Have fishes had their chips? The dilemma of threatened fishes

Synopsis The conservation status and factors threatening fishes worldwide are reviewed in order to introduce a series of one-page articles on Threatened fishes of the world, and to encourage the incorporation of information on threatened fishes into international conservation programmes. Information on fish extinction and threat rates are compared with those of other animal groups, and the unique characteristics of fish conservation problems are highlighted. At present 979 species of fishes are listed as threatened in the IUCN Red List and at least 36 species and three subspecies are listed as recently extinct. It is argued that these figures are probably gross underestimates and that they may mislead conservation authorities and resource users about the seriousness of the situation. Freshwater fishes may be the most threatened group of vertebrates after the Amphibia. Urgent action is required to save many narrowly endemic, stenotopic species from extinction, especially in Africa, Asia and South America. The conservation of common species that drive essential ecological processes is also important. Anthropogenic pressures, especially habitat degradation, the introduction of invasive species and pollution, on inland and coastal waters are particularly severe and many major fish communities are threatened with elimination throughout the world. The conservation of marine fishes is complicated by the fact that it is difficult to ascertain their rarity. The importance of the retention of genetic variation is highlighted, and both orthodox and innovative conservation measures are encouraged. Further research on minimum viable populations, genetics, and the factors that cause fishes to become vulnerable to extinction, is urgently required.Invited editorial     

Are all fishes ancient polyploids?

Euteleost fishes seem to have more copies of many genes than their tetrapod relatives. Three different mechanisms could explain the origin of these 'extra' fish genes. The duplicates may have been produced during a fish-specific genome duplication event. A second explanation is an increased rate of independent gene duplications in fish. A third possibility is that after gene or genome duplication events in the common ancestor of fish and tetrapods, the latter lost more genes. These three hypotheses have been tested by phylogenetic tree reconstruction. Phylogenetic analyses of sequences from human, mouse, chicken, frog (Xenopus laevis), zebrafish (Danio rerio) and pufferfish (Takifugu rubripes) suggest that ray-finned fishes are likely to have undergone a whole genome duplication event between 200 and 450 million years ago. We also comment here on the evolutionary consequences of this ancient genome duplication.     

Non-laminar cerebral cortex in teleost fishes?

A large skull is disadvantageous to animals that move quickly in three-dimensional space, such as fishes and birds in water or air. A cerebral neocortex with a six-layered sheet has not evolved, most likely due to the limited cranial space. Instead of the laminar cortex, telencephalic nuclear masses seem to have evolved as the pallium in teleost fishes. We consider that the nuclear masses contain rather simple neural circuits sharing a skeleton of simple circuits in the mammalian cortex, which have been elaborated by additional circuits in mammals. Such basic similarities at the connectional level shared by nuclear and cortical pallium might underlie similar or equivalent functions.     

Does respiratory sinus arrhythmia occur in fishes?

The hypothesis that respiratory modulation of heart rate variability (HRV) or respiratory sinus arrhythmia (RSA) is restricted to mammals was tested on four Antarctic and four sub-Antarctic species of fish, that shared close genotypic or ecotypic similarities but, due to their different environmental temperatures, faced vastly different selection pressures related to oxygen supply. The intrinsic heart rate (fH) for all the fish species studied was approximately 25% greater than respiration rate (fV), but vagal activity successively delayed heart beats, producing a resting fH that was synchronized with fV in a progressive manner. Power spectral statistics showed that these episodes of relative bradycardia occurred in a cyclical manner every 2-4 heart beats in temperate species but at >4 heart beats in Antarctic species, indicating a more relaxed selection pressure for cardio-respiratory coupling. This evidence that vagally mediated control of fH operates around the ventilatory cycle in fish demonstrates that influences similar to those controlling RSA in mammals operate in non-mammalian vertebrates.     

Does diet in Celtic Sea fishes reflect prey availability?

Feeding preferences of Celtic Sea fishes were investigated using a database of stomach content records, collected between 1977 and 1994. The diet of cod Gadus morhua , hake Merluccius merluccius , megrim Lepidorhombus whiffiagonis , whiting Merlangius merlangus and saithe Pollachius virens changed markedly as the animals grew larger, and although large predators generally chose larger bodied prey, the variability of prey sizes consumed also increased. Large predators continued to select small, low value, benthic prey ( e.g . Callionymus spp. and Trisopterus spp.) which were easier to catch, rather than larger, more energy lucrative pelagic prey ( e.g . mackerel Scomber scombrus ), even though these pelagic prey‐fishes were nearly always available and were often very abundant. Stock estimates of the International Council for the Exploration of the Sea and U.K. groundfish survey catches were used as indices of prey abundance. Blue‐whiting Micromesistius poutassou and other small pelagic fishes ( Argentina spp. and clupeoids) were identified as being particularly important, and were consumed by some predators more often than would be expected given the abundance of these prey in the environment. There was no evidence for density‐dependent feeding by predators on mackerel and only hake exhibited density‐dependent feeding on horse‐mackerel. Hake, cod and megrim consumed more blue‐whiting when this prey was at higher abundance in the environment. In choosing what prey to consume, predators must balance costs and benefits, considering the quality of prey and the energy expended during search, capture and handling.     

Mhc class II B gene evolution in East African cichlid fishes

 A distinctive feature of essential major histocompatibility complex (Mhc) loci is their polymorphism characterized by large genetic distances between alleles and long persistence times of allelic lineages. Since the lineages often span several successive speciations, we investigated the behavior of the Mhc alleles during or close to the speciation phase. We sequenced exon 2 of the class II B locus 4 from 232 East African cichlid fishes representing 32 related species. The divergence times of the (sub)species ranged from 6000 to 8.4 million years. Two types of evolutionary analysis were used to elucidate the pattern of exon 2 sequence divergence. First, phylogenetic methods were applied to reconstruct the most likely evolutionary pathways leading from the last common ancestor of the set to the extant sequences, and to assess the probable mechanisms involved in allelic diversification. Second, pairwise comparisons of sequences were carried out to detect differences seemingly incompatible with origin by nonparallel point mutations. The analysis revealed point mutations to be the most important mechanism behind allelic divergences, with recombination playing only an auxiliary part. Comparison of sequences from related species revealed evidence of random allelic (lineage) losses apparently associated with speciation. Sharing of identical alleles could be demonstrated between species that diverged 2 million years ago. The phylogeny of the exon was incongruent with that of the flanking introns, indicating either a high degree of convergent evolution at the peptide-binding region-encoding sites, or intron homogenization. Received: 6 December 1999 / Revised: 15 February 2000     

Coexistence of coral reef fishes — a lottery for living space

Synopsis Data are summarised from studies of two reef fish communities — pomacentrids territorial on rubble patches, and fishes resident in small isolated colonies of coral. In each case there is evidence that availability of living sites limits numbers of fishes, and that similar species of fish use the same kinds of spaces. Priority of arrival as recruits, rather than subtle differences in requirements or competitive abilities of adults, appears to determine which species holds each site. Faced with a limited and patchy supply of living space, most reef fishes are sedentary as adults, and produce frequent clutches of pelagic larvae over extended breeding seasons In this way they maximise their chances of getting offspring into suitable living sites as such sites appear. It is argued that by adopting this strategy, reef fishes are preadapted for forming inter-specific lotteries for living space if several species with similar requirements occur together. Such lotteries among similar species may be a feature common to many reef fish communities, and may explain the typically high within-site diversity found in them.This paper forms part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Helfman, ed.).     

Incipient speciation driven by hypertrophied lips in Midas cichlid fishes?

Sympatric speciation has been debated in evolutionary biology for decades. Although it has gained in acceptance recently, still only a handful of empirical examples are seen as valid (e.g. crater lake cichlids). In this study, we disentangle the role of hypertrophied lips in the repeated adaptive radiations of Nicaraguan crater lake cichlid fish. We assessed the role of disruptive selection and assortative mating during the early stages of divergence and found a functional trade‐off in feeding behaviour between thick‐ and thin‐lipped ecotypes, suggesting that this trait is a target of disruptive selection. Thick‐lipped fish perform better on nonevasive prey at the cost of a poorer performance on evasive prey. Using enclosures in the wild, we found that thick‐lipped fish perform significantly better in rocky than in sandy habitats. We found almost no mixed pairs during two breeding seasons and hence significant assortative mating. Genetic differentiation between ecotypes seems to be related to the time since colonization, being subtle in L. Masaya (1600 generations ago) and absent in the younger L. Apoyeque (<600 generations ago). Genome‐wide differentiation between ecotypes was higher in the old source lakes than in the young crater lakes. Our results suggest that hypertrophied lips might be promoting incipient sympatric speciation through divergent selection (ecological divergence in feeding performance) and nonrandom mating (assortative mating) in the young Nicaraguan crater lakes. Nonetheless, further manipulative experiments are needed in order to confirm the role of hypertrophied lips as the main cue for assortative mating.     

Is post‐feeding thermotaxis advantageous in elasmobranch fishes?

The effects of post‐feeding thermotaxis on ileum evacuation and absorption rates were examined in the laboratory using two elasmobranch species, the Atlantic stingray Dasyatis sabina, which inhabits thermally variable environments, and the whitespotted bamboo shark Chiloscyllium plagiosum, a stenothermic fish living on Indo‐Pacific reefs. Experiments at temperatures similar to those experienced in nature revealed temperature change had no significant effect on C. plagiosum absorption or evacuation rates, suggesting stenothermic sharks cannot exploit temperature differences as a means to improve digestion efficiency. On the other hand, D. sabina showed significantly lower evacuation and absorption rates at lower temperatures. The relative decrease was greater for evacuation (Q₁₀ = 3·08) than absorption rates (Q₁₀ = 2·20), resulting in a significant increase in total absorption, suggesting D. sabina can benefit from using shuttling behaviour to exploit thermal variability in their environment to maximize energetic uptake.     

Foraging habitat determines predator–prey size relationships in marine fishes

Predator–prey size (PPS) relationships are determined by predator behaviour, with the likelihood of prey being eaten dependent on their size relative to that of the consumer. Published PPS relationships for 30 pelagic or benthic marine fish species were analysed using quantile regression to determine how median, lower and upper prey sizes varied with predator size and habitat. Habitat effects on predator foraging activity/mode, morphology, growth and natural mortality are quantified and the effects on PPS relationships explored. Pelagic species are more active, more likely to move by caudal fin propulsion and grow more rapidly but have higher mortality rates than benthic species, where the need for greater manoeuvrability when foraging in more physically complex habitats favours ambush predators using pectoral fin propulsion. Prey size increased with predator size in most species, but pelagic species ate relatively smaller prey than benthic predators. As pelagic predators grew, lower prey size limits changed little, and prey size range increased but median relative prey size declined, whereas the lower limit increased and median relative prey size was constant or increased in benthic species.     

Species richness of exotic and endangered fishes in Japan’s reservoirs

We analyzed the patterns of total, endangered and exotic fish species richness in 80 reservoirs throughout Japan using Generalized Linear Models (GLMs) with variables of dam characteristics (e.g. reservoir size, isolated period, altitude, DO, pH, rainfall, and air temperature) and watershed characteristics (e.g. watershed area, human population, and land-use patterns). Exotic species richness was positively correlated with total species richness but negatively correlated with endangered species richness. Largemouth bass, rainbow trout, and bluegill were the three most widely distributed species, occupying 47.5, 37.5, and 33.8% of the reservoirs, respectively. The patterns of total and endangered species richness were largely explained by watershed area and annual air temperature. Exotic species richness was determined primarily by the proportion of developed areas within each watershed. Therefore, reservoirs in more developed areas tend to support more exotic fish species that in turn threaten endangered fish species more severely.

How many species of cichlid fishes are there in African lakes?

