Population dynamics of Hawaiian seabird colonies vulnerable to sea-level rise

Globally, seabirds are vulnerable to anthropogenic threats both at sea and on land. Seabirds typically nest colonially and show strong fidelity to natal colonies, and such colonies on low-lying islands may be threatened by sea-level rise. We used French Frigate Shoals, the largest atoll in the Hawaiian Archipelago, as a case study to explore the population dynamics of seabird colonies and the potential effects sea-level rise may have on these rookeries. We compiled historic observations, a 30-year time series of seabird population abundance, lidar-derived elevations, and aerial imagery of all the islands of French Frigate Shoals. To estimate the population dynamics of 8 species of breeding seabirds on Tern Island from 1980 to 2009, we used a Gompertz model with a Bayesian approach to infer population growth rates, density dependence, process variation, and observation error. All species increased in abundance, in a pattern that provided evidence of density dependence. Great Frigatebirds (Fregata minor), Masked Boobies (Sula dactylatra), Red-tailed Tropicbirds (Phaethon rubricauda), Spectacled Terns (Onychoprion lunatus), and White Terns (Gygis alba) are likely at carrying capacity. Density dependence may exacerbate the effects of sea-level rise on seabirds because populations near carrying capacity on an island will be more negatively affected than populations with room for growth. We projected 12% of French Frigate Shoals will be inundated if sea level rises 1 m and 28% if sea level rises 2 m. Spectacled Terns and shrub-nesting species are especially vulnerable to sea-level rise, but seawalls and habitat restoration may mitigate the effects of sea-level rise. Losses of seabird nesting habitat may be substantial in the Hawaiian Islands by 2100 if sea levels rise 2 m. Restoration of higher-elevation seabird colonies represent a more enduring conservation solution for Pacific seabirds.     

Ecology and conservation of the rare annual Petrorhagia nanteuilii (Childing Pink) on the vegetated shingle spits of Pagham Harbour, West Sussex

Petrorhagia nanteuilii (Childing Pink), a nationally rare annual, exists at only one location in the UK on the vegetated shingle spits at Pagham Harbour, West Sussex. Primarily found in dry grassland habitats throughout Spain and Portugal, at present its only known threat in the UK is the invasive perennial, Centranthus ruber (Red Valerian); however for conservation of Petrorhagia nanteuilii to be sustainable, its presence at Pagham Harbour must be better understood. The aim of this study was therefore to investigate the local distribution and ecological associations of Petrorhagia nanteuilii in order to understand the relationships between Petrorhagia nanteuilii and the vegetated shingle communities and thereby inform their conservation. Field surveys of the shingle flora were undertaken in 2007 and 2011. Vegetation was sampled and the edaphic and physical environment was also investigated using TWINSPAN, multivariate analyses and GIS. The results of the study show that Petrorhagia nanteuilii has little association with the true vegetated shingle communities on the shingle spits. They also reveal that in recent years the species has spread rapidly across parts of the site due to a number of adaptations aiding its survival and recent shifts in conservation management that have influenced spit morphology. The ecological disconnection that exists between Petrorhagia nanteuilii and the true shingle communities means it is unlikely that management strategies will be conflicting. The study concludes that conservation of both is feasible, however for management to be sustainable, both short-term and long-term strategies must be considered.     

Klimawandel in Küstenzonen

The IPCC-TAR (2001) depicts the consequences of climate change on a global scale with a central focus on vulnerability and adaptation. The objective of this article is the important regional/local scale: First, sea-level rise, storm surges and related issues are perceived here, and second, decisions about precautionary measures, and coastal protection are mostly initialised on this scale. The Integrated Coastal Zone Management (ICZM) is described as an auspicious approach managing conflicts in coastal zones.     

