Life-history characteristics of eleotrid fishes of the western hemisphere, and perils of life in a vanishing environment

The term amphidromous was coined (Myers in Copeia 1949:89–97, 1949b) to describe diadromous life histories that include migrations, not associated with reproduction, that are between fresh and marine waters. This concept has facilitated evaluations of life cycles among a number of groups of fishes including some belonging to the family Eleotridae. Information was gathered on life history patterns of eleotrid fish species that have been recorded from both fresh and brackish or marine waters of the east and west coasts of North, Middle, and South America, including adjacent islands, seeking evidence of diadromy and especially of amphidromy. Convincing evidence was found of diadromy in the life cycles of four species from the east coast (with another two species possibly in this category), and of three species from the west coast of the Americas and adjacent islands. However, there was convincing evidence of amphidromy in only one species from the east coast and adjacent islands, with another three species or species populations possibly in this category, and in three species from the west coast and adjacent islands. It seems possible from the available information that there may be variation among populations in life cycles of some of these species. However, it remains for the use of modern techniques to allow more definitive determinations of life history patterns of these eleotrid species populations. Serious concern exists with respect to the conservation of all diadromous species because of the worldwide emphasis on river manipulations, especially of dam construction. Resident diadromous organisms are being impeded in their migrations, and thus imperiled in their potential for survival, depending on the nature of river alterations and the abilities of the organisms to cope with them.     

A recent marine ancestry for diadromous fishes? Sometimes yes,but mostly no!

Synopsis Diadromy in its three forms (anadromy, catadromy and amphidromy) necessarily involves fish spending part of their lives in the sea. It is not uncommon contention that diadromous fish have a recent marine ancestry and that they are in the process of moving from a marine to a freshwater lifestyle. However, analysis of the distribution of diadromy in fishes suggests that, for most species, it is an ancient life history style though, for a few, a recent marine ancestry seems plausible. In general, there is no reason to consider diadromy anything but a stable and successful life history style, and no reason to regard it as transitional between marine and freshwater habitats.Invited Editorial     

The evolutionary origins of diadromy inferred from a time-calibrated phylogeny for Clupeiformes (herring and allies)

One of the most remarkable types of migration found in animals is diadromy, a life-history behaviour in which individuals move between oceans and freshwater habitats for feeding and reproduction. Diadromous fishes include iconic species such as salmon, eels and shad, and have long fascinated biologists because they undergo extraordinary physiological and behavioural modifications to survive in very different habitats. However, the evolutionary origins of diadromy remain poorly understood. Here, we examine the widely accepted productivity hypothesis, which states that differences in productivity between marine and freshwater biomes determine the origins of the different modes of diadromy. Specifically, the productivity hypothesis predicts that anadromous lineages should evolve in temperate areas from freshwater ancestors and catadromous lineages should evolve in tropical areas from marine ancestors. To test this, we generated a time-calibrated phylogeny for Clupeiformes (herrings, anchovies, sardines and allies), an ecologically and economically important group that includes high diversity of diadromous species. Our results do not support the productivity hypothesis. Instead we find that the different modes of diadromy do not have predictable ancestry based on latitude, and that predation, competition and geological history may be at least as important as productivity in determining the origins of diadromy.     

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.     

Tropical and temperate freshwater amphidromy: a comparison between life history characteristics of Sicydiinae,ayu, sculpins and galaxiids

Amphidromy is a distinctive form of diadromy, but differences in the life histories of tropical and temperate amphidromous fishes suggest that there are two types of freshwater amphidromy. The life histories of Sicydiinae gobies, ayu (Plecoglossus altivelis), Japanese sculpins (Cottus) and galaxiids (Galaxiidae), suggest that the Sicydiinae are representatives of tropical freshwater amphidromy, whereas ayu, sculpins and galaxiids are representatives of temperate freshwater amphidromy. The Sicydiine larval stage may be required to occur in the ocean for all species, but ayu, sculpins and galaxiids have landlocked or fluvial forms with larvae that do not need to enter the ocean for larval feeding and growth. This suggests that Sicydiine larvae have a high oceanic dependency whereas ayu, sculpins and galaxiid larvae have a low oceanic dependency. Freshwater amphidromous fish in tropical and temperate zones appear to have developed two different strategies in the evolution of their life histories. It is likely that the evolutionary direction of the larval stage of tropical amphidromy is to remain in the sea and that of temperate amphidromy is towards having the ability to remain in freshwater if needed. Tropical and temperate amphidromy appear to be biologically informative categories and evaluations of this hypothesis will facilitate better understanding of the various forms of amphidromy in the future.     

