whole life in a freshwater environment, although Rhinogobius species, except for the fluvial type, were thought to have an amphidromous life history according to previous studies. These species are thought to have a landlocked life cycle. The otolith Sr:Ca ratios of R. flumineus showed consistently low ratios towards the edge except for only around the core, although they were collected from a river connected to the sea. These species could have a fluvial life history corresponding to a previous study. The present study clearly suggests that the migratory histories of Rhinogobius spp. are highly different both within and between species and that they have flexible migratory patterns allowing them to utilize the full range of salinity during their life history.     

Heterogeneity of otolith chemical composition from two-dimensional mapping: Relationship with biomineralization mechanisms and implications for microchemistry analyses

Although otoliths are widely used as archives to infer life-history traits and habitat use in fishes, their biomineralization process remains poorly understood. This lack of knowledge is problematic as it can lead to misinterpretation of the different types of signals (e.g., optical or chemical) that provide basic data for research in fish ecology, fisheries management, and species conservation. Otolith calcification relies on a complex system involving a pericrystalline fluid, the endolymph, whose organic and inorganic compositions are spatially heterogeneous for some constituents. This property stems from the particular structure of the calcifying saccular epithelium. In this study, we explored the spatial heterogeneity of elemental incorporation in otoliths for two species of high economic interest, European hake Merluccius merluccius (L. 1758) and European sea bass Dicentrarchus labrax (L. 1758). Two-dimensional mappings of chemical elements were obtained using UV high-repetition-rate femtosecond laser ablation (fs-LA) system coupled to a high-resolution inductively coupled plasma sector field mass spectrometer analyses on transverse sections of sagittae. Results highlighted a clear asymmetry between proximal (sulcus) and distal (antisulcus) concentrations for elements such as magnesium (Mg), phosphorus (P), manganese (Mn), and potassium (K) with concentration gradient directions that varied depending on the element. Strontium (Sr) and barium (Ba) did not show a proximo-distal gradient. These results are discussed in light of current knowledge on the endolymph composition and the mechanisms that lead to its compartmentalization, highlighting the need for further research on otolith biomineralization. Operational implications for studies based on otolith chemical composition are also discussed with emphasis on advice for sampling strategies to avoid analytical biases and the need for in-depth analyses of analytical settings before comparing otolith signatures between species or geographical areas.     

Use of otolith chemistry to examine patterns of diadromy in the threatened Australian grayling Prototroctes maraena

Otolith chemical analyses (proton-induced X-ray emission and laser ablation inductively coupled plasma mass spectrometry) were used to examine patterns of diadromy in Australian grayling Prototroctes maraena collected from three rivers in south-eastern Australia. Concentrations of Sr:Ca and Ba:Ca were measured in the sagittal otoliths of 25 fish and in water samples collected on two occasions from multiple sites within freshwater, estuarine and marine reaches of the rivers. The results provide evidence of marine residency during the juvenile phase, with high Sr:Ca and low Ba:Ca in the inner region of the otoliths that were consistent across all three rivers. This general pattern was apparent for all individuals examined, suggesting that diadromous migration may be an obligate aspect of the life history. The chemical signatures of the outer regions of the otoliths were distinct between the three rivers, whilst there was no difference in the inner regions of the otoliths. This suggests that juveniles from different rivers may reside in a relatively homogenous chemical environment, such as the sea, and populations in coastal Victorian rivers may share a common marine recruitment source.