Void development on carbonate coasts: creation of anchialine habitats

Anchialine caves in coastal locations develop in two ways: by pseudokarst processes that form talus caves, sea caves, tafoni, fissure caves and lava tubes, and by karst dissolutional processes that form stream caves, flank margin caves, and blue holes. Pseudokarst caves are of minor importance in anchialine cave habitat development, with some lava tubes being notable exceptions. Dissolution caves provide the most extensive, variable, and long-term environments for anchialine habitats. The Carbonate Island Karst Model (CIKM) allows dissolutional cave development in carbonate coasts to be understood as the interplay between freshwater and marine water mixing, sea-level change, rock maturity, and interaction with adjacent non-carbonate rocks. Glacioeustatic sea-level changes of the Quaternary have moved all coastal anchialine cave environments repeatedly through a vertical range of over 100 m, and modern anchialine environments could not develop at their current elevations until ~4,000 years ago when sea level reached its present position. Blue holes form by a variety of mechanisms, but the most common is upward stoping and collapse from deep dissolutional voids. As a result, they provide vertical connection between different levels of horizontal cave development produced by a variety of earlier sea-level positions. Blue holes are overprinted by successive sea-level fluctuations; each sea-level event adds complexity to the habitats within blue holes and the cave systems they connect. Blue holes can reach depths below the deepest glacioeustatic sea-level lowstand, and thereby provide a refugia for anchialine species when cave passages above are drained by Quaternary sea-level fall. Blue holes represent the most significant anchialine cave environment in the world, and may provide clues to anchialine cave species colonization and speciation events.     

Phylogeny and historical biogeography of the cave-adapted shrimp genus Typhlatya (Atyidae) in the Caribbean Sea and western Atlantic

Aim To infer phylogenetic relationships among five species of the cave‐adapted shrimp genus Typhlatya in order to test competing hypotheses of dispersal and colonization of the disjunct cave localities occupied by these five species. Location Typhlatya species are found in caves and anchialine ponds across the northern margin of the Caribbean Sea, along the Mediterranean and Adriatic coasts and on oceanic islands in the Atlantic and eastern Pacific oceans. This study focuses on five species, one from Bermuda, one from the Caicos Islands and three from the Yucatan Peninsula of Mexico. Methods Partial sequences (c. 1400 bp) from the mitochondrial cytochrome b, 16S rDNA and COI genes were obtained from representative samples of the five species. Phylogenetic inference was carried out with maximum parsimony and maximum likelihood analyses. Parsimony networks were constructed for the Bermudian species Typhlatya iliffei and one Yucatan species Typhlatya mitchelli, to determine the degree of connectivity among populations inhabiting different cave systems. Results All three land masses were recovered as monophyletic. The two insular marine species from Bermuda and the Caicos Islands formed a clade, while the three continental freshwater species from the Yucatan Peninsula formed another. Within both Bermuda and the Yucatan, shared haplotypes were found in different cave systems, suggesting recent or ongoing gene flow among populations in both locales. Main conclusions The two insular marine Typhlatya species originated from an ancestral marine population, possibly already cave‐adapted, that is suggested to have colonized the Caicos Islands and subsequently dispersed to Bermuda via the Gulf Stream. Divergence estimates suggest that colonization occurred before the formation of present‐day anchialine cave habitat, which did not form on either island until the late Pliocene to early Pleistocene. Divergence estimates also indicate that the Yucatan freshwater species split before the formation of freshwater cave habitat in the Yucatan. These species could have inhabited crevicular marine habitats before the late Pliocene/early Pleistocene in the Yucatan or elsewhere in the Caribbean, and subsequently migrated to freshwater caves once they formed.     

A new species of Typhlatya (Crustacea: Decapoda: Atyidae) from anchialine caves on the French Mediterranean coast

A new species of the thermophylic Tethyan relict prawn Typhlatya is described from two anchialine caves near Perpignan (southern France). The new species is closely related to a congener known only from a freshwater cave at Castellón (eastern Spain), about 400 km to the south-west, differing apparently only in the size and shape of the rostrum and the armature of the dactylus of the fifth pereiopod. Based on palaeogeographical evidence and assuming a sister-group relationship between both species, we suggest that their common ancestor could not be older than early Pliocene in age, and that it was already a stygobiont taxon adapted to live in shallow-water marine crevicular habitats. This ancestor would have vanished from the western Mediterranean after the cooling associated with the onset of northern Hemisphere glaciation, about 3 Mya, as documented for other Mediterranean marine taxa. Indeed, the genus is completely stygobiont and does not occur in fluvial environments. The Pyrenees represent a watershed boundary that eliminates the possibility of the derivation of one species from the other by active dispersal after establishment in continental waters.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 387–414.     