Fighting the flow: downstream–upstream linkages in the ecology of diadromous fish faunas in West Coast New Zealand rivers

1. Diadromy is a dominating behavioural characteristic of fish faunas in New Zealand rivers, with amphidromy and catadromy being the most common strategies.
2. Juvenile life stages of amphidromous and catadromous species migrate from the sea, through river systems, to find habitats for feeding, growth, maturation and reproduction.
3. Studies of fish distributions in rivers of the West Coast of the South Island of New Zealand show that, in most species, these migrations result in more or less continuous distributions from the lower reaches to the upstream limits of each species' range.
4. Upstream penetration of rivers varies widely between species and this generates downstream–upstream trajectories of declining species richness.
5. Parallel trajectories of declining downstream–upstream abundance are likely in each species.
6. These patterns demonstrate the presence of downstream–upstream linkages in the community ecology of freshwater fishes in New Zealand rivers.     

Potential interactions between diadromous fishes of U.K. conservation importance and the electromagnetic fields and subsea noise from marine renewable energy developments

The considerable extent of construction and operation of marine renewable energy developments (MRED) within U.K. and adjacent waters will lead, among other things, to the emission of electromagnetic fields (EMF) and subsea sounds into the marine environment. Migratory fishes that respond to natural environmental cues, such as the Earth's geomagnetic field or underwater sounds, move through the same waters that the MRED occupy, thereby raising the question of whether there are any effects of MRED on migratory fishes. Diadromous species, such as the Salmonidae and Anguillidae, which undertake large-scale migrations through coastal and offshore waters, are already significantly affected by other human activities leading to national and international conservation efforts to manage any existing threats and to minimize future concerns, including the potential effect of MRED. Here, the current state of knowledge with regard to the potential for diadromous fishes of U.K. conservation importance to be affected by MRED is reviewed. The information on which to base the review was found to be limited with respect to all aspects of these fishes' migratory behaviour and activity, especially with regards to MRED deployment, making it difficult to establish cause and effect relationships. The main findings, however, were that diadromous species can use the Earth's magnetic field for orientation and direction finding during migrations. Juveniles of anadromous brown trout (sea trout) Salmo trutta and close relatives of S. trutta respond to both the Earth's magnetic field and artificial magnetic fields. Current knowledge suggests that EMFs from subsea cables may interact with migrating Anguilla sp. (and possibly other diadromous fishes) if their movement routes take them over the cables, particularly in shallow water (<20 m). The only known effect is a temporary change in swimming direction. Whether this will represent a biologically significant effect, for example delayed migration, cannot yet be determined. Diadromous fishes are likely to encounter EMFs from subsea cables either during the adult movement phases of life or their early life stages during migration within shallow, coastal waters adjacent to natal rivers. The underwater sound from MRED devices has not been fully characterized to determine its acoustic properties and propagation through the coastal waters. MRED that require pile driving during construction appear to be the most relevant to consider. In the absence of a clear understanding of their response to underwater sound, the specific effects on migratory species of conservation concern remain very difficult to determine in relation to MRED. Based on the studies reviewed, it is suggested that fishes that receive high intensity sound in close proximity to construction may be physiologically affected to some degree, whereas those at farther distances, potentially up to several km, may exhibit behaviour responses; the effect of which is unknown and will be dependent on the properties of the received sound and receptor characteristics and condition. Whether there are behavioural effects on the fishes during operation is unknown but any change to the environment and subsequent response by the fishes would need to be considered over the lifetime of the MRED. It is not yet possible to determine if effects relating to sound exposure are biologically significant. The current assumptions of limited effects are built on an incomplete understanding of how the species move around their environment and interact with natural and anthropogenic EMFs and subsea sound. A number of important knowledge gaps exist, principally whether migratory fish species on the whole respond to the EMF and the sound associated with MRED. Future research should address the principal gaps before assuming that any effect on diadromous species results in a biological effect.     