The endemic cichlid fishes of Lakes Malawi, Tanganyika and Victoria are textbook examples of explosive speciation and adaptive radiation, and their study promises to yield important insights into these processes. Accurate estimates of species richness of lineages in these lakes, and elsewhere, will be a necessary prerequisite for a thorough comparative analysis of the intrinsic and extrinsic factors influencing rates of diversification. This review presents recent findings on the discoveries of new species and species flocks and critically appraises the relevant evidence on species richness from recent studies of polymorphism and assortative mating, generally using behavioural and molecular methods. Within the haplochromines, the most species-rich lineage, there are few reported cases of postzygotic isolation, and these are generally among allopatric taxa that are likely to have diverged a relatively long time in the past. However, many taxa, including many which occur sympatrically and do not interbreed in nature, produce viable, fertile hybrids. Prezygotic barriers are more important, and persist in laboratory conditions in which environmental factors have been controlled, indicating the primary importance of direct mate preferences. Studies to date indicate that estimates of alpha (within-site) diversity appear to be robust. Although within-species colour polymorphisms are common, these have been taken into account in previous estimates of species richness. However, overall estimates of species richness in Lakes Malawi and Victoria are heavily dependent on the assignation of species status to allopatric populations differing in male colour. Appropriate methods for testing the specific status of allopatric cichlid taxa are reviewed and preliminary results presented.     

Filial cannibalism in fishes: Why do parents eat their offspring?

Filial cannibalism (the eating of one's own offspring) occurs in a variety of taxa, but is especially prevalent in fishes with parental care. Recent research supports a central tenet of parental-investment theory; that is, parents consume their offspring when it maximizes their lifetime reproductive success. This review outlines the theoretical framework used to explain the adaptive significance of filial cannibalism, evaluates experimental studies to test some predictions of this theory and discusses how the occurrence of filial cannibalism affects other aspects of a species' reproductive ecology.     

Determinants of the steep species–area relationship of coral reef fishes

The increase in species richness with area is known as the species–area relationship (SPAR). Although several mutually non-exclusive processes may produce the SPAR, the null, often ignored, hypothesis states that a SPAR can be generated by random placement alone. The log–log-transformed SPAR of coral reef fishes on small patch-reefs revealed a steep slope of 0.55. However, this slope was dependent on the cumulative area of the reef examined and was therefore affected by random placement. After statistically removing the contribution of random placement from the SPAR, the slope was estimated to be 0.21. This is consistent with estimates from other, mostly terrestrial, systems. Furthermore, a randomization procedure, where the probability of fishes to reach a patch was proportional to reef area, showed that the field measured SPAR did not differ from random placement. In addition, fish assemblages on species poor reefs did not form subsets of species rich reefs (i.e., no nestedness) beyond that expected from random placement. Steep log–log-transformed SPARs can be formed by random placement alone, indicating that caution should be used when assigning an ecological meaning to SPARs generated from small spatial scales.     

Is density-dependent growth in young-of-the-year fishes a question of critical weight?

We develop a conceptual argument thatdensity-dependent growth via reductions in preyresources are most likely to occur in the late-larvalor juvenile stage in both marine and freshwaterfishes. We use results from a suite ofindividual-based models and literature examples of avariety of marine, estuarine and freshwater species toprovide evidence of the effects of age-0 fish on theirprey. We conclude that larval-stage survival relatedto food-limited growth contributes significantly torecruitment variability. However, density-dependentregulation of cohort biomass via feedbacks derivedfrom reductions in prey resources is most likely tooccur at a ``critical-weight' during the late-larval orjuvenile stage. This occurs when fish densitiesremain relatively high, and population consumption ishighest relative to prey density and replenishmentrate. We compare our critical weight concept toHoude's (1997) critical size hypothesis.     

Can otolith elemental chemistry retrospectively track migrations in fully marine fishes?

