Genetic variation among populations of the Antarctic toothfish: evolutionary insights and implications for conservation

Commercial fishing is having an increasingly negative impact on marine biodiversity, with over 70% of the world's fish stocks being fully exploited and, in many cases, overexploited. On top of this, the Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR) has granted commercial fishing permits in the most remote marine environment on earth, the high-latitude Southern Ocean. The primary target of these new commercial fishing ventures is the large pelagic piscivorous predator, the Antarctic toothfish (Dissostichus mawsoni). Unfortunately, little information is available on the demography, genetics, or life history of this large fish. Without such information we have little idea as to the effects of commercial fishing on the population structure and survival of this species. In this study, we focus on patterns of genetic diversity within and between geographically disparate populations of the Antarctic toothfish, using randomly amplified polymorphic DNA markers. Results of our study showed high levels of genetic similarity within and between populations. Despite high levels of genetic similarity, genetic analyses detected significant population structure, including fixed differences among populations, a significant fixation index (Fst) and between-population differentiation via a Mantel test. From a conservation perspective, low levels of genetic diversity may be indicative of relatively small populations that would not be able to withstand heavy commercial fishing pressures. Given that there is evidence for significant genetic structure, it will be important to manage these fisheries in a manner that will help prevent the loss of unique genetic variation from regional overfishing.     

Responses of population structure and genomic diversity to climate change and fishing pressure in a pelagic fish

The responses of marine species to environmental changes and anthropogenic pressures (e.g., fishing) interact with ecological and evolutionary processes that are not well understood. Knowledge of changes in the distribution range and genetic diversity of species and their populations into the future is essential for the conservation and sustainable management of resources. Almaco jack (Seriola rivoliana) is a pelagic fish with high importance to fisheries and aquaculture in the Pacific Ocean. In this study, we assessed contemporary genomic diversity and structure in loci that are putatively under selection (outlier loci) and determined their potential functions. Using a combination of genotype–environment association, spatial distribution models, and demogenetic simulations, we modeled the effects of climate change (under three different RCP scenarios) and fishing pressure on the species' geographic distribution and genomic diversity and structure to 2050 and 2100. Our results show that most of the outlier loci identified were related to biological and metabolic processes that may be associated with temperature and salinity. The contemporary genomