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.     

Contribution of water chemistry and fish condition to otolith chemistry: comparisons across salinity environments

This study quantified the per cent contribution of water chemistry to otolith chemistry using enriched stable isotopes of strontium (⁸⁶Sr) and barium (¹³⁷Ba). Euryhaline barramundi Lates calcarifer, were reared in marine (salinity 40), estuarine (salinity 20) and freshwater (salinity 0) under different temperature treatments. To calculate the contribution of water to Sr and Ba in otoliths, enriched isotopes in the tank water and otoliths were quantified and fitted to isotope mixing models. Fulton's K and RNA:DNA were also measured to explore the influence of fish condition on sources of element uptake. Water was the predominant source of otolith Sr (between 65 and 99%) and Ba (between 64 and 89%) in all treatments, but contributions varied with temperature (for Ba), or interactively with temperature and salinity (for Sr). Fish condition indices were affected independently by the experimental rearing conditions, as RNA:DNA differed significantly among salinity treatments and Fulton's K was significantly different between temperature treatments. Regression analyses did not detect relations between fish condition and per cent contribution values. General linear models indicated that contributions from water chemistry to otolith chemistry were primarily influenced by temperature and secondly by fish condition, with a relatively minor influence of salinity. These results further the understanding of factors that affect otolith element uptake, highlighting the necessity to consider the influence of environment and fish condition when interpreting otolith element data to reconstruct the environmental histories of fish.     

Density-dependent growth as a key mechanism in the regulation of fish populations: evidence from among-population comparisons.

It is generally assumed that fish populations are regulated primarily in the juvenile (pre-recruit) phase of the life cycle, although density dependence in growth and reproductive parameters within the recruited phase has been widely reported. Here we present evidence to suggest that density-dependent growth in the recruited phase is a key process in the regulation of many fish populations. We analyse 16 fish populations with long-term records of size-at-age and biomass data, and detect significant density-dependent growth in nine. Among-population comparisons show a close, inverse relationship between the estimated decline in asymptotic length per unit biomass density, and the long-term average biomass density of populations. A simple population model demonstrates that regulation by density-dependent growth alone is sufficient to generate the observed relationship. Density-dependent growth should be accounted for in fisheries' assessments, and the empirical relationship established here can provide indicative estimates of the density-dependent growth parameter where population-specific data are lacking.     

Competition in Umbra-Perca fish assemblages: experimental and field evidence

Summary One potentially important effect of interspecific competition in freshwater fish communities is to increase predation intensity from gape-limited piscivores by lowering growth rates of prey species. We investigated the operation and consequences of competition between central mudminnows (Umbra limi) and yearling yellow perch (Perca flavescens) in a system where size-limited predation on mudminnows by larger perch is a principle structuring mechanism. During laboratory experiments in which mudminnows foraged for patchily-presented food in the presence and absence of yearling perch, the food intake of mudminnows decreased at both the population and individual-fish levels when perch were present. Mudminnows were neither less active nor did they occupy lower-quality food patches in the presence of perch; exploitation competition, rather than interference or predator avoidance, appeared responsible for reduced feeding success. To assess effects of competition in the field, we examined size distributions and condition factors of yearling mudminnows and perch in small Wisconsin lakes having mudminnow-only and mudminnow-perch assemblages. Yearling mudminnows were smaller and in poorer condition in an assemblage composed (by mass) of 45% yearling perch than in two mudminnow-only assemblages. Conversely, yearling perch were larger and in better condition when the mudminnow-perch assemblage contained only 45% perch then when it contained 92% perch. Our experimental and field results indicated that an asymmetrical competitive relationship could contribute directly to the interspecific population dominance of perch over mudminnows in Umbra-Perca assemblages through reduced food intake, growth, and condition of mudminnows, and indirectly through increased vulnerability of mudminnows to size-limited predation.     