Cladogenesis and loss of the marine life-history phase in freshwater galaxiid fishes (Osmeriformes: Galaxiidae)

Switches from migratory (diadromous) to nonmigratory (freshwater) life histories are known to have occurred repeatedly in some aquatic taxa. However, the significance of the loss of diadromy as an initiator for speciation remains poorly understood. The rivers of New Zealand's South Island house a species flock of recently derived nonmigratory galaxiid fishes known as the Galaxias vulgaris complex. Members of this complex are morphologically and genetically similar to the diadromous G. brevipinnis found in New Zealand and southeastern Australia. We hypothesised that South Island's G. vulgaris complex (at least 10 nonmigratory lineages) represents a number of independent radiations from a migratory G. brevipinnis stock, with repeated loss of diadromy. Sequence data were obtained for 31 ingroup samples (G. vulgaris complex and G. brevipinnis) plus four outgroup taxa. A well-resolved phylogeny based on 5039 base pairs of the mitochondrial genome suggests that diadromy has been lost on three separate occasions. Thus, speciation in these galaxiid fishes is partly an incidental phenomenon caused by switches from diadromous to nonmigratory strategies. However, much of the subsequent nonmigratory diversity is monophyletic, suggesting that drainage evolution (vicariance) has also played a major role in cladogenesis. Levels of sequence divergence among major ingroup lineages (1.6-12.7%) suggest that the radiation is considerably older relative to Northern Hemisphere (postglacial) complexes of salmonid, osmerid, and gasterosteid fishes. Sympatric taxa are not monophyletic, suggesting that their coexistence reflects secondary contact rather than sympatric speciation. The monophyly of New Zealand G. brevipinnis is well supported, but both mitochondrial DNA and nuclear sequences indicate that G. brevipinnis is paraphyletic on an intercontinental scale. The divergence (maximum 11.5%) between Tasmanian and New Zealand G. brevipinnis, although large, supports marine dispersal rather than vicariance as the principle biogeographic mechanism on an intercontinental scale.     

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

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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.     

Fish ecology and management of the Sepik-Ramu,New Guinea,a large contemporary tropical river basin

Synopsis The ichthyofauna of the Sepik-Ramu basin is composed of diadromous species and the freshwater derivatives of marine families. Fish species diversity, ichthyomass and fish catches are low even by Australasian standards. Three major factors have produced the depauperate ichthyofauna and restricted fishery within the basin: First, the zoogeographic origins of the ichthyofauna. Australasian freshwater fishes, being mainly derived from marine families, generally exhibit ecological characteristics that have evolved for life in estuaries, not rivers. This has led to peculiarities in river fish ecology and explains the probable low fish production from rivers in this region in general. Several important riverine trophic resources are not exploited by the Australasian freshwater ichthyofauna. The modes of reproduction amongst the Australasian freshwater ichthyofauna have limited the colonisation and exploitation of floodplain habitats. Second, Sepik-Ramu lowland habitats, especially floodplains, are very young. This has resulted in low fish species diversity in lowlands, whilst diversity at higher altitudes is equable, in comparison to river systems in southern New Guinea/ northern Australia. Third, the Sepik-Ramu lacks an estuary in sharp contrast to river systems in southern New Guinea or northern Australia. Most of the 18 families of Australasian fishes missing from the Sepik-Ramu are probably absent because of this factor alone. In particular, the Sepik-Ramu has not been colonised by any family of fishes having pelagic eggs, resulting in the loss from the fauna of the few Australasian fish taxa with high reproductive rates. Consequently, the general problems with river fish ecology in Australasia are exacerbated within the Sepik-Ramu by the particular development and morphology of the basin. Fish species diversity in the Sepik-Ramu is low, even in comparison with those taxa representative of marine families resident in rivers in nearby zoogeographic regions (S.E. Asia) whose ichthyofaunas are otherwise dominated by freshwater dispersant groups. The Sepik-Ramu ichthyofauna is considered noteworthy for what is absent, not what is present. Ichthyomass and fish production can be increased by fish species introductions whilst, in theory, biodiversity of the native fish fauna can be maintained. The directions in which ecological evaluations of proposed introductions might proceed in practice for the Sepik-Ramu are discussed but are constrained by the lack of knowledge on species interactions from other areas.     

Potential re-establishment of diadromous fish species in the River Scheldt (Belgium)

This study (2002) documents on the inland penetration off diadromous fish species into the tidal and non-tidal part of the River Scheldt and on the impact of two lock-weir complexes. Long-term trends in oxic conditions show the river is undergoing a natural recovery process, nevertheless five of ten diadromous species recorded were still restricted to the brackish part of the estuary. Despite poor conditions, five species reached the upper freshwater part of the estuary. Erratic free-flowing conditions at the tidal weir offered limited opportunities for some diadromous species to migrate into the non-tidal part of the river. Upstream migration over the second barrier is almost completely blocked. Rehabilitation schemes to restore self-sustaining populations of diadromous fish in the cross-border River Scheldt need to include the building of fish bypasses and improved wastewater treatment and habitat restoration programmes. Guest editors: S. Dufour, E. Prévost, E. Rochard & P. Williot Fish and diadromy in Europe (ecology, management, conservation)     

