Larval dispersal and population connectivity among a network of marine reserves

A major challenge in marine ecology is describing patterns of larval dispersal and population connectivity, as well as their underlying processes. We used a biophysical model to simulate dispersal of eastern oyster, Crassostrea virginica, larvae and connectivity among a network of 10 no‐take reserves in a shallow, wind‐driven estuary to assess the relative importance of spawning location, spawning date, larval behavior, larval mortality, and adult reproductive output to predicted dispersal and connectivity patterns. The location (i.e., natal reserve) and date of spawning relative to physical processes, particularly frequency of wind reversals, were the dominant drivers of dispersal and connectivity patterns. To a lesser extent, larval behavior (i.e., 3D vertical advection and ontogenetic depth regulation) and mortality modified dispersal and connectivity, whereas spatiotemporal variability in adult reproductive output was of minimal importance. Over a 21‐day larval duration, mean dispersal distance of passive surface particles ranged from 5 to 40 km. Reserves were too small (1 km²) relative to mean dispersal distances to promote extensive local retention (median 2%) and spaced too far apart (typically ~50 km) to promote extensive inter‐reserve connectivity (median 2%). Limited connectivity and local retention may preclude the network from being self‐sustainable, thereby limiting its long‐term conservation and management benefits. In reserve systems characterized by limited connectivity, management efforts should focus on increasing connectivity by increasing the number or size of reserves to realize the benefits of improved adult demographics within reserves.     

Dispersal and tributary immigration by juvenile coho salmon contribute to spatial expansion during colonisation

Anadromous fishes are frequently restricted by artificial barriers to movement such as dams and culverts, so measuring dispersal helps identify sites where improved connectivity could promote range expansion and population viability. We used a combination of DNA‐based parentage analysis and mark–recapture techniques to evaluate dispersal by juvenile coho salmon (Oncorhynchus kisutch) in a population in the initial stages of colonisation following installation of fish passage structures at a previously impassable dam on the Cedar River, WA, USA. The spatial distribution of individuals within maternal families revealed that dispersal was common. Among the offspring of radio‐tagged mothers, 28% were collected outside the spawning reach and dispersed up to 6.3 km (median = 1.5 km). Most juveniles captured in a tributary (Rock Creek, where few adults spawned) had immigrated from the Cedar River and represented many different families. Juvenile dispersal therefore provided a secondary phase of spatial expansion following initial colonisation by adults. Consistent with the condition‐dependent dispersal hypothesis, juveniles that dispersed farther upstream in the tributary were larger than fish collected near the tributary mouth. Overall, the results demonstrated widespread dispersal in a system with low coho salmon densities, and this might increase the rate of population growth if it reduces the effects of local density dependence. By implication, juveniles can take advantage of rearing habitats reconnected through barrier removal, even when such areas are located several kilometres from adult breeding grounds.     

Genetic differentiation between and within ecotypes of pike (Esox lucius) in the Baltic Sea

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Patterns and predictors of fish dispersal in rivers

Quantifying fish dispersal and identifying its general predictors is key for understanding temporal patterns in population dynamics, emigration and immigration, meta‐community dynamics, many ecological processes and predicting recovery time or population responses to environmental changes. This is the first comprehensive quantitative meta‐analysis of heterogeneous freshwater fish movement, aiming to determine mobile and stationary shares of fish communities, their dispersal distances and key predictors of dispersal patterns. By reviewing and analysing 160 empirical data sets from 71 studies covering 62 fishes in streams, it goes beyond previous studies of salmonids’ heterogeneous movement. Based on fitted leptokurtic dispersal kernels, the movement distances of (i) a stationary component (σ_stat) and (ii) a mobile component (σ_mob) as well as the (iii) share of each component (p) were calculated. The median movement distance of the stationary and mobile component of a fish population was 36.4 and 361.7 m, respectively. The share of the stationary individuals was high (median = 66.6%), but unrelated to movement distance. Single and multiple linear regressions as well as mixed‐effects models revealed movement distances positively related to fish length, aspect ratio of the caudal fin, stream size and duration of the study. Furthermore, movement distance differed between taxonomic families. The quantitative parameters of heterogeneous fish movement provided are prerequisite to estimate time lags in fish response to river rehabilitation, temporal patterns in species dispersal, and minimum effective size of potential founder populations for species conservation and stock recovery based on minimum numbers of specimen to disperse.     

Genetic structure of brown trout,Salmo trutta,populations from differently sized tributaries of Lake Mjøsa in south‐east Norway

This study was based on genotyping eight microsatellite loci of 463 brown trout, Salmo trutta L., sampled in nine differently sized tributaries in three areas on the eastern shore of Lake Mjøsa, south‐eastern Norway. The populations were genetically structured, and Mantel's test showed that genetic distance correlated positively with geographical distance. Temporal differentiation F_ST over a 2‐year period was estimated in five streams and was non‐significant after Bonferroni correction. Effective population size N_e was positively correlated with the habitat length available from the lake (0.3–22 km) and negatively with the number of full sib pairs in the sample. There was no correlation between N_e and genetic diversity, and private alleles were recorded in three medium‐sized streams, but not in the largest two. The importance of small spawning and nursery streams for the maintenance of genetic diversity of brown trout was demonstrated.     

Novel insights into the marine phase and river fidelity of anadromous twaite shad Alosa fallax in the UK and Ireland

1. Most research on anadromous fishes has been invested in their freshwater life‐phases, resulting in a relatively sparse understanding of their spatial ecology during marine life‐phases. However, understanding the marine dispersal of anadromous fishes is essential to identify threats and to implement conservation measures that fully encompass their lifecycle.
2. The twaite shad Alosa fallax is an anadromous fish increasingly imperilled across its range due to pollution, harvesting, and impediments to freshwater migration, but little is known about its distribution and movements during its marine life‐phase. Here, the application of acoustic telemetry provided novel insights into the coastal dispersal of twaite shad in the UK and Ireland during 2018–2019, and the freshwater entry of individuals during the 2019 spawning season.
3. Of 73 twaite shad acoustic‐tagged during their upstream migration in the River Severn in May 2018, 58 emigrated from the river. Twelve were subsequently detected 200 km to the south‐west at the Taw–Torridge Estuary between July 2018 and April 2019, where estuarine movements up to 5.8 km inland occurred in summer, winter, and spring. One was subsequently detected in the Munster Blackwater Estuary (Ireland) and then in the River Severn, indicating a minimum movement distance of 950 km. Thirty‐four (59%) of the emigrating individuals from 2018 re‐entered fresh water in the rivers Severn (n = 33) and Wye (n = 2) in April and May 2019.
4. These results suggest year‐round use of estuarine and nearshore habitats by at least a subset of the twaite shad population during their marine phase, providing evidence of potential range overlap between populations that spawn in different areas in the UK and Ireland, which may be facilitated by substantial dispersal. The results also highlight the potential of telemetry for estimating freshwater and marine mortality, and the benefits of sharing detection data across networks.

     

First evidence of spawning migration by goldfish (Carassius auratus); implications for control of a globally invasive species

Goldfish (Carassius auratus) was one of the first fishes to be domesticated and has been widely introduced across the globe, but is now considered one of the world's worst invasive aquatic species. Surprisingly, there is a dearth of information on its spatial and temporal movement patterns, which hampers the development of effective control programmes. We examined the movement patterns of an introduced population of C. auratus in a south‐western Australian river using passive acoustic telemetry. The study population had a high residency index within the array (i.e. proportion of all days at liberty that, on average, each fish was detected by a receiver) with fish being detected on 64% of days. The individuals were also reasonably mobile, travelling a mean of 0.30 km (linear river kilometres).day^‐1 within the array, and one fish moved 231.3 km over the 365‐day study period (including 5.4 km in a 24 hr period). Importantly, C. auratus displayed significant seasonal movement patterns including a clear shift in habitats during its breeding period with most mature individuals being detected in an off‐channel wetland during that time. The results of this study strongly suggest that C. auratus undertook a spawning migration into a lentic habitat. These results have important implications for developing control programmes for the species, such as targeting connections to off‐channel lentic systems during its breeding period.     