Naïveté in novel ecological interactions: lessons from theory and experimental evidence

The invasion of alien species into areas beyond their native ranges is having profound effects on ecosystems around the world. In particular, novel alien predators are causing rapid extinctions or declines in many native prey species, and these impacts are generally attributed to ecological naïveté or the failure to recognise a novel enemy and respond appropriately due to a lack of experience. Despite a large body of research concerning the recognition of alien predation risk by native prey, the literature lacks an extensive review of naïveté theory that specifically asks how naïveté between novel pairings of alien predators and native prey disrupts our classical understanding of predator–prey ecological theory. Here we critically review both classic and current theory relating to predator–prey interactions between both predators and prey with shared evolutionary histories, and those that are ecologically ‘mismatched’ through the outcomes of biological invasions. The review is structured around the multiple levels of naïveté framework of Banks & Dickman (2007), and concepts and examples are discussed as they relate to each stage in the process from failure to recognise a novel predator (Level 1 naïveté), through to appropriate (Level 2) and effective (Level 3) antipredator responses. We discuss the relative contributions of recognition, cue types and the implied risk of cues used by novel alien and familiar native predators, to the probability that prey will recognise a novel predator. We then cover the antipredator response types available to prey and the factors that predict whether these responses will be appropriate or effective against novel alien and familiar native predators. In general, the level of naïveté of native prey can be predicted by the degree of novelty (in terms of appearance, behaviour or habitat use) of the alien predator compared to native predators with which prey are experienced. Appearance in this sense includes cue types, spatial distribution and implied risk of cues, whilst behaviour and habitat use include hunting modes and the habitat domain of the predator. Finally, we discuss whether the antipredator response can occur without recognition per se, for example in the case of morphological defences, and then consider a potential extension of the multiple levels of naïveté framework. The review concludes with recommendations for the design and execution of naïveté experiments incorporating the key concepts and issues covered here. This review aims to critique and combine classic ideas about predator–prey interactions with current naïveté theory, to further develop the multiple levels of naïveté framework, and to suggest the most fruitful avenues for future research.     

Structuring of the cyanobacterial community by pelagic fish in subtropical reservoirs: experimental evidence from Australia

1. Subtropical reservoirs of Australia are commonly subject to summer blooms of cyanobacteria. The potential for food web manipulation to control cyanobacterial blooms was investigated in Lake Maroon, south east Queensland using enclosures in which the density of the Australian gudgeon Hypseleotris spp. was manipulated. 2. Zooplankton biomass and community structure were strongly affected by fish density. A size dependent predation effect of Hypseleotris on zooplankton was observed at ambient fish densities, and the community shifted towards a dominance of copepod juveniles and nauplii. Substantial increases in the populations of Ceriodaphnia and calanoid copepods were observed at low fish densities and in the absence of fish. 3. At ambient fish densities total phytoplankton and the proportion of cyanobacteria were maintained at levels similar to those prevailing at day 0. Total phytoplankton and the proportion of cyanobacteria decreased substantially at low fish densities and in the absence of fish. Chlorophytes became dominant in the ‘no fish’ treatment and the grazing‐resistant species Oocystis and Dictyosphaerium were significantly higher than at ambient fish densities. 4. The experiment demonstrated a strong positive relationship between Hypseleotris density and cyanobacteria, and the results suggest that subtropical reservoirs may be suited to food web manipulation as a means of controlling summer cyanobacterial blooms.     

Ontogenic development of three GnRH systems in the brain of a pleuronectiform fish, barfin flounder

A pleuronectiform fish, the barfin flounder Verasper moseri, has three molecular forms of gonadotropin-releasing hormone (GnRH) in the brain, salmon GnRH (sGnRH), chicken GnRH-II (cGnRH-II) and seabream GnRH (sbGnRH). To elucidate the ontogenic origin of the neurons that produce these GnRH molecules, the development of three GnRH systems was examined by in situ hybridization and immunocytochemistry. Neuronal somata that express sGnRH mRNA were detected first in the vicinity of the olfactory epithelium 21 days after hatching (Day 21), and then in the transitional area between the olfactory nerve and olfactory bulb and the terminal nerve ganglion on Day 28. cGnRH-II mRNA-expressing neuronal somata were first identified in the midbrain tegmentum near the ventricle on Day 7. cGnRH-II-immunoreactive (ir) fibers were first found in the brain on Day 7. sbGnRH mRNA-expressing neuronal somata were first detected in the preoptic area on Day 42. sbGnRH-ir fibers were localized in the preoptic area-hypothalamus, and formed a distinctive bundle of axons projecting to the pituitary on Day 70. These results indicate that three forms of GnRH neurons have separate embryonic origins in the barfin flounder as in other perciform fish such as tilapia Oreochromis niloticus and red seabream Pagrus major: sGnRH, cGnRH-II and sbGnRH neurons derive from the olfactory placode, the midbrain tegmentum near the ventricle and the preoptic area, respectively.     

