Pelagic Larval Duration and Population Structure of a Widely Distributed Amphidromous Fish in the Indo‐Pacific Region

Pelagic larval duration (PLD) is a commonly used proxy for dispersal potential in coral reef fishes. Here we examine the relationship between PLD, genetic structure and genetic variability in geographically widespread and ecological generalist species from one coral reef fish family (Pomacentridae) that differs in mean larval duration by more than a month. The genetic structure was estimated in eight species using a mitochondrial molecular marker (D-loop) and in a sub-set of five species using nuclear molecular markers (ISSRs). Estimates of genetic differentiation were similar among species with pelagic larvae, but differed between molecular markers. The mtDNA indicated no structure in all species except one, while the ISSR indicated some structure between the sampling locations in all species. We detected a relationship between PLD and genetic structure using both markers. These relationships, however, were caused by a single species, Acanthochromis polyacanthus, which differs from all the other species examined here in lacking a larval phase. With this species excluded, there was no relationship between PLD and genetic structure using either marker despite a range of PLDs of more than 20 days. Genetic diversities were generally high in all species and did not differ significantly among species and locations. Nucleotide diversity and total heterozygosity were negatively related to maximum PLD but again these relationships were caused by A. polyacanthus and disappeared when this species was excluded. These genetic patterns are consistent with moderate gene flow among well-connected locations and indicate that at this phylogenetic level (i.e., within family) the duration of the pelagic larval phase is unrelated to the patterns of genetic differentiation.

Mennesson, Marion Isabelle, Tabouret, Hélène, Pécheyran, Christophe, Feunteun, Eric, Keith, Philippe (2015): Amphidromous life cycle of Eleotris fusca (Gobioidei : Eleotridae), a widespread species from the Indo-Pacific studied by otolith analyses. Cybium 39 (4): 249-260, DOI: 10.26028/cybium/2015-394-002, URL: http://dx.doi.org/10.26028/cybium/2015-394-002

The relationship between pelagic larval duration (PLD) and population connectivity in marine fishes has been controversial, but most studies to date have focused on tropical taxa. Here, we examine PLD in 11 species of triplefin fishes from a temperate environment in the Hauraki Gulf, New Zealand, to describe daily increment patterns and settlement marks in the otoliths. The formation of daily increments was validated using larvae of known age and tetracycline marking of settled juveniles. Settlement mark identity was verified by comparing total increment counts from otoliths of recently settled fishes with PLD counts from post-settlement fishes. A similar pattern of three groups of increments across the otolith was found in all specimens examined. The settlement mark was similar in all species and occurred as a sharp drop in increment width within the area of transition in optical density. PLD was lengthy, compared to species of triplefins from elsewhere, and ranged between 54.4 ± 1.7 SE days in Bellapiscis lesleyae to 86.4 ± 2.6 SE days in Forsterygion malcolmi. Variation in PLD within species was high but did not mask interspecific differences. PLD was not phylogenetically constrained, as sister species differed significantly in PLD. PLD was compared with genetic population connectivity for eight of the study species using mitochondrial gene flow data from Hickey, Lavery, Hannan, Baker, Clements. Mol Ecol 18:680–696 (2009). The observed lack of correlation between PLD and gene flow suggests that dispersal is limited by other factors, such as larval behaviour and the availability of settlement habitat.

Understanding connectivity remains a fundamental challenge to marine ecology due to technical limitations of tracking larval dispersal. Marine population genetic analyses are often used to make inferences about the scale of population connectivity. For species with a larval phase, pelagic larval duration (PLD) is assumed to influence the scale of connectivity. If PLD and genetic metrics are reliable proxies of connectivity, the 2 should be well correlated. Previous tests report conflicting results, with many reports that global FST (Wright's fixation index) correlates poorly with PLD, and one very high correlation of isolation-by-distance (IBD) slope, which is derived from FST, with PLD. First we clarify the expectations for the performance of these different proxies in light of the latest understanding of larval dispersal dynamics. We then test the hypothesis that IBD slope may be a more robust correlate with dispersal scale than global FST with a new dataset of recent marine genetic studies. Re-evaluation of previously published and new datasets revealed a consistent, moderate fit (R 2 ~0.30) between genetic and PLD proxies of dispersal (using either IBD slope or global FST), with significant improvement for small-scale (<650 km) studies (R 2 = 0.50), and important effects of marker type. Significant effects of number of individuals and number of populations sampled on the genetic metrics in our dataset suggest a common need for more robust sampling designs. These results syn- chronize previous studies on this topic and provide validation that PLD and genetic metrics typically reflect scales of dispersal, as intended, at least when sampling design is robust.