Search for Bermuda’s deep water caves

The mid-Atlantic islands of Bermuda harbor one of the richest and most diverse anchialine communities known from anywhere on Earth. However, all known anchialine caves in Bermuda (maximum depth—26 m) were dry during the last glacial period extending from approximately 9,000 to 115,000 years ago when glacial sea levels were as much as 127 m lower. Since it is highly unlikely that Bermuda’s endemic cave species evolved since the caves were flooded by sea level rise, alternate deeper habitats must have existed to shelter anchiane fauna for prolonged periods of lower sea level during the Pleistocene. In order to systematically search for such now deep water cave habitats, high-resolution multibeam sonar and remotely operated vehicles were used to map and explore the seafloor off Bermuda in 60–200 m depths along the outer shelf break edge of the submarine escarpment surrounding the Bermuda Platform and an adjacent seamount. Specific goals were to discover deep water cave and/or crevicular habitats and to characterize the nature, geological stratification and composition, and sea level history of the platform margin, in particular focusing on features directly relating to Pleistocene low sea stand events. During this sea floor survey, clearly defined paleo-shoreline features generated by wave and current erosion were found to encircle the Bermuda seamount and Challenger Bank at 60 and 120 m depths.     

Local adaptation despite high gene flow in the waterfall‐climbing Hawaiian goby,Sicyopterus stimpsoni

Environmental heterogeneity can promote the emergence of locally adapted phenotypes among subpopulations of a species, whereas gene flow can result in phenotypic and genotypic homogenization. For organisms like amphidromous fishes that change habitats during their life history, the balance between selection and migration can shift through ontogeny, making the likelihood of local adaptation difficult to predict. In Hawaiian waterfall‐climbing gobies, it has been hypothesized that larval mixing during oceanic dispersal counters local adaptation to contrasting topographic features of streams, like slope gradient, that can select for predator avoidance or climbing ability in juvenile recruits. To test this hypothesis, we used morphological traits and neutral genetic markers to compare phenotypic and genotypic distributions in recruiting juveniles and adult subpopulations of the waterfall‐climbing amphidromous goby, Sicyopterus stimpsoni, from the islands of Hawai'i and Kaua'i. We found that body shape is significantly different between adult subpopulations from streams with contrasting slopes and that trait divergence in recruiting juveniles tracked stream topography more so than morphological measures of adult subpopulation differentiation. Although no evidence of population genetic differentiation was observed among adult subpopulations, we observed low but significant levels of spatially and temporally variable genetic differentiation among juvenile cohorts, which correlated with morphological divergence. Such a pattern of genetic differentiation is consistent with chaotic genetic patchiness arising from variable sources of recruits to different streams. Thus, at least in S. stimpsoni, the combination of variation in settlement cohorts in space and time coupled with strong postsettlement selection on juveniles as they migrate upstream to adult habitats provides the opportunity for morphological adaptation to local stream environments despite high gene flow.     

Diurnal dynamics of the microbial loop in peatlands: structure, function and relationship to environmental parameters

Globally, introductions of alien species are increasingly common, with invasive predators potentially having detrimental effects via predation on native species. However, native prey may avoid predation by adopting new behaviors. To determine whether invasive fish populations consume endemic shrimp in invaded Hawaiian anchialine habitats or if adopted patterns of diel migration prevents predation as previously hypothesized, a total of 183 invasive poeciliids (158 Gambusia affinis and 25 Poecilia reticulata) were collected for gut content analyses from four anchialine sites during wet and dry seasons on the islands of Hawai‘i and Maui. Predation on shrimp was not detected in habitats where they retreat exclusively into the underlying aquifer diurnally and only emerge nocturnally. However, low levels of predation were detected (7/65 fishes, only by Gambusia affinis) at Waianapanapa Cave, Maui, where shrimp retreat into both the aquifer and a cave during the day. Thus, adopted behavioral responses to invasive fishes generally, though not universally, prevent predation on endemic Hawaiian anchialine shrimps. However, non-consumptive effects resulting from behavioral modification of shrimps may have appreciable impacts on the Hawaiian anchialine ecosystem and warrant further study.     