Relationships between fish size and otolith size of four bathydemersal fish species from the south eastern Arabian Sea,India

The objective of this study was to estimate a prey body size from the hard parts (e.g. otoliths) of a fish species frequently found in the guts of predators. Length–weight relationships between otolith size (length, height, weight and aspect ratio) and fish size (total length and weight) were determined for four fish species captured in the Arabian Sea by bottom trawl (2015 survey on‐board FORV Sagar Sampada, 200–300 m depth), off the west coast of India: Psenopsis cyanea, Pterygotrigla hemisticta, Bembrops caudimacula and Hoplostethus rubellopterus. No significant differences were noted between the size of the left and right otoliths (t test) in any of the four species. The length–weight relationship of the otolith in all four species showed a negative allometric growth pattern (t test, p < .05). The data fitted well to the regression model for otolith length (OL), otolith height (OH) and otolith weight (OW) to total length (TL) and total weight (TW). Results showed that these relationships are a helpful tool in predicting fish size from the otoliths and in calculating the biomass of these less‐studied fish species during feeding studies and palaentology.     

Orange roughy otolith growth rates: a direct experimental test of the Romanek–Gauldie otolith growth model

Endolymph chemistry (Ca²⁺, pH, Na⁺) was measured in 17 orange roughy (Hoplostethus atlanticus; Teleostei; Trachichthyidae). Theoretical daily microincrement widths based on the Romanek and Gauldie model of otolith growth were calculated from the measure of otolith chemistry. Theoretical daily microincrement widths were well-correlated (r²=59%) with the width of the last daily microincrement growing at the posterior edge of otoliths taken from the sampled fish.     

Ultrastructure of lamellae, mineral and matrix components of fish otolith twinned aragonite crystals: implications for estimating age in fish

Atomic force microscopy (AFM) of the crystalline ultrastructure of otoliths fromPagrus major(Sparidae),Macruronus novaezelandiae(Merlucciidae),Oncorhynchus tshawytscha(Salmonidae),Sebastes alutus(Scorpaenidae), andHoplostethus atlanticus(Trachichthyidae) showed regular deposition of lamellae in the size range 13-490 nm. The orientation of lamellae in the {010} plane was the same as lamellae in crystals of mineral aragonite. Lamellae in mineral aragonite were in the size range 15-45 nm. Lamellae observed in the otolith ofM. novaezelandiaeby transmission electron microscopy showed a range of widths (25-225 nm) similar to lamellae observed by AFM. The observed lamella widths were in the size range that has been described for sub-daily and daily microincrements in otoliths. Observed lamellae widths were also in the size range of alpha-recoil trajectories of(222)Rn and provide a potential diffusion sink correction for the(222)Rn losses in radionuclide method of ageing otoliths. Comparison of the orientations of lamellae to templates based on the Bragg unit cell structure of twinned aragonite indicated that the lamellae resulted from polysynthetic twinning on the {010} aragonite crystal face. Additional cyclic twinning occurred on the {110} face of the aragonite crystal and sometimes led to pseudohexagonal crystals, whose sizes were orders of magnitude larger than lamellae. The organic matrix of the otolith was visible by atomic force and transmission electron microscopy at the nanometer level of resolution, but the organic matrix was confined to the {110} twinning plane of symmetry of the otolith crystal.     

Molecular,cellular and histological changes in skin from a larval to an adult phenotype during bony fish metamorphosis

Developmental models for skin exist in terrestrial and amphibious vertebrates but there is a lack of information in aquatic vertebrates. We have analysed skin epidermal development of a bony fish (teleost), the most successful group of extant vertebrates. A specific epidermal type I keratin cDNA (hhKer1), which may be a bony-fish-specific adaptation associated with the divergence of skin development (scale formation) compared with other vertebrates, has been cloned and characterized. The expression of hhKer1 and collagen 1α1 in skin taken together with the presence or absence of keratin bundle-like structures have made it possible to distinguish between larval and adult epidermal cells during skin development. The use of a flatfish with a well-defined larval to juvenile transition as a model of skin development has revealed that epidermal larval basal cells differentiate directly to epidermal adult basal cells at the climax of metamorphosis. Moreover, hhKer1 expression is downregulated at the climax of metamorphosis and is inversely correlated with increasing thyroxin levels. We suggest that, whereas early mechanisms of skin development between aquatic and terrestrial vertebrates are conserved, later mechanisms diverge. This work was carried out within the project “Arrested development: The Molecular and Endocrine Basis of Flatfish Metamorphosis” (Q5RS-2002-01192) with financial support from the Commission of the European Communities. It does not necessarily reflect the Commission’s views and in no way anticipates its future policy in this area. This project was further supported by Pluriannual funding to CCMAR by the Portuguese Science and Technology Council. M.A. Campinho was sponsored by the Portuguese Ministry of Science (grant no. SFRH/BD/6133/2001).     

Ungulates can control tree invasions: experimental evidence from nonnative conifers and sheep herbivory

Invasive conifer species are increasingly recognized as a serious problem in many parts of the world, where they are having large ecological and economic impacts. Understanding factors that trigger and can control invasion is key to management. Grazing and browsing by large herbivores have been suggested as a mechanism that may halt conifer invasions, although conflicting results have been reported (i.e. positive, negative or no effect of grazing on invasion). We believe that some of these opposing responses arise due to the absence of well-planned and replicated experiments, since current evidence is mostly observational, and for example, differences in animal densities can produce different results. Thus, in this study, we tested whether large herbivores can control invasion by nonnative conifers and whether the severity of the invasion process would be lessened by increased herbivory intensity. We evaluate experimentally herbivore damage on Pinus contorta, a highly invasive species in many countries of the Southern Hemisphere, under different sheep stocking rates in Patagonia, Argentina. We used four stocking rates, corresponding to 1, 2, 4 and 8 times the recommended sheep herbivory intensity for the study area. The response was not linear but rather presented a threshold. The greater stocking rate, the greater the browsing, higher reduction in seedling height, and decrease of survival of P. contorta. Also, the highest stocking rate damaged and killed 99% of them. This study provides evidence that large domestic herbivores can play a key role in the invasion process and, if managed properly, would provide a tool to help limit conifer invasion.     

Inter‐individual variation promotes ecological success of populations and species: evidence from experimental and comparative studies

Biological diversity is threatened by exploitation, fragmentation of natural habitats, pollution, climate change, and anthropogenic spread of species. The question of how among‐individual variation influences the performance of populations and species is a poorly explored but currently growing field of research. Here, we review 31 experimental and 14 comparative studies and first investigate whether there is empirical support for the propositions that higher levels of among‐individual phenotypic and genetic variation promote the ecological and evolutionary success of populations and species in the face of environmental change. Next, we examine whether and how the effect of diversity depends on environmental conditions. Finally, we explore whether the relationship linking population fitness to diversity is typically linear, asymptotic, or whether the benefits peak at intermediate diversity. The reviewed studies provide strong, almost invariable, evidence that more variable populations are less vulnerable to environmental changes, show decreased fluctuations in population size, have superior establishment success, larger distribution ranges, and are less extinction prone, compared with less variable populations or species. Given the overwhelming evidence that variation promotes population performance, it is important to identify conditions when increased variation does not have the theoretically expected effect, a question of considerable importance in biodiversity management, where there are many other practical constraints. We find that experimental outcomes generally support the notion that genetic and phenotypic variation is of greater importance under more stressful than under benign conditions. Finally, population performance increased linearly with increasing diversity in the majority (10 of 12) of manipulation studies that included four or more diversity levels; only two experiments detected curvilinear relationships.     

Sodium ion cycle in bacterial pathogens: evidence from cross-genome comparisons.

Analysis of the bacterial genome sequences shows that many human and animal pathogens encode primary membrane Na+ pumps, Na+-transporting dicarboxylate decarboxylases or Na+ translocating NADH:ubiquinone oxidoreductase, and a number of Na+ -dependent permeases. This indicates that these bacteria can utilize Na+ as a coupling ion instead of or in addition to the H+ cycle. This capability to use a Na+ cycle might be an important virulence factor for such pathogens as Vibrio cholerae, Neisseria meningitidis, Salmonella enterica serovar Typhi, and Yersinia pestis. In Treponema pallidum, Chlamydia trachomatis, and Chlamydia pneumoniae, the Na+ gradient may well be the only energy source for secondary transport. A survey of preliminary genome sequences of Porphyromonas gingivalis, Actinobacillus actinomycetemcomitans, and Treponema denticola indicates that these oral pathogens also rely on the Na+ cycle for at least part of their energy metabolism. The possible roles of the Na+ cycling in the energy metabolism and pathogenicity of these organisms are reviewed. The recent discovery of an effective natural antibiotic, korormicin, targeted against the Na+ -translocating NADH:ubiquinone oxidoreductase, suggests a potential use of Na+ pumps as drug targets and/or vaccine candidates. The antimicrobial potential of other inhibitors of the Na+ cycle, such as monensin, Li+ and Ag+ ions, and amiloride derivatives, is discussed.     