Increasing dispersal duration should result in increasing dispersal distance, facilitating higher gene flow among populations. As such, it has long been predicted that genetic structure (e.g. F(ST) ) among populations of marine species should be strongly correlated with pelagic larval duration (PLD). However, previous studies have repeatedly shown a surprisingly poor correspondence. This result has been frequently interpreted as evidence for larval behaviours or physical oceanographic processes that result in larvae failing to reach their dispersal potential, or error inherent in estimating PLD and F(ST) . This study employed a computer modelling approach to explore the impacts of various uncertainties on the correlation between measures of genetic differentiation such as F(ST) and PLD. Results indicate that variation resulting from PLD estimation error had minor impacts on the correlation between genetic structure and PLD. However, variation in effective population size between species, errors in F(ST) estimation and non-equilibrium F(ST) values all had major impacts, resulting in dramatically weaker correlations between PLD and F(ST) . These results suggest that poor correlations between PLD and F(ST) may result from variation and uncertainty in the terms associated with the calculation of F(ST) values. As such, PLD may be a much stronger determinant of realized larval dispersal than suggested by the weak-to-moderate correlations between PLD and F(ST) reported in empirical studies.

Duration of the pelagic phase of benthic marine fishes has been related to dispersal distance, with longer pelagic larval duration (PLD) expected to result in greater dispersal potential. Here, we examine PLDs of 2 species of coral-reef butterflyfish (Chaetodon auriga and C. flavirostris) across latitudes (14°S–37°S) along the Great Barrier Reef into south-eastern Australia; we predict that PLD will be higher for fish collected below the breeding latitudes of 24°S. For C. auriga, apart from significantly longer PLDs at Lord Howe Island and Jervis Bay (means of 54 and 52 days, respectively), all locations had similar PLDs (mean 41 days). For C. flavirostris, there was no significant location effect on PLD (mean 41.5 days); however, PLD at Lord Howe Island was 58 days with high variance precluding significance. Also, there was no significant variation in PLD among years for either species despite considerable variation in East Australian Current strength.

Migratory fishes are natural wonders. For many people, the term migratory fish evokes images of salmon audaciously jumping at waterfalls as they return to their own riverine birthplace to spawn after years of growth in the ocean, but freshwater fishes actually show a broad spectrum of migration strategies. Migratory fishes include small species – three-spined sticklebacks that spawn in coastal streams around the northern Pacific and gobies that move from the ocean into tropical island streams by climbing waterfalls (McDowall, 1988) – as well as some of the largest freshwater fishes in the world, such as the Mekong dog-eating catfish and the Chinese paddlefish (Stone, 2007). Aside from migratory habits, these species have few shared characteristics; they encompass numerous evolutionary lineages, enormous differences in life history, and every possible direction and distance of migration. Biologists treat migratory freshwater fishes as a functional group because their life-history strategy revolves around long-distance movement between ecosystems in a perilous quest to take advantage of both high-quality breeding sites and bountiful feeding areas. As humans have physically blocked fish migrations, degraded breeding and feeding grounds and relentlessly harvested migrants for their flesh and roe, many populations have declined or been extirpated. This chapter will provide an overview of fundamental and applied research that is helping to guide efforts to conserve migratory freshwater fishes. For practical purposes, we define migratory behaviour as the synchronized movement of a substantial proportion of a population between distinct habitats, which is repeated through time within or across generations. Modern definitions of fish migrations typically recognise both the adaptive benefits of migrating and individual variation in executing the general strategy (see McDowall, 1988; Lucas & Baras, 2001). Not every individual must move, the timing may vary somewhat from year to year, and the motive for migrating may include seeking refuge from harsh conditions in addition to breeding and feeding. Nonetheless, in most cases, migration is critical to individual fitness and population persistence because it enables specialised use of different habitats for growth and reproduction. Where their migration routes are blocked or key habitats are lost, migratory fishes often suffer rapid and catastrophic losses. Human appropriation and degradation of the Earth's freshwater ecosystems (Vorosmarty et al. , 2010; Carpenter et al. , 2011) have transformed this reliance on multiple habitats into a detriment for many migratory fishes.