Spatial distribution patterns of the non‐native European catfish,Silurus glanis,from multiple online sources – a case study for the River Tagus (Iberian Peninsula)

Effective management of invasive fishes depends on the availability of updated information about their distribution and spatial dispersion. Forensic analysis was performed using online and published data on the European catfish, Silurus glanis L., a recent invader in the Tagus catchment (Iberian Peninsula). Eighty records were obtained mainly from anglers’ fora and blogs, and more recently from www.youtube.com . Since the first record in 1998, S. glanis expanded its geographic range by ≈700 km of river network, occurring mainly in reservoirs and in high‐order reaches. Human‐mediated and natural dispersal events were identified, with the former occurring during the first years of invasion and involving movements of >50 km. Downstream dispersal directionality was predominant. The analysis of online data from anglers was found to provide useful information on the distribution and dispersal patterns of this non‐native fish, and is potentially applicable as a preliminary, exploratory assessment tool for other non‐native fishes.     

Microsatellites markers to depict the reproductive and genetic patterns of farmed gilthead seabream (Sparus aurata): illustration by a case study on mass spawning

In the absence of breeding strategy, natural spawning constitutes the breeding ground for fish farmers to empirically manage their commercial broodstock. In this context, we used six microsatellite markers to characterize the genetic pattern of six commercial seabream broodfish tanks having a common history spanning four generations. The progeny of one tank single‐day mass‐spawning event, reared in two different environments, was used to estimate the genetic parameters for body weight. Limited genetic differentiation was observed among broodfish groups. A panel of nine loci allowed us to unambiguously assign 95.4% of the offspring (1692) and identify 37 parents (65% of the total broodfish). The limited effective population size (N_e = 15.3) was due to the elevated variance of parental contributions and to broodfish failing to contribute to the progeny. The fluctuation of the allele frequency highlighted the risks of genetic drift and reduction in the heterozygosity in the next generations. Heritability for body weight was moderate at commercial size (0.40 ± 0.10) and the high genetic correlation at later stages laid the groundwork for precocious selection criteria for growth. The discussion opens on the opportunity to use mass spawning for selective breeding.     

Multiple spawning events promote increased larval dispersal of a predatory fish in a western boundary current

Transport of larvae by ocean currents is an important dispersal mechanism for many species. The timing and location of spawning can have a large influence on settlement location. Shifts in the known spawning habitat of fish, whether due to climate or the discovery of new spawning stock, can influence the distribution of juveniles and our understanding of connectivity. The globally distributed species; Pomatomus saltatrix, is one such example where a previously unrecognised summer spawning event and a more southern latitudinal extent was recently reported for the southwest Pacific population. Although restrictions are in place to protect the traditional spawning event, the importance of the newly recognised summer spawning event is uncertain. Here, we investigate larval dispersal of P. saltatrix using particle tracking simulations to identify the contributions of the different spawning events to settlement. By modelling dispersal of larvae released in northern and mid‐latitude regions over the Austral spring and summer, we show that the newly recognised mid‐latitude summer spawning event contributes over 50% of the larvae reaching southern latitudes. This is due to a reduced (1–2 days) pelagic larval duration (associated with temperature), resulting in reduced larval mortality, and the seasonal (summer) strengthening of the East Australian Current (EAC) transporting particles ~50 km further south. These findings demonstrate that in dynamic boundary current systems such as the EAC, the final settlement location of larvae that are transported by ocean currents can vary considerably depending on the timing and location of spawning and that multiple spawning events are important for maximum dispersal.     

March of the sculpin: measuring and predicting short‐term movement of banded sculpin Cottus carolinae

Movement of fishes through space and time is critical for population regulation and community structuring, but the dispersal of many benthic stream fishes remains unstudied. We used passive integrated transponders to track the short‐term dispersal of 51 banded sculpin Cottus carolinae throughout a 600 m reach of Little Creek in central Tennessee during April and May. Our objectives were to assess the efficacy of recently developed dispersal models, evaluate temporal variability in movement and determine whether individuals switched between stationary and mobile movement behaviours. Observed movement distances did not differ from modelled leptokurtic dispersal kernels estimated using the fishmove package in the R Statistical Environment for 12 of 13 recapture occasions. Leptokurtic dispersal kernel parameters including the mobile component (σ_mob) and shared stationary component (p) were temporally dynamic and differed from static median values reported for fishes in fishmove, while the more abundant stationary component (σ_stat) showed agreement with fishmove. The recapture occasion during which model predictions were not validated was associated with a large flow pulse that stimulated increased movement at the population scale. At the individual scale, 28 of 51 fish switched between stationary and mobile dispersal behaviour and the frequency distribution of switches was leptokurtic. Collectively, our findings reveal an emergent property characterised by consistent upstream movement of banded sculpin despite variability in population‐scale responses to flow and individual‐scale switches in movement behaviour. This paradox represents the march of the sculpin, in which fish diffusively spread upstream at a constant rate despite multiscale variability in movement behaviours.     