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

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Stretched to the limit; can a short pelagic larval duration connect adult populations of an Indo‐Pacific diadromous fish (Kuhlia rupestris)?

Freshwater species on tropical islands face localized extinction and the loss of genetic diversity. Their habitats can be ephemeral due to variability in freshwater run‐off and erosion. Even worse, anthropogenic effects on these ecosystems are intense. Most of these species are amphidromous or catadromous (i.e. their life cycle includes a marine larval phase), which buffers them against many of these effects. A long pelagic larval duration (PLD) was thought to be critical to ensure the colonization and persistence in tropical islands, but recent findings indicated that several species with short PLDs are successful in those ecosystems. To test the potential of a short PLD in maintaining genetic connectivity and forestalling extirpation, we studied Kuhlia rupestris, a catadromous fish species with an extensive distribution in the western Pacific and Indian Oceans. Using a combination of molecular genetic markers (13 microsatellite loci and two gene regions from mtDNA) and modelling of larval dispersal, we show that a short PLD constrains genetic connectivity over a wide geographical range. Molecular markers showed that the short PLD did not prevent genetic divergence through evolutionary time and speciation has occurred or is occurring. Modelling of larvae dispersal suggested limited recent connectivity between genetically homogeneous populations across the Coral Sea. However, a short PLD can maintain connectivity on a subocean basin scale. Conservation and management of tropical diadromous species needs to take into account that population connectivity may be more limited than previously suspected in those species.     

Amphipod crustaceans from anchihaline cave waters of the Galapagos Islands

Three species of blind Amphipoda are recorded from anchihaline cave waters in the Galapagos Islands. One of these, Galapsiellus leleuporum (Monod, 1970) was previously know from the islands; the male is described for the first time. The other two belong to families with predominantly (though not exclusively) deep-sea members: Valettietta cavernicola sp. nov. (family Lysianassidae; and Antronicippe serrata gen. et sp. nov. (family Pardaliscidae). The genus Valettietta was not previously recorded from caves; it contained two abyssal species in the Atlantic, while a third species was recorded from the Pacific in a vertical haul between 0 and 5300 m. Antronicippe is closely related to Spelaeonicippe , a genus with two species known from anchihaline caves in the Canary Islands and the West Indies.     

Patterns of genetic connectivity among anchialine habitats: a case study of the endemic Hawaiian shrimp Halocaridina rubra on the island of Hawaii

Anchialine habitats, landlocked bodies of mixohaline water that fluctuate with the tides but have no surface connection to the sea, are known from around the world. Many anchialine organisms have widespread distributions and it has been hypothesized that high levels of gene flow and low levels of genetic differentiation are characteristic of populations from these habitats. However, the generality of this hypothesis requires further assessment, particularly in light of the significant negative impact these habitats and their biota have experienced from anthropogenic causes. This study investigated the population structure and demography of an endemic Hawaiian anchialine species, the atyid shrimp Halocaridina rubra, using mitochondrial cytochrome c oxidase subunit I (COI) gene sequences. A survey of 305 individuals from 16 populations collected on the island of Hawaii revealed 135 haplotypes. These haplotypes belonged to one of two divergent (2.7-4.9%) lineages; notably, no haplotypes were shared between the two coasts of the island. Along each coast, strong subdivision and little to no gene flow occurs between populations separated by > 30 km. The population structure and demography of H. rubra on Hawaii are influenced by regional hydrology, geology, volcanism and two distinct colonization events of the island. Thus, H. rubra on Hawaii demonstrates that populations of endemic anchialine organisms may exhibit significant levels of genetic structure and restricted levels of gene flow over limited geographic scales. This report brings novel insight into the biology of anchialine organisms and has important implications for the future management of these habitats and their biota.     