Otolith microchemistry can provide valuable information about stock structure and mixing patterns when the magnitude of environmental differences among areas is greater than the cumulative influence of any vital effects. Here, the current understanding of the underlying mechanisms governing element incorporation into the otolith is reviewed. Hard and soft acid and base (HSAB) theory is employed to explore the differences in chemical behaviours, distributions and affinities between elements. Hard acid cations (e.g. Mg(2+) , Li(+) and Ba(2+) ) tend to be less physiologically influenced and accepted more readily into the otolith crystal lattice but are relatively homogeneous in seawater. Soft acid cations (e.g. Zn(2+) and Cu(2+) ) on the other hand, exhibit more varied distributions in seawater, but are more likely to be bound to blood proteins and less available for uptake into the otolith. The factors influencing the geographical distribution of elements in the sea, and their incorporation into the otoliths of marine fishes are reviewed. Particular emphasis is placed on examining physiological processes, including gonad development, on the uptake of elements commonly used in population studies, notably Sr. Finally, case studies are presented that either directly or indirectly compare population structuring or movements inferred by otolith elemental fingerprints with the patterns indicated by additional, alternative proxies. The main obstacle currently limiting the application of otolith elemental microchemistry to infer movements of marine fishes appears to lie in the largely homogeneous distribution of those elements most reliably measured in the otolith. Evolving technologies will improve the discriminatory power of otolith chemistry by allowing measurement of spatially explicit, low level elements; however, for the time being, the combination of otolith minor and trace element fingerprints with alternative proxies and stable isotopic ratios can greatly extend the scope of migration studies. Among the otolith elements that routinely occur above instrument detection limits, Ba, Mn and Li were deemed the most likely to prove reliable geographic markers in marine species.     

Can filter‐feeding fishes improve water quality in lakes?

1.  In this paper we examine the potential of a cichlid fish species ( Sarotherodon galilaeus ) to both maintain positive growth rates through filter-feeding on phytoplankton and improve water quality in Lake Kinneret through suppression of dinoflagellate ( Peridinium gatunense ) blooms.
2.  Seasonal plankton consumption by S. galilaeus from Lake Kinneret was examined experimentally by monitoring changes in plankton assemblages during 24 h in 5-m³ mesocosms containing varying densities of fish. Taxon-specific grazing rates ranged from 0 to 17 mg g_fish day^–1, with mean total consumption of 1.6% fish body weight per day. During the spring bloom of P. gatunense , S. galilaeus consumed mostly (94%) netphytoplankton (≥20 μm). The remaining 6% consisted mostly of nanophytoplankton (<20 μm). During the summer and fall, net- and nanophytoplankton accounted for 54 and 42%, respectively, of the diet of S. galilaeus . Zooplankton and flagellated and ciliated protozoans made up the remaining 4%.
3.  Simulations using a fish bioenergetics model indicated that consumption rates ( C ) were near maximum in spring (90% C _max), while consumption was reduced in summer-fall (59% C _max). Sarotherodon galilaeus obtains sufficient energy through filter-feeding year-round, although most growth (≥60%) occurs during the spring P. gatunense bloom.
4.  Despite efficient feeding on P. gatunense and nanophytoplankton by S. galilaeus , estimates of instantaneous plankton mortality caused by ingestion were two orders of magnitude lower than maximum potential plankton growth rates. Thus the potential for the S. galilaeus population in Lake Kinneret to positively affect water quality through algal suppression is low.     

Is metamorphosis a critical interval in the early life of marine fishes?

Synopsis Small volumes of food in ‘stomach’, large proportions of empty ‘stomachs’ and small maximum sizes of prey during metamorphosis of cod,Gadus morhua, all point to an energy crisis during this developmental interval. The small size of the developing stomach of the 10–14 mm long cod may be partly responsible for the above mentioned difficulties during metamorphosis. A hypothesis based on Hjort's ‘critical stage theory’ is proposed, adding another critical interval at metamorphosis. Based on the presented data on cod and reports on other species in the literature, the new critical interval is discussed for larvae of marine fish species in general.     

Can upwelling signals be detected in intertidal fishes of different trophic levels?

For intertidal fishes belonging to three species, the herbivore Scartichthys viridis (Blenniidae), the omnivore Girella laevifrons (Kyphosidae) and the carnivore Graus nigra (Kyphosidae), mass and body size relationships were higher in individuals from an upwelling zone compared with those from a non‐upwelling zone. RNA:DNA were higher in the herbivores and omnivores from the upwelling zone. Higher biomass and RNA:DNA in the upwelling intertidal fishes may be a consequence of an increased exposure to higher nutrient availability, suggesting that increased physiological conditioning in vertebrates from upwelling areas can be detected and measured using intertidal fishes of different trophic levels.