Possible impact of sea-level rise on some habitat types at the Baltic coast of Denmark

According to estimates from the Danish Meteorological Institute global warming until 2080 may cause a relative sea-level rise in Danish waters of 33–46 cm. In the present paper the possible impact of a sea-level rise of this magnitude on coastal habitat types is discussed for three case studies, based on previous investigations of vegetation, topography and soil of localities at the Baltic coast of Denmark. The case studies include the following types of localities and habitats: (1) an off-shore barrier complex: sandy beach, sand dune, geolittoral, brackish, low-tidal meadow, reed bed; (2) a protected bay: geolittoral, brackish meadow, coastal grassland; (3) a dune area: mobile and fixed dune communities, and adjoining sea wall: coastal grassland. In the geolittoral meadow and coastal grassland habitats the sea-level rise is expected to cause a horizontal displacement of vegetation zones and a reduction in area, depending on accretion rate (sedimentation, peat formation), local topography and inland land-use. In the beach and sand dune habitats the sea-level rise is expected to cause a change in groundwater level, influencing slack vegetation, and a change in the erosion/accretion pattern, resulting in landward rebuilding of the mobile dune as well as in a more or less diffuse inland sand drift, causing destabilization of fixed dune vegetation.     

Predicting the Impact of Sea-Level Rise on Caribbean Sea Turtle Nesting Habitat

Abstract:  The projected rise in sea level is likely to increase the vulnerability of coastal zones in the Caribbean, which are already under pressure from a combination of anthropogenic activities and natural processes. One of the major effects will be a loss of beach habitat, which provides nesting sites for endangered sea turtles. To assess the potential impacts of sea-level rise on sea turtle nesting habitat, we used beach profile measurements of turtle nesting beaches on Bonaire, Netherlands Antilles, to develop elevation models of individual beaches in a geographic information system. These models were then used to quantify areas of beach vulnerable to three different scenarios of a rise in sea level. Physical characteristics of the beaches were also recorded and related to beach vulnerability, flooding, and nesting frequency. Beaches varied in physical characteristics and therefore in their vulnerability to flooding. Up to 32% of the total current beach area could be lost with a 0.5-m rise in sea level, with lower, narrower beaches being the most vulnerable. Vulnerability varied with land use adjacent to the beach. These predictions about loss of nesting habitat have important implications for turtle populations in the region.     

Estuarine Vegetated Habitats as Corridors for Predator Movements

Abstract: The spatial proximity of one habitat to another can strongly influence population and community dynamics. We investigated whether the proximity of intertidal oyster reefs to vegetated estuarine habitats, salt marshes, and seagrass beds, affects the abundance and community structure of benthic macroinvertebrates on reefs and predator-prey interactions between mobile predators and bivalves living on reefs. Benthic macroinvertebrate abundance was highest on reefs spatially separated from salt marshes. Macroinvertebrate species richness was highest on reefs separated from both salt marshes and seagrass beds. Comparisons of predation on juvenile bivalves transplanted to reefs for 7–12 days indicated that survivorship of clams was greatest on reefs spatially separated from both salt marshes and seagrass beds, whereas reef proximity to vegetated habitats did not affect the survivorship of oysters. The foraging behavior of blue crabs may explain patterns of macroinvertebrate abundance and clam survivorship among reefs with different proximity to vegetated habitats. In experiments conducted in 30-m² field enclosures, blue crabs had higher predation rates on hard clams transplanted onto artificial reefs adjacent to salt marshes or seagrass beds than on reefs separated from both habitats by unvegetated sand bottom. Thus, vegetated habitats appeared to act as corridors by facilitating the access of blue crabs to oyster reefs and enhancing the intensity of blue crab predation. Such an understanding of the effects of landscape characteristics of estuarine habitats on their value as habitats for estuarine organisms can be used to predict the consequences of habitat fragmentation on ecosystem function and to improve strategies for habitat and species conservation and restoration.     

Factors Mediating Co‐Occurrence of an Economically Valuable Introduced Fish and Its Native Frog Prey

Habitat characteristics mediate predator–prey coexistence in many ecological systems but are seldom considered in species introductions. When economically important introduced predators are stocked despite known negative impacts on native species, understanding the role of refuges, landscape configurations, and community interactions can inform habitat management plans. We measured these factors in basins with introduced trout (Salmonidae) and the Cascades frog (Rana cascadae) to determine, which are responsible for observed patterns of co‐occurrence of this economically important predator and its native prey. Large, vegetated shallows were strongly correlated to co‐occurrence, and R. cascadae larvae occur in shallower water when fish are present, presumably to escape predation. The number of nearby breeding sites of R. cascadae was also correlated to co‐occurrence, but only when the western toad (Anaxyrus boreas) was present. Because A. boreas larvae are unpalatable to fish and resemble R. cascadae, they may provide protection from trout via Batesian mimicry. Although rescue‐effect dispersal from nearby populations may maintain co‐occurrence, within‐lake factors proved more important for predicting co‐occurrence. Learning which factors allow co‐occurrence between economically important introduced species and their native prey enables managers to make better‐informed stocking decisions. Factores que Median la Co‐Ocurrencia de un Pez Introducido con Valor Económico y su Presa, una Rana Nativa     