Fish Assemblages of Mediterranean Marine Caves

Fish assemblages associated with 14 marine caves and adjacent external rocky reefs were investigated at four Marine Protected Areas (MPAs) along the coasts of Italy. Within the caves sampling was carried out in different sub-habitats: walls, ceilings, bottoms and ends of caves. On the whole, 38 species were recorded inside the 14 caves investigated. Eighteen species were exclusively found inside the caves: they were mainly represented by speleophilic (i.e. species preferentially or exclusively inhabiting caves) gobids (e.g. Didogobius splechtnai) and nocturnal species (e.g. Conger conger). Forty-one species were censused outside, 20 of which were shared with cave habitats. Apogon imberbis was the most common fish found in all 14 caves investigated, followed by Thorogobius ephippiatus (recorded in 13 caves), and Diplodus vulgaris and Scorpaena notata (both censused in 12 caves). Distinct fish assemblages were found between external rocky reefs and the different cave sub-habitats. New data on the distribution of some speleophilic gobids were collected, showing the existence of a pool of species shared by marine caves on a large scale (i.e. hundreds of km). Considering the uniqueness of cave fishes (18 exclusive species and different assemblage structures), the inclusion of marine caves among the habitats routinely investigated for fish biodiversity monitoring could facilitate the achievement of more comprehensive inventories. Due to their contribution to local species diversity and the shelter they provide to species valuable for conservation, marine caves should be prioritized for their inclusion not only within future MPAs through the Mediterranean Sea, but also into larger management spatial planning.     

Evolution of cave Axiokebuita and Speleobregma (Scalibregmatidae,Annelida)

The evolutionary history of Axiokebuita and Speleobregma, two poorly known lineages of annelids exclusive from deep‐sea or marine caves but always from crevicular habitats, is explored here. Speleobregma lanzaroteum Bertelsen, 1986, and Axiokebuita cavernicola sp. n. are described from anchialine and marine caves of the Canary Islands using light and electron microscopy. Speleobregma lanzaroteum is previously known only from a single specimen from the water column of an anchialine cave in Lanzarote. Emended diagnosis, details on the ciliary patterns and behavioural observations are provided based on newly collected material and in situ observations. Axiokebuita cavernicola sp. n. is found in Pleistocene gravel deposits in a shallow water marine cave in Tenerife (Canary Islands). The new species is characterized by the presence of dorsal ciliary bands and short knob‐like neuropodial cirri from segment two. The porosity and permeability of the gravelly environment of A. cavernicola sp. n. are shown to be equivalent to the water column or crevices of Speleobregma and other Axiokebuita spp. Phylogenetic analyses of five gene fragments and 44 terminals using maximum‐likelihood and Bayesian methods support a derived position of A. cavernicola sp. n. within Axiokebuita and confirm a sister‐group relationship of Axiokebuita with Speleobregma with high nodal support. The Axiokebuita–Speleobregma clade is morphologically characterized by a globular pygidium with adhesive glands and ventral ungrooved ciliated palps. Our results support two independent cave colonization events, favoured by the preadaptation of the members of Axiokebuita–Speleobregma lineage to crevicular habitats.     

Finding answers in the dark: caves as models in ecology fifty years after Poulson and White

The use of semi‐isolated habitats such as oceanic islands, lakes and mountain summits as model systems has played a crucial role in the development of evolutionary and ecological theory. Soon after the discovery of life in caves, different pioneering authors similarly recognized the great potential of these peculiar habitats as biological model systems. In their 1969 paper in Science, ‘The cave environment’, Poulson and White discussed how caves can be used as natural laboratories in which to study the underlying principles governing the dynamics of more complex environments. Together with other seminal syntheses published at the time, this work contributed to establishing the conceptual foundation for expanding the scope and relevance of cave‐based studies. Fifty years after, the aim of this review is to show why and how caves and other subterranean habitats can be used as eco‐evolutionary laboratories. Recent advances and directions in different areas are provided, encompassing community ecology, trophic‐webs and ecological networks, conservation biology, macroecology and climate change biology. Special emphasis is given to discuss how caves are only part of the extended network of fissures and cracks that permeate most substrates and, thus, their ecological role as habitat islands is critically discussed. Numerous studies have quantified the relative contribution of abiotic, biotic and historical factors in driving species distributions and community turnovers in space and time, from local to regional scales. Conversely, knowledge of macroecological patterns of subterranean organisms at a global scale remains largely elusive, due to major geographical and taxonomical biases. Also, knowledge regarding subterranean trophic webs and the effect of anthropogenic climate change on deep subterranean ecosystems is still limited. In these research fields, the extensive use of novel molecular and statistical tools may hold promise for quickly producing relevant information not accessible hitherto.     