Expression and hormonal control of a new larval cuticular multigene family at the onset of metamorphosis of the tobacco hornworm

The pattern of cuticular protein synthesis by the epidermis of the tobacco hornworm larva changes during the final day of feeding, leading to an alteration in cuticular structure and a stiffening of the cuticle. We have isolated a small multigene family which codes for at least three of the new cuticular proteins made at this time. The five genes which were isolated from this family map to two different genomic regions. Sequencing shows that one of the genes is 1.9 kb and consists of three exons coding for a 12.2-kDa acidic (pI = 5.26) protein that is predominantly hydrophilic. The deduced amino acid sequence shows regions of similarity to proteins from flexible lepidopteran cuticles and from Drosophila larval and pupal cuticles, but not to proteins found in highly sclerotized cuticles. This gene family is first expressed late on the penultimate day (Day 2) of feeding in the final larval instar and ceases expression 2 days later when metamorphosis begins. In situ hybridization shows that this gene family is expressed in all the epidermal cells of Day 3 larvae except the bristle cells and those at the muscle attachment site. Expression can be induced in Day 1 epidermis by exposure to 50 ng/ml 20-hydroxyecdysone in vitro, but only if juvenile hormone is absent. Its developmental expression, tissue specificity, and hormonal regulation strongly suggest that this multigene family is involved in the structural changes that occur in the larval cuticle just prior to the onset of metamorphosis.     

Comparison of spatial variation in otolith chemistry of two fish species and relationships with water chemistry and otolith growth

Spatial variation in the chemistry (Mg, Mn, Sr and Ba) of recently deposited otolith material (last 20–30 days of life) was compared between two demersal fish species; snapper Pagrus auratus (Sparidae) and sand flathead Platycephalus bassensis (Platycephalidae), that were collected simultaneously at 12 sites across three bays in Victoria, south-eastern Australia. Otolith chemistry was also compared with ambient water chemistry and among three sampling positions adjacent to the proximal otolith margin. For both species, variation in otolith chemistry among bays was significant for Ba, Mn and Sr; however, differences among bays were only similar between species for Ba and Mn. Only Ba showed significant variation at the site level. Across the 12 sites, mean otolith Ba levels were significantly positively correlated between species. Further, although incorporation rates differed, mean ambient Ba levels for both species were positively correlated with ambient Ba levels. Spatial variation in multi-element otolith chemistry was also broadly similar between species and with multi-element water chemistry. Partition coefficients clearly indicated species-specific incorporation of elements into otoliths. Mg and Mn were consistently higher in snapper than sand flathead otoliths (mean ±s .d ., Mg snapper 22·1 ± 3·8 and sand flathead 9·9 ± 1·5 μg g⁻¹, Mn snapper 4·4 ± 2·6 and sand flathead 0·5 ± 0·3 μg g⁻¹), Sr was generally higher in sand flathead otoliths (sand flathead 1570 ± 235 and snapper 1346 ± 104 μg g⁻¹) and Ba was generally higher in snapper otoliths (snapper 12·1 ± 12·8 and sand flathead 1·8 ± 1·4 μg g⁻¹). For both species, Mg and Mn were higher in the faster accreting regions of the otolith margin, Sr was lower in the slower accreting region and Ba showed negligible variation among the three sampling regions. This pattern was consistent with the higher Mg and Mn, and generally lower Sr observed in the faster accreting snapper otoliths. It is hypothesized that the differences between species in the incorporation of these elements may be at least partly related to differences in metabolic and otolith accretion rate. Although rates of elemental incorporation into otoliths appear species specific, for elements such as Ba where incorporation appears consistently related to ambient concentrations, spatial variation in otolith chemistry should show similarity among co-occurring species.