Physiological condition and migratory experience affect fitness‐related outcomes in adult female sockeye salmon

Relating fish physiology, behaviour and experience to fitness‐related outcomes at the individual scale is ecologically significant, but presents difficulties for free‐ranging fishes in natural systems. Physiological state (e.g. level of stress or maturity) and experience (e.g. habitat use or exposure to stressors) may alter the probability of survival or reproduction. This study examined the relative influence of physiology and migratory experience on survival, migration duration, reproductive longevity, and egg retention in adult female sockeye salmon (Oncorhynchus nerka) from a Fraser River population. One hundred and thirty‐five females were plasma sampled and tagged with radio transmitters and archival temperature loggers. Fish were tracked 55 km through two natal lakes to spawning grounds, following passage of a hydroelectric dam. For 39 females, we assessed the proportion of time within an optimal temperature (T_optAS) window (13.4–19.5°C), which provides ≥90% of maximum aerobic scope. Females with lower plasma glucose concentrations were more likely to reach spawning grounds. Early migrants spent longer in natal lakes. More time in the T_optAS window was associated with greater reproductive longevity and lower probability of egg retention. Later arriving females had reduced longevity on spawning grounds, as did females that retained eggs. Exposure to higher dam discharge was associated with reduced reproductive longevity and greater probability of egg retention, but not lower survival, indicating a delayed effect of dam passage. Our results underscore the complexity of factors governing fitness‐related outcomes for salmonids, particularly the importance of female experience in the days and weeks prior to spawning.     

Estimating effective population size of large marine populations,is it feasible?

Sustainable exploitation of marine populations is a challenging task relying on information about their current and past abundance. Fisheries‐related data can be scarce and unreliable making them unsuitable for quantitative modelling. One fishery independent method that has attracted attention in this context consists in estimating the effective population size (N_e), a concept founded in population genetics. We reviewed recent empirical studies on N_e and carried out a simulation study to evaluate the feasibility of estimating N_e in large fish populations with the currently available methods. The detailed review of 26 studies found that published empirical N_e values were very similar despite differences in species and total population sizes (N). Genetic simulations for an age‐structured fish population were carried out for a range of population and samples sizes, and N_e was estimated using the Linkage Disequilibrium method. The results showed that already for medium‐sized populations (1 million individuals) and common sample sizes (50 individuals), negative estimates were likely to occur which for real applications is commonly interpreted as indicating very large (infinite) N_e. Moreover, on average, N_e estimates were negatively biased. The simulations further indicated that around 1% of the total number of individuals might have to be sampled to ensure sufficiently precise estimates of N_e. For large marine populations, this implies rather large samples (several thousands to millions of individuals). If however such large samples were to be collected, many more population parameters than only N_e could be estimated.     

Parentage Determination and Effective Population Size Estimation in Mass Spawning Pacific Oyster,Crassostrea gigas,Based on Microsatellite Analysis

Seven high polymorphic microsatellite loci were used to determine the pedigrees in a mass spawning of Pacific oyster, Crassostrea gigas, and to estimate the genetic variability between broodstock and offspring. Parental assignment was performed on a total of 155 individuals, including 141 offspring, 8 candidate mothers, and 6 candidate fathers. The assignment results of real offspring were generally in agreement with simulation with a success rate over 99% using only six of these loci. The allelic diversity and observed heterozygosity (H_o) exhibited similarity between parents and offspring populations, but the expected heterozygosity (H_e) had a significant decrease in offspring. Although all the males and females contributed to the next generation, the variances of reproductive success and unequal sex ratio resulted in a decline in effective population size (N_e = 11.42). The inbreeding rate of this small‐scale, mass spawning population was estimated at approximately 16.5% per generation. This gave us an insight that when designing breeding programs based on mass spawning for future oyster cultivation generations, the higher inbreeding and lower effective population size should be considered.     