Life history correlates and extinction risk of capital-breeding fishes

We consider a distinction for fishes, often made for birds and reptiles, between capital-breeding and income-breeding species. Species that follow a capital-breeding strategy tend to evolve longer intervals between reproductive events and tend to have characteristics that we associate with higher extinction risk. To examine whether these ideas are relevant for fishes, we assembled life history data for fish species, including an index of extinction risk, the interval between spawning events, the degree of parental care, and whether or not the species migrates to spawn. These data were used to evaluate two hypotheses: (1) fish species with a major accessory activity to spawning (migration or parental care) spawn less often and (2) fish species that spawn less often are at greater risk of extinction. We tested these hypotheses by applying two alternative statistical methods that account for phylogenetic correlation in cross-taxon comparisons. The two methods predicted average intervals between spawning events 0.13–0.20 years longer for fishes with a major accessory activity. Both accessories, above-average parental care and spawning migration, were individually associated with longer average spawning intervals. We conclude that the capital-breeding paradigm is relevant for fishes. We also confirmed the second hypothesis, that species in higher IUCN extinction risk categories had longer average spawning intervals. Further research is needed to understand the relationship between extinction risk and spawning interval, within the broader context of life history traits and aquatic habitats. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: S. Dufour, E. Prévost, E. Rochard & P. Williot Fish and diadromy in Europe (ecology, management, conservation)     

Reconstructing labroid evolution with single-copy nuclear DNA.

Fifteen per cent of all living fishes are united in a single suborder (Labroidei) and display a dazzling array of behavioural and ecological traits. The labroids are considered monophyletic and members share a pharyngeal jaw apparatus (PJA) modified for crushing and processing prey. Outside of the explicitly functional PJA, there is no corroborative evidence for a monophyletic Labroidei. Here, we report the first molecular phylogenetic analysis of the suborder. Contrary to morphology-based phylogenies, our single-copy nuclear DNA data do not support labroid families as a natural group. Our data indicate that pharyngognathy has evolved independently among labroid families and that characters of the PJA are not reliable markers of perciform evolution. This work ''crushes'' conventional views of fish phylogeny and should engender novel concepts of piscine life history evolution.     

The origin of the salmonid fishes: marine, freshwater... or neither?

The evolutionary origins of the salmonidfishes, whether in freshwater or the sea, havebeen debated for centuries. Early viewsfavoured a group of marine ancestry invadingfreshwaters; more recently, there was a shifttowards a freshwater ancestry, on grounds thata return to freshwater to spawn indicates theancestral biome. Salmonids are widely believedto share an ancient common ancestry with thenorthern hemisphere Osmeridae and southernhemisphere Retropinnidae and Galaxiidae. Salmonidae are diadromous, as are Osmeridae,Retropinnidae and Galaxiidae. This suggeststhat diadromy is an ancient behavioralphenomenon across all these groups, that theshared common ancestry of these groups was alsodiadromous, and that the ancestry of Salmonidaewas neither marine nor freshwater, but wasamongst diadromous fishes. This begs thequestion of whether this common ancestor wasmarine or freshwater, a question for which ananswer seems likely to be elusive.     

Untested Assumptions: The Role of Canals in the Dispersal of Sea Lamprey, Alewife, and other Fishes in the Eastern United States

Canals provide aquatic organisms with an excellent thoroughfare to disperse, expand their range, and gain access to new drainages. The evidence used to support this contention is vast and dates to at least the early nineteenth century. In most cases however, the evidence used to defend the contention is not a direct observation of fish dispersing through a canal. Instead, the transfer is generally inferred after the new species is observed in a new watershed, one connected by a canal to the watershed in which the fish was already established. Often, these inferences are developed without considering aspects of the life history and behavior of the fishes involved, or the structure of the canal. I explore the historic transfer of sea lamprey, alewife and other fishes from one watershed to another in the inland waters of eastern North America, primarily New York. New York is an ideal area for examining this phenomenon as a canal has connected each of the five major drainages in the state during part of the previous 200 years. These twelve major canals, and the Welland Canal that connects lakes Ontario and Erie, bypassed major obstructions and created a continuous water route among drainages. Also, many of the 167 freshwater and diadromous fishes of New York exhibited distribution patterns historically limited to a watershed or set of neighboring watersheds. When several important cases are re-examined, dispersal through a canal is not the most parsimonious explanation for the presence of the species in a new drainage. To argue fish dispersal through navigation canals, researchers must consider the natural history and ecological requirements of the species, the characteristics and environmental conditions of the canal, and alternative explanations. The mere presence of a canal does not demonstrate that fish used the canal for dispersal.     