Periodic Migration and Lowland Forest Refugia in a "Sedentary" Neotropical Bird, Wetmore's Bush-Tanager

Although the concept of tropical birds as sedentary is pervasive, evidence suggests many are not. Our grasp of movement status in tropical birds is decidedly poor, but the successful long-term conservation of these birds depends on such information. Sedentariness will likely doom much tropical avian diversity, but increased vagility is a two-edged sword: beneficial in promoting immigration, but detrimental in that more than one habitat may be required. Birds requiring more than one habitat may be unable to locate a particular type as landscape modifications increase. Our long-term data set from the Sierra de Los Tuxtlas in southern Veracruz, México, reveals infrequent, large-scale movements in a local highland endemic. Wetmore's Bush-Tanager ( Chlorospingus ophthalmicus wetmorei) seems occasionally dependent upon lowland forests (now greatly diminished) as a refugium from temporarily unsuitable highlands. Our data and observations lead us to three conclusions: 1) assumptions of sedentariness in tropical birds should be made with extreme caution; 2) normal, but periodic phenomena may be easily overlooked, even in relatively long-term studies; and 3) missing such phenomena jeopardizes the success of any conservation plan because over the long term a population may be dependent upon refugia only occasionally occupied.     

Predation, habitat complexity, and variation in density-dependent mortality of temperate reef fishes

Density dependence in demographic rates can strongly affect the dynamics of populations. However, the mechanisms generating density dependence (e.g., predation) are also dynamic processes and may be influenced by local conditions. Understanding the manner in which local habitat features affect the occurrence and/or strength of density dependence will increase our understanding of population dynamics in heterogeneous environments. In this study I conducted two separate field experiments to investigate how local predator density and habitat complexity affect the occurrence and form of density-dependent mortality of juvenile rockfishes (Sebastes spp.). I also used yearly censuses of rockfish populations on nearshore reefs throughout central California to evaluate mortality of juvenile rockfish at large spatial scales. Manipulations of predators (juvenile bocaccio, S. paucispinus) and prey (kelp, gopher, and black-and-yellow [KGB] rockfish, Sebastes spp.) demonstrated that increasing the density of predators altered their functional response and thus altered patterns of density dependence in mortality of their prey. At low densities of predators, the number of prey consumed per predator was a decelerating function, and mortality of prey was inversely density dependent. However, at high densities of predators, the number of prey killed per predator became an accelerating response, and prey mortality was directly density dependent. Results of field experiments and large-scale surveys both indicated that the strength of density-dependent mortality may also be affected by the structural complexity of the habitat. In small-scale field experiments, increased habitat complexity increased the strength of density-dependent mortality. However, at large scales, increasing complexity resulted in a decrease in the strength of density dependence. I suggest that these differences resulted from scale-dependent changes in the predatory response that generated mortality. Whether increased habitat complexity leads to an increase or a decrease in the strength of density-dependent mortality may depend on how specific predatory responses (e.g., functional or aggregative) are altered by habitat complexity. Overall, the findings of this study suggest that rates of demographic density dependence and the resulting dynamics of local populations may largely depend upon attributes of the local habitat.     

Improving accuracy of LiDAR-derived digital terrain models for saltmarsh management