Genetic estimates of population structure and dispersal in a benthic stream fish

Abstract–  Comparing genetic and demographic estimates of dispersal in freshwater fish can improve understanding of movement distributions and population connectivity. Here we examined genetic variation among mottled sculpin (Cottus bairdi) in the Nantahala River (North Carolina, USA) to compare genetic estimates of dispersal with estimates derived from mark–recapture studies of individual movement. Microsatellite‐based analysis of gene flow revealed evidence of strong isolation by distance among locations spanning only 5.6 km and limited dispersal among clusters of sites separated by swift cascades. Estimates of between‐cluster contemporary dispersal rates derived from Bayesian assignment tests ranged from 1% to 6%, with most movement occurring among adjacent clusters in a downstream direction. Evidence of a long‐term net immigration asymmetry and decreasing genetic diversity from downstream to upstream locations indicates that historical patterns of stream colonisation contrast with contemporary dispersal patterns. Our findings are largely consistent with predictions from individual movement patterns but suggest that long moves (>500 m) are more frequent, and maximum dispersal distances are greater than what has been reported from mark–recapture studies. The discrepancy may reflect spatial limitations of mark–recapture methods or temporal variation in dispersal in individuals and populations.     

Riverine spawning,long distance larval drift,and floodplain recruitment of a pelagophilic fish: A case study of golden perch (Macquaria ambigua) in the arid Darling River,Australia

1. Pelagic spawning riverine fish (pelagophils) spawn in free‐flowing river habitats with downstream drift of eggs and larvae but the spatial scale is often unknown, and this constitutes a major ecological knowledge gap.
2. In the arid Darling River in south‐eastern Australia, the present objectives were: (i) to determine the potential downstream dispersal distance of young golden perch (Macquaria ambigua); and (ii) to evaluate whether provision of environmental water enhanced dispersal of young fish from Menindee Lakes to the lower Darling River (LDR) while also cueing further spawning in downstream lotic reaches.
3. Golden perch spawned in unregulated lotic tributaries on a flood pulse and larvae drifted or dispersed >1,600 km downstream and entered large ephemeral productive floodplain lake nursery habitats as fully scaled fingerlings.
4. Planned releases of environmental water cued golden perch spawning in the LDR and enabled juvenile fish to disperse downstream from the Menindee Lakes nursery into receiving populations in the LDR, Great Darling Anabranch, and southern Murray River, with some fish potentially completing an active migration of >2,100 km by age 1 year.
5. The Darling River case study highlights the need for a system‐scale approach to the conservation management of pelagophilic fish, along with multi‐year perennial flow strategies to improve ecosystem integrity in large rivers globally.

     

Success of a large riverine cyprinid smallmouth yellowfish Labeobarbus aeneus in a southern African impoundment

Rheophilic cyprinids have been detrimentally affected by impoundments, which alter flow regimes, the physical environment and block migration routes. Initial evidence for a southern African cyprinid, smallmouth yellowfish Labeobarbus aeneus, suggested that dependence on riverine spawning areas limits their success within the turbid impoundments of the Orange River system. However, well‐established impoundment populations have since been documented. A comprehensive assessment of L. aeneus biology and population dynamics was undertaken to understand the drivers of successful colonisation in Lake Gariep, South Africa's largest impoundment. Labeobarbus aeneus exhibited slow growth, delayed maturity and longevity, most likely resulting in slow population growth rates. Highest adult and juvenile abundances were observed >100 km from the inflowing Orange River during the peak and post‐spawning seasons, respectively, suggesting that the species displays high ecological plasticity and may not be dependent on inflowing rivers for spawning. Meta‐analyses of life history parameters indicated consistency among riverine and lacustrine populations. Although experiencing an initial lag phase, L. aeneus has successfully adapted to the lacustrine environment.     