The European Tertiary Neritiliidae (Mollusca, Gastropoda, Neritopsina): indicators of tropical submarine cave environments and freshwater faunas

The oldest freshwater neritiliid, Neritilia bisinuata , is described from the Middle Eocene of the Loire Basin. Another European species, N. neritinoides , ranging from the Lower Oligocene to Lower Miocene (Upper Burdigalian) is recognized; its habitat appears to have been freshwater, but very close to the sea. Two new marine neritiliid species from the Aquitaine Basin are described: Bourdieria favia sp. nov. from the Upper Oligocene and Pisulinella aucoini  sp. nov. from the Lower Miocene. A third undescribed species from the Lower Miocene is referred to the same family and related to Pisulinella . The Oligocene species has a strong spiral sculpture, a character completely absent in previously known neritiliid species. The genus Agapilia , founded on juvenile N. neritinoides and adult Vitta picta , appears to be a junior synonym of the genus Vitta. The associated occurrence of shells of the families Neritiliidae, Neritopsidae and Pickworthiidae (well-known inhabitants of Indo-West Pacific submarine caves) at Peyrère suggest the first occurrence of a characteristic assemblage of dark submarine caves during the Oligocene. Both factorial analysis and relative abundance show that at Peyrère these families are associated with other cryptic fossils (various gastropods, bivalves, Brachiopoda, corals, Annelida). However, there are indications of other submarine cave assemblages in various Cenozoic deposits from the Palaeocene to the middle Miocene.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 447–467.     

Phytoplankton of cenotes and anchialine caves along a distance gradient from the northeastern coast of Quintana Roo,Yucatan Peninsula

This work details the taxonomic composition of suspended algae (phytoplankton and tycoplankton) communities in five cenotes (sinkholes) and two anchialine caves in northeastern Quintana Roo, Mexico. The sample set of cenotes are Casa, Nohoch Nah Chich, Maya Blue, Cristal, and Carwash, as well as the two associated caves leading from the cenotes of Maya Blue and Cristal. The site distribution represents a distance gradient with respect to the coastline with which we observe the effects of tidal movement and the mixing of waters (e.g. saline water and freshwater) on the composition of the suspended algae communities. Two sample sets were taken, one at the end of the dry season (March–April 1995) and the second at the end of the rainy season (September–October 1995) with the goal of comparing the contrasting climatic conditions of the region. A total of 79 species were identified, of which, diatoms were the most important with respect to species richness with a total of 75% of species. The floristic composition is very similar between the freshwater cenotes. The distance of a cenote site with respect to the coastline was a determining factor in the species composition. Casa Cenote is the most distinct of the sample set for the presence of marine species due to its proximity to the coastline. The tides are a large determining factor of the floristic composition of Casa Cenote with 24% all species identified in this study found exclusively in this system. The anchialine system species are transported from the cenotes and the adjacent cave systems. The largest percentage or species (95%) are freshwater, and only 5% of the total number of identified species are of marine origin. It is recognized that the most distant cenotes from the coast, Carwash and Cristal, as well as Maya Blue and Nohoch Nah Chich, are the most similar, despite being part of different cave systems. In these inland systems the marine species decreased drastically (2.4% in Nohoch Nah Chich and no marine species in the remaining cenotes). Marine species are found at the halocline of the caves.     

Mitochondrial OXPHOS genes provides insights into genetics basis of hypoxia adaptation in anchialine cave shrimps

Cave shrimps from the genera Typhlatya, Stygiocaris and Typhlopatsa (TST complex) comprises twenty cave-adapted taxa, which mainly occur in the anchialine environment. Anchialine habitats may undergo drastic environmental fluctuations, including spatial and temporal changes in salinity, temperature, and dissolved oxygen content. Previous studies of crustaceans from anchialine caves suggest that they have possessed morphological, behavioral, and physiological adaptations to cope with the extreme conditions, similar to other cave-dwelling crustaceans. However, the genetic basis has not been thoroughly explored in crustaceans from anchialine habitats, which can experience hypoxic regimes. To test whether the TST shrimp-complex hypoxia adaptations matched adaptive evolution of mitochondrial OXPHOS genes. The 13 OXPHOS genes from mitochondrial genomes of 98 shrimps and 1 outgroup were examined. For each of these genes was investigated and compared to orthologous sequences using both gene (i.e. branch-site and Datamonkey) and protein (i.e. TreeSAAP) level approaches. Positive selection was detected in 11 of the 13 candidate genes, and the radical amino acid changes sites scattered throughout the entire TST complex phylogeny. Additionally, a series of parallel/convergent amino acid substitutions were identified in mitochondrial OXPHOS genes of TST complex shrimps, which reflect functional convergence or similar genetic mechanisms of cave adaptation. The extensive occurrence of positive selection is suggestive of their essential role in adaptation to hypoxic anchialine environment, and further implying that TST complex shrimps might have acquired a finely capacity for energy metabolism. These results provided some new insights into the genetic basis of anchialine hypoxia adaptation.     