The giant clam subfamily Tridacninae (family Cardiidae) is an important group of bivalve molluscs found throughout the Red Sea and Indo-Pacific, from East Africa to the Eastern Pacific biogeographic region. The Tridacna genus is currently revised with numerous cryptic species identified with molecular markers. New Tridacna records from the fringe of the known distribution areas are extremely useful to identify genetically unique species, geographic ranges, and to examine processes associated with species differentiation. While Tridacna maxima is abundant in French Polynesia (Central South Pacific Ocean) the larger fluted giant clam Tridacna squamosa was formerly reported only in the Austral Islands in the south. Following a recent survey that spanned 23 islands and atolls of the Society, Tuamotu and Gambier Archipelagos, the presence of T. squamosa between the Cook Islands and Pitcairn Islands is confirmed using both morphological and molecular information, suggesting a relic distribution across the Central Pacific Ocean. Tridacna squamosa is rare, but present throughout Tuamotu and Gambier. However, it remained undetected from the Society Islands, probably due to historical over-fishing. This species is valued by local inhabitants, and is sought after mainly as gifts and also for a limited local shell trade. The rarity of T. squamosa may call for conservation measures in the near future.

Chromosomal rearrangements associated with pelagic larval duration in Labridae (Perciformes)

Diadromous fish species in the family Kuhliidae are able to colonise freshwater systems in Indo-Pacific islands, but their life cycle and the mechanisms involved in the colonisation of such ecosystems are poorly documented. After validating the daily rate of increment deposition in otoliths of Kuhlia rupestris, we estimated the pelagic larval duration (PLD) of K. rupestris, widely distributed in the Indo-Pacific area, and K. sauvagii, endemic to the Indian Ocean. Median PLD of K. rupestris was significantly longer than that of K. sauvagii (40.6 ± 6.9 and 32.3 ± 3.4 days (± s.d.), respectively), implying that the PLD is probably one factor controlling the extent of distribution range in Kuhlia. Within K. rupestris, individuals from New Caledonia had longer PLDs than those from Réunion Island (44.3 ± 6.7 and 37.3 ± 4.7 days (± s.d.) respectively). Further research on larval migration is needed to determine whether this was due to different environmental conditions or is population-specific. Interestingly, the PLD of these Kuhlia species is shorter than the PLD of other tropical diadromous fishes. These results improve our understanding of the dispersal strategies of freshwater fauna, to colonise and persist in tropical islands.

AimTo assess how climate change may decouple the ecosystems used by a migratory fish, and how decoupling influences priorities for stream restoration.LocationAustralia.MethodsWe modelled changes in habitat suitability under climate change in both riverine and marine habitats for a threatened diadromous species, the Australian Grayling Prototroctes maraena, using niche models. The loss of riverine habitats for Grayling was compared with or without considering the impact of climate change on adjacent marine habitats. We also asked whether considering marine climate change changed the locations where removing dams had the greatest benefit for Grayling conservation.ResultsClimate change is expected to cause local extinction in both marine and river habitats regardless of whether dams are retained or removed at the trailing edge of the Grayling's range (north‐eastern). Decoupling of habitats was most apparent in the eastern and south‐eastern portion of the Grayling's range, where ocean warming may cause a decline in the suitability of marine habitats for larvae, while many freshwater habitats retained suitable habitat for adults. Removing dams to restore connectivity between ocean and freshwater habitats was predicted to have the greatest benefit for Grayling in southern portions of their range. Under climate change, the priorities for barrier removal gradually shift towards dams at higher elevation because of increasing suitability of freshwater habitats at higher elevations.Main conclusionsOur study highlights the importance of assessing climate range shifts in multiple ecosystems for migratory species and can help inform priorities for stream restoration under a changing climate.

Climate change is predicted to increase ocean temperatures and influence weather patterns. Here, we examine the influence of temperature and other environmental variables on key early life traits of the coral reef damselfish, Pomacentrus moluccensis, based on ten cohorts of newly settled fish collected over 13 years from around Lizard Island (Great Barrier Reef, Australia). Pelagic larval duration (PLD), larval growth and size at settlement were estimated through otolith microstructure analysis. Multiple regression techniques were used to measure the strength of the associations between these traits and developmental temperature, rain, wind speed and solar radiation. Temperature accounted for 18.4, 26.7 and 25.0 % of the variability in PLD, growth rates and settlement size, respectively. PLDs generally declined and growth rates generally increased with increasing temperatures to ~28 °C, above which PLDs tended to increase and growth rates tended to decrease. Size at settlement did not differ between ~25 and ~28 °C, but tended to decrease with increasing temperature above ~28 °C. Together rain, wind speed and solar radiation explained 6.3, 26.3 and 33.7 % of the remaining variability in PLD, growth rates and size at settlement, respectively. Higher wind speeds were generally associated with longer PLDs. Increasing wind, high rainfall and increasing solar radiation were associated with slower growth rates and smaller sizes at settlement. Overall, results suggest that ~28 °C is likely to be a thermal optimum for larval development for this species and other environmental factors associated with climate change including rainfall, wind speed and solar radiation should be considered in predictions of effects on larval fish.

Estimates of pelagic larval duration (PLD) for 10 species of Pomacentridae and two species of Gobiidae were made. In eight of the 12 species examined, within‐population mean PLDs differed between sampling times, locations within regions and among regions. In contrast, the range of these same PLD estimates overlapped at all spatial and temporal scales examined in 11 of the 12 species, but not between regions in one species (Amphiprion melanopus). Therefore, despite tight error estimates typically associated with estimates of PLD taken from a particular population at a particular time in some taxa, the overlapping ranges in PLD reported here indicate that the length of the pelagic larval phase is a much more plastic trait than previously appreciated. Within‐species variation in PLD has considerable potential to provide further insights into the ecology and evolution of tropical reef fishes.

We examined prey availability, prey consumed, and diet energy content as sources of variation in growth of natural fall Chinook Salmon Oncorhynchus tshawytscha subyearlings rearing in riverine and reservoir habitats in the Snake River. Subyearlings in riverine habitat primarily consumed aquatic insects (e.g., Diptera, Ephemeroptera, Trichoptera), of which a high proportion was represented by adult, terrestrial forms. In the reservoir, subyearlings also consumed aquatic insects but also preyed heavily at times on nonnative lentic amphipods Corophium spp. and the mysid Neomysis mercedis, which were absent in riverine habitats. The availability of prey was typically much higher in the reservoir due to N. mercedis often composing over 90% of the biomass, but when this taxon was removed from consideration, biomass estimates were more often higher in the riverine habitat. Subyearling diets during 2009–2011were generally 17–40% higher in energy in the riverine habitat than in the reservoir. Observed growth in both length and weight were significantly higher in the riverine habitat than in the reservoir. Little is known about how temporal and spatial changes in the food web in large river landscapes influence populations of native anadromous fishes. Our results provide a glimpse of how the spread and establishment of nonnative prey species can reduce juvenile salmon growth in a large river impoundment, which in turn can affect migration timing and survival.

The relationships between pelagic larval duration (PLD) and geographic distribution patterns or population genetic structures of fishes remain obscure and highly variable among species. To further understand the early life history of the tidepool snake moray Uropterygius micropterus and the potential relationship between PLD and population genetic structure of this species, otolith microstructure and population genetics based on concatenated mtDNA sequence (cytochrome b and cytochrome oxidase subunit I, 1,336 bp) were analyzed for 195 specimens collected from eight locations around the southern Ryukyu Islands, Taiwan, and the central Philippines. Eels with longer PLD and lower otolith growth rates were observed at relatively higher latitudes with lower water temperatures (54.6 ± 7.7 days and 1.28 ± 0.16 µm day−1 on Ishigaki Island, Japan, vs. 43.9 ± 4.9 days and 1.60 ± 0.19 µm day−1 on Badian, the Philippines), suggesting that leptocephali grew faster and had shortened pelagic periods in warmer waters. Meanwhile, the eels along the southwest coast of Taiwan had relatively longer PLD (57.9 ± 10.5 days), which might be associated with the more complex ocean current systems compared to their counterparts collected along the east coast of Taiwan (52.6 ± 8.0 days). However, the southwestern and eastern Taiwan groups had similar otolith growth rates (1.33 ± 0.19 µm day−1 vs. 1.36 ± 0.16 µm day−1). Despite the intergroup variation in PLD, genetic analysis revealed fluent gene flow among the tidepool snake morays in the study regions, implying that intraspecies PLD variation had a weak effect on genetic structure. The leptocephalus stage might have ensured the widespread gene flow among the study areas and leptocephalus growth was likely influenced by regional water temperature.