Three ecological factors influencing riverine fish diversity in the Shubuto River system,Japan: habitat capacity,habitat heterogeneity and immigration

A major goal of stream ecology is to identify environmental gradients that shape riverine communities. We examined the relative importance of three ecological factors that have been hypothesized to influence a longitudinal pattern of fish diversity: habitat capacity, heterogeneity and immigration of diadromous fishes. Field surveys were carried out in the entire network of the Shubuto River system, Hokkaido, Japan. A hierarchical partitioning approach revealed that distance from the sea, a proxy for immigration potential of diadromous fishes, had the greatest explanatory capacity, by which 24.9 % of variation in fish species richness was explained. Habitat capacity (approximated by catchment area) was also identified as a significant predictor of fish diversity, whereas habitat heterogeneity brought little improvement to the model performance. These results reflect the fish fauna of the Shubuto River system, in which diadromous fishes are dominant in both abundance and species richness.     

The joint evolution of the Myxozoa and their alternate hosts: A cnidarian recipe for success and vast biodiversity

The relationships between parasites and their hosts are intimate, dynamic and complex; the evolution of one is inevitably linked to the other. Despite multiple origins of parasitism in the Cnidaria, only parasites belonging to the Myxozoa are characterized by a complex life cycle, alternating between fish and invertebrate hosts, as well as by high species diversity. This inspired us to examine the history of adaptive radiations in myxozoans and their hosts by determining the degree of congruence between their phylogenies and by timing the emergence of myxozoan lineages in relation to their hosts. Recent genomic analyses suggested a common origin of Polypodium hydriforme, a cnidarian parasite of acipenseriform fishes, and the Myxozoa, and proposed fish as original hosts for both sister lineages. We demonstrate that the Myxozoa emerged long before fish populated Earth and that phylogenetic congruence with their invertebrate hosts is evident down to the most basal branches of the tree, indicating bryozoans and annelids as original hosts and challenging previous evolutionary hypotheses. We provide evidence that, following invertebrate invasion, fish hosts were acquired multiple times, leading to parallel cospeciation patterns in all major phylogenetic lineages. We identify the acquisition of vertebrate hosts that facilitate alternative transmission and dispersion strategies as reason for the distinct success of the Myxozoa, and identify massive host specification‐linked parasite diversification events. The results of this study transform our understanding of the origins and evolution of parasitism in the most basal metazoan parasites known.     

The Covert World of Fish Biofluorescence: A Phylogenetically Widespread and Phenotypically Variable Phenomenon

The discovery of fluorescent proteins has revolutionized experimental biology. Whereas the majority of fluorescent proteins have been identified from cnidarians, recently several fluorescent proteins have been isolated across the animal tree of life. Here we show that biofluorescence is not only phylogenetically widespread, but is also phenotypically variable across both cartilaginous and bony fishes, highlighting its evolutionary history and the possibility for discovery of numerous novel fluorescent proteins. Fish biofluorescence is especially common and morphologically variable in cryptically patterned coral-reef lineages. We identified 16 orders, 50 families, 105 genera, and more than 180 species of biofluorescent fishes. We have also reconstructed our current understanding of the phylogenetic distribution of biofluorescence for ray-finned fishes. The presence of yellow long-pass intraocular filters in many biofluorescent fish lineages and the substantive color vision capabilities of coral-reef fishes suggest that they are capable of detecting fluoresced light. We present species-specific emission patterns among closely related species, indicating that biofluorescence potentially functions in intraspecific communication and evidence that fluorescence can be used for camouflage. This research provides insight into the distribution, evolution, and phenotypic variability of biofluorescence in marine lineages and examines the role this variation may play.     

The evolution of migration and some life history parameters in marine fishes

Synopsis Migration is a common phenomenon in marine fishes but the consequences with respect to life history variation have been little explored. Migration both influences the evolution of other traits and is contigent upon the evolution of other behavioural and demographic characters. The interaction between such factors is illustrated by considering the relationship between the cost of migration in relation to fecundity and the advantages and disadvantages of schooling, a phenomenon hypothesized to favour the evolution of migration. These analyses predict that migratory species will be relatively large, mature late and at a relatively large size and grow relatively fast. Data from north temperate marine fish support these predictions.