Accurate digital elevation models of saltmarshes are crucial for both conservation and management goals. Light detection and ranging (LiDAR) is increasingly used for topographic surveys due to the ability to acquire high resolution data over spatially-extensive areas. This capability is ideally suited to saltmarsh environments, which are often vast, inaccessible systems where topographic variations can be very subtle. Derivation of surface (DSMs) (ground elevation plus vegetation) versus terrain (bare ground elevation) models (DTMs) relies on the ability of the LiDAR sensor to accurately record multiple returns. In saltmarshes however, the dense stands of low (< 1 m) vegetation commonly found precludes the acquisition of more than one return, and the resulting DTM is not different to the DSM. Establishing the offset between ground and vegetation surface in order to correct the LiDAR-derived DTM can be challenging due to the spatial variability in saltmarsh habitats. Here we show the development and application of a habitat-specific correction factor (HSCF) for the Odiel Saltmarshes using a combination of habitat object-based classification (82% overall accuracy) and ground control surveys that reduces the DTM error to within that associated with the LiDAR sensor (average error 0.1 m). We also show that the true accuracy of supplied (unmodified) DTMs can be >0.5 m in saltmarshes dominated by dense vegetation such as Spartina densiflora. In particular, global projections of sea-level rise across the next 80 years (0.18–0.59 m) significantly overlaps this accuracy margin, implying that assessments and modelling of sea-level impacts in saltmarsh systems will likely be erroneous if based on Lidar-derived DTMs. Erroneous assumptions and conclusions can result if the real accuracy of DTMs (bare ground) on vegetated saltmarshes is not considered, and the consequences of the propagation of this misinformation through to management decisions should not be over-looked.     

Global climate change: an opportunity for coastal dunes??

The predictions for coastal change under the scenario of global sea-level rise offer impending disaster for the variety of coastal morphologies, their associated habitats, and the accompanying infrastructure. However, the predictions tend to ignore the role of sediment budget in the maintenance of coastal morphology and the dynamics of sediment transfers in the beach-dune sand-sharing system. Accepting that shoreline displacement may be an outcome of sea-level rise and a negative sediment budget, conditions are presented that could lead to a positive or equilibrium sediment budget in the coastal foredune and the retention of the foredune system even as it is being displaced. Accommodation space is a key requirement for the continued functioning of the foredune morphologies during periods of sea-level rise.     

Predicted effects of climate change,vegetation and tree cover on dune slack habitats at Ainsdale on the Sefton Coast,UK

Dune slack habitats are highly dependent on the availability of water to support flora and fauna. Typically this is provided by shallow groundwater. This paper describes the seasonal and long term variation in groundwater levels in part of the Sefton coastline between 1972 and 2007. The effects of climate change, vegetation management and coastline realignment on groundwater levels are modelled. The observed annual water table levels rise and fall with an amplitude of 1.5 m, but longer term variations and trends are apparent. A stochastic water balance model was used to describe the changes in water table levels in slack floors in the open dunes and also in areas afforested with pine trees. It was found that the pine trees evaporated 214 mm/year more than open dunes vegetation, resulting in the water table being 0.5–1.0 m lower under the trees than under the open dunes. The effects of climate change on the ground water was simulated using predictions of future climate conditions based on the UKCIP02 medium high emissions scenario. The increase in temperature and change in rainfall patterns will result in a decrease in mean ground water levels by 1.0–1.5 mm in the next 90 years. Typical patterns consist of sequences of 5–10 years of low water table levels interspersed by infrequent sequences consisting of 2–5 years of relatively high or “normal” levels. These results indicate that that flora and fauna that cannot survive a 5–10 year period of water table levels >2.5 m below ground level are unlikely to survive or persist in many slack areas and a change in the ecology of these slack may become inevitable. Other effects of climate change include sea level rise which will result in a gradual rise in water table levels. Coastal erosion will increase the water table gradient to the sea and result in a slight lowering of the ground water levels. Conversely coastal accretion will reverse this process. The spatial distribution of coastal erosion and accretion along the Sefton coastline and its likely impacts on groundwater levels are discussed. The modelling work described in this paper has identified the factors which have the largest effect on groundwater levels in temperate coastal dune systems.     

Grizzly bear habitat selection is scale dependent.

The purpose of our study is to show how ecologists' interpretation of habitat selection by grizzly bears (Ursus arctos) is altered by the scale of observation and also how management questions would be best addressed using predetermined scales of analysis. Using resource selection functions (RSF) we examined how variation in the spatial extent of availability affected our interpretation of habitat selection by grizzly bears inhabiting mountain and plateau landscapes. We estimated separate models for females and males using three spatial extents: within the study area, within the home range, and within predetermined movement buffers. We employed two methods for evaluating the effects of scale on our RSF designs. First, we chose a priori six candidate models, estimated at each scale, and ranked them using Akaike Information Criteria. Using this method, results changed among scales for males but not for females. For female bears, models that included the full suite of covariates predicted habitat use best at each scale. For male bears that resided in the mountains, models based on forest successional stages ranked highest at the study-wide and home range extents, whereas models containing covariates based on terrain features ranked highest at the buffer extent. For male bears on the plateau, each scale estimated a different highest-ranked model. Second, we examined differences among model coefficients across the three scales for one candidate model. We found that both the magnitude and direction of coefficients were dependent upon the scale examined; results varied between landscapes, scales, and sexes. Greenness, reflecting lush green vegetation, was a strong predictor of the presence of female bears in both landscapes and males that resided in the mountains. Male bears on the plateau were the only animals to select areas that exposed them to a high risk of mortality by humans. Our results show that grizzly bear habitat selection is scale dependent. Further, the selection of resources can be dependent upon the availability of a particular vegetation type on the landscape. From a management perspective, decisions should be based on a hierarchical process of habitat selection, recognizing that selection patterns vary across scales.     