Distributional dynamics of a specialized subterranean community oppose the classical understanding of the preferred subterranean habitats

Troglobionts are organisms that are specialized for living in a subterranean environment. These organisms reside prevalently in the deepest zones of caves and in shallow subterranean habitats, and complete their entire life cycles therein. Because troglobionts in most caves depend on organic matter resources from the surface, we hypothesized that they would also select the sections of caves nearest the surface, as long as environmental conditions were favorable. Over 1 year, we analyzed, in monthly intervals, the annual distributional dynamics of a subterranean community consisting of 17 troglobiont species, in relation to multiple environmental factors. Cumulative standardized annual species richness and diversity clearly indicated the existence of two ecotones within the cave: between soil and shallow subterranean habitats, inhabited by soil and shallow troglobionts; and between the transition and inner cave zones, where the spatial niches of shallow and deep troglobionts overlap. The mean standardized annual species richness and diversity showed inverse relationships, but both contributed to a better insight into the dynamics of subterranean fauna. Regression analyses revealed that temperatures in the range 7–10°C, high moisture content of substrate, large cross section of the cave, and high pH of substrate were the most important ecological drivers governing the spatiotemporal dynamics of troglobionts. Overall, this study shows general trends in the annual distributional dynamics of troglobionts in shallow caves and reveals that the distribution patterns of troglobionts within subterranean habitats may be more complex than commonly assumed.     

Milyeringa Veritas (Eleotridae), a remarkably versatile Cave Fish from the Arid Tropics of Northwestern Australia

The blind cave gudgeon Milyeringa veritas is restricted to groundwaters of Cape Range and Barrow Island, northwestern Australia. It occurs in freshwater caves and in seawater in anchialine systems. It is associated with the only other stygobitic cave vertebrate in Australia, the blind cave eel, Ophisternon candidum, the world's longest cave fish, and a diverse stygofauna comprising lineages with tethyan tracks and widely disjunct distributions, often from North Atlantic caves. The cave gudgeon inhabits a karst wetland developed in Miocene limestones in an arid area. There is an almost complete lack of information on the basic biology of this cave fish, despite it being listed as threatened under the Western Australian Wildlife Conservation Act. Allozyme frequencies and distributions indicate significant population sub-structuring on the Cape Range peninsula such that the populations are essentially isolated genetically suggesting that more than one biological species is present. Further, they suggest that the vicariant events may have been associated with a series of eustatic low sealevels. Analysis of intestinal contents indicates that they are opportunistic feeders, preying on stygofauna and accidentals trapped in the water, at least at the sites sampled which were open to the surface, a conclusion supported by the results of stable isotope ratio analysis. The gudgeons are found in freshwater caves and throughout deep anchialine systems in which they occur in vertically stratified water columns in which there is a polymodal distribution of water chemistries (temperature, pH, salinity, dissolved oxygen, redox, dissolved inorganic nitrogen series, hydrogen sulphide).     

Genetic analysis of threatened Australian grayling Prototroctes maraena suggests recruitment to coastal rivers from an unstructured marine larval source population

Population genetic variation of Australian grayling Prototroctes maraena was examined to determine whether the dispersal strategy of this amphidromous species favours retention of larvae and juveniles in close proximity to their natal river, or mixing of populations via marine dispersal. Variation in microsatellite and mitochondrial DNA markers was unstructured and differentiation was indistinguishable from zero across four coastal rivers spanning approximately one-quarter of the continental range of the species. This result indicates that the marine larval and juvenile phase probably facilitates extensive gene flow among coastal rivers and agrees with a previous analysis of otolith chemistry that suggested larvae probably move into the sea rather than remain in estuaries. It appears likely that the dispersal strategy of P. maraena would enable recolonization of rivers that experience localized extinction provided that connectivity between freshwater habitats and the sea is sufficient to permit migration and that enough source populations remain intact to support viability of the wider population.