Bonga shad (Ethmalosa fimbriata) spawning tactics in an upwelling environment

Successful recruitment in small pelagic fish populations inhabiting upwelling zones is subject to variation in fecundity and is driven by spatial and temporal fluctuations in environmental conditions, that is, mainly sea surface temperature, salinity and food availability. These fluctuations in abiotic factors have stimulated small pelagic fish populations to exhibit specifically adapted spawning tactics. To better understand to what extent a short‐lived exploited fish species such as bonga shad Ethmalosa fimbriata has adapted to an upwelling environment, we have investigated the interrelationship between upwelling intensity as a proxy for productivity and population fecundity by means of a virtual population analysis. We found that females of intermediate size contributed significantly more eggs to the population's fecundity than smaller or larger ones. Our model results further indicate that E. fimbriata exhibits a spawning preference at water temperatures of around 25°C and upwelling intensities of around 2.5 m³ s^?1 m^?1. Hence, we hypothesize that climate change‐driven increases in sea temperatures and modifications of upwelling‐favourable winds could significantly impact the species’ reproductive biology. To understand how climate change might impact fisheries, spawning tactics of small pelagic fishes are important to assess as well as their recruitment success. Such information is particularly relevant in countries where the fishery is critical at socio‐economic level, to better implement fisheries management addressing multiple stressors.     

Dispersal of Eastern King Prawn larvae in a western boundary current: New insights from particle tracking

Patterns in larval transport of coastal species have important implications for species connectivity, conservation, and fisheries, especially in the vicinity of a strengthening boundary current. An Ocean General Circulation Model for the Earth Simulator particle tracking model was used to assess the potential dispersal of Eastern King Prawn (EKP) larvae Melicertus (Penaeus) plebejus, an important commercial and recreational species in Eastern Australia. Particles were exposed to a constant natural mortality rate, and temperature‐dependent growth (degree‐days) was used to determine the time of settlement. Forward and backward simulations were used to identify the extent of larval dispersal from key source locations, and to determine the putative spawning regions for four settlement sites. The mean dispersal distance for larvae was extensive (~750–1,000 km before settlement), yet the northern spawning locations were unlikely to contribute larvae to the most southern extent of the EKP range. There was generally great offshore dispersal of larvae, with only 2%–5% of larvae on the continental shelf at the time of settlement. Our particle tracking results were combined with existing site‐specific reproductive potentials to identify the relative contributions of larvae from key source locations. Although mid‐latitude sites had only moderate reproductive potential, they delivered the most particles to the southern coast and are probably the most important sources of larval EKP for the two southern estuaries. Our modelling suggests that mesoscale oceanography is a strong determinant of recruitment success of the EKP, and highlights the importance of both larval dispersal and reproductive potential for understanding connectivity across a species’ range.     

Is Guam a regional source,destination, or stepping‐stone for larvae of three fisheries species?

The pelagic larval duration (PLD) period of fish can influence dispersal, recruitment, and population connectivity, thereby potentially informing best strategies for fisheries management. Computer models were used to simulate the dispersal of larvae of three species, representing a range of PLDs, from the Pacific island of Guam and neighboring islands for a 9‐year period (2004–2012) to gain insight into the best management strategies for these species. The species included two springtime spawners with shorter and longer PLDs, scribbled rabbitfish (Siganus spinus; 33 days) and yellowfin goatfish (Mulloidichthys flavolineatus; ~90 days maximum), and a fall spawner with a similarly long PLD, bluespine unicornfish (Naso unicornis; ~94 days maximum). An ocean circulation model coupled with a particle dispersal model provided simulated numbers of larvae settling at each island in relation to the island where they were spawned. Graph analysis was used to examine generational connections between islands. For S. spinus, self‐seeding was the dominant means of replenishment at Guam. Local management actions to maintain adequate spawning stock may be a primary control on long‐term sustainability for that fishery. In contrast, N. unicornis and M. flavolineatus populations at Guam were reliant on outside sources for 92%–98% of larval supply. For them, identifying and negotiating the preservation of upstream spawning potential in the Marshall Islands and Federated States of Micronesia will be needed. Guam played a relatively minor role in generational connectivity across the region. Shortest paths spanning the region often did not pass through Guam, or there were equally short paths through other islands.