Conserving international coastal habitat networks on migratory waterfowl flyways

Migratory waterfowl depend on habitat networks at local, national and international scales for their survival. Coastal habitats are key areas for many waterfowl. Different species use different biotopes and in different places, so overall many parts of the coastal resource are important. As well as national conservation efforts, waterfowl conservation is increasingly focussed worldwide on collaborative international conservation, catalysed by several measures e.g. the Ramsar Convention, the EC Conservation of Wild Birds Directive, and the Bonn Convention Agreement on the Conservation of African/Eurasian Migratory Waterbirds. Several international conservation plans are under development for single species, but a more effective approach may be to develop plans, for assemblages of migratory birds with similar habitat requirements. All such plans must incorporate future sustainable use of the habitats on which the birds depend. Yet migratory bird and coastal habitat conservation is still often approached separately, despite the two being now closely linked to the development of the Natura 2000 site network in the European Community. Implementing the 1992 EC Habitats Directive requires the selection of coastal habitat sites for designation, set in national and international contexts of resource distribution. International coastal habitat inventories are needed to underpin this process. Combining such inventories with assessment of the flyway habitat requirements of waterfowl species and assemblages offers great potential for identifying international coastal habitat networks that meet the objectives of both habitat and migratory waterfowl conservation.     

Seasonal changes in the structure of an intertidal community

Quantitative (0.25 m²) samples of macrofaunal (>1.0 mm) invertebrates were taken in each season from one habitat of an intertidal sandbar in the North Inlet estuary near Georgetown, South Carolina, USA. During all seasons the community inhabiting the sample site was numerically dominated by two species of haustoriid amphipods (Acanthohaustorius millsi and Pseudohaustorius caroliniensis). Seasonal changes at the community level were clearly controlled by the population dynamics of the numerically dominant species, and qualitative information on life histories was important to the interpretation of analyses' results.This work was supported by the Environmental Technology Center of Martin Marietta Corporation and the Belle W. Baruch Institute for Marine Biology and Coastal Research. It is Contribution No. 138 of the Belle W. Baruch Institute for Marine Biology and Coastal research.     

An ecological model of the habitat mosaic in estuarine nursery areas: Part I—Interaction of dispersal theory and habitat variability in describing juvenile fish distributions

Identification of critical habitat in estuarine nursery areas is an important conservation and management objective. Habitat can be viewed as a mosaic of both temporally variable environmental features and spatially variable structural features that combine to define optimal habitat. Effective models of juvenile distributions should account for individual movement, as well as the full suite of habitat variability including both spatial and temporal components. We have extended a terrestrial model of small-scale movement patterns to describe habitat choices of an index juvenile fish in an estuarine nursery system. Movement of small juvenile fishes was found to be influenced by both spatial and temporal patterns in habitat quality, and it was a balanced mix of both that resulted in an optimal distribution. Fishes that perceive habitat on a scale much smaller than the scale of spatial heterogeneity may respond to temporal change as a movement cue allowing for more deterministic outcomes at larger scales despite perceptual limitations. These model outcomes suggest a hierarchical approach is best for describing habitat choice in juvenile fishes and this approach will be used in the future to explore individual and population responses to predictable habitat change.     

Simulating the fate of Florida Snowy Plovers with sea-level rise: Exploring research and management priorities with a global uncertainty and sensitivity analysis perspective

Changes in coastal habitats due to sea-level rise provide an uncertain, yet significant threat to shoreline dependent birds. Rising sea levels can cause habitat fragmentation and loss which can result in considerable reduction in their foraging and nesting areas. Computational models and their algorithmic assumptions play an integral role in exploring potential mitigation responses to uncertain and potentially adverse ecological outcomes. The presence of uncertainty in metapopulation models is widely acknowledged but seldom considered in their development and evaluation, specifically the effects of uncertain model inputs on the model outputs. This paper was aimed to (1) quantify the contribution of each uncertain input factor to the uncertainty in the output of a metapopulation model which evaluated the effects of long-term sea-level rise on the population of Snowy Plovers (Charadrius alexandrinus) found in the Gulf Coast of Florida, and (2) determine the ranges of model inputs that produced a specific output for the purpose of formulating environmental management decisions. This was carried out by employing global sensitivity and uncertainty analysis (GSA) using two generic (model independent) methods, the qualitative screening Morris method and a quantitative variance-based Sobol’ method coupled with Monte Carlo filtering. The analyses were applied to three density dependence scenarios: assuming a ceiling-type density dependence, assuming a contest-type density dependence, and assuming that density dependence is uncertain as to being ceiling- or contest-dependent. The sources of uncertainty in the outputs depended strongly on the type of density dependence considered in the model. In general, uncertainty in the outputs highly depended on the uncertainty in stage matrix elements (fecundity, adult survival, and juvenile survival), dispersal rate from central areas with low current populations (the “Big Bend” area of Florida) to the northern, panhandle populations, the maximum growth rate, and density dependence type. Our results showed that increasing the maximum growth rate to a value of 1.2 or larger will increase the final average population of Snowy Plovers assuming a contest-type density dependence. Results suggest that studies that further quantify which density dependence relationship best describes Snowy Plover population dynamics should be conducted since this is the main driver of uncertainty in model outcomes. Furthermore, investigating the presence of Snowy Plovers in the Big Bend region may be important for providing connection between the panhandle and peninsula populations.     

Evaluating the effects of foraging habitat restoration on shorebird reproduction: the importance of performance criteria and comparative design

Coastal development and engineering projects preclude ecosystem processes that provide habitat for beach nesting birds. Management for coastal species may depend on actions that attempt to restore important habitat features and mitigate disturbance effects. However, species response to restoration or other management actions may be difficult to predict or measure. At Jones Beach State Park, on Long Island, New York, a 0.49 ha restoration project provided moist substrate foraging habitat for breeding Piping Plovers (Charadrius melodus) from 2002 to 2005. We examined whether foraging habitat restoration affected Piping Plover breeding population size, productivity, and fledgling production within 300 m of the restoration site. We found a positive relationship between habitat restoration and the number of fledglings produced per year. However, foraging habitat restoration did not significantly increase the number of Plover pairs breeding at Jones Beach. Our ability to evaluate restoration effects on Plovers depended on: 1) use of multiple performance criteria; 2) a design that allowed comparison of pre- and post-restoration data; and 3) a spatial control that allowed comparison of similar areas that were near and far from the restoration site. Despite the small size of the restoration project, there were measurable benefits to Plovers, indicating that foraging habitat restoration may be an effective tool for species recovery.     

Predator recognition and anti-predator responses in the rainbowfish Melanotaenia eachamensis

Predator evasion behaviour patterns of three populations of rainbowfish (Melanotaenia eachamensis) were compared. The populations differed in the level of complexity of their natural habitats and the type and extent of predation. The predator recognition abilities of fish were assessed by exposing them to models differing in their degree of predator realism. The availability of vegetated cover and the location of the models with respect to cover were manipulated. Fish from Lake Tinaroo, a relatively open habitat containing numerous predators, showed strong changes in elective group size (EGS) in response to the different models but did not rely on cover as a place of refuge. In contrast, Dirran Creek fish originate from a small, fast-flowing, structurally complex stream lacking predatory fish species, and they showed little ability to distinguish between the different models and responded to threat by spending longer in vegetated areas. Members of the Lake Eacham captive stock increased their EGS in response to models representing low threat and with more threatening models increased the amount of time spent in vegetated regions of the arena. The contrasting reactions to predatory threat displayed by these populations highlights the need to use a number of different response indices when comparing the anti-predator responses of different fish populations. These data suggest that the level of habitat complexity as well as prior predator experience influence anti-predator responses of different fish popu-lations. Received: 2 October 1996 / Accepted after revision: 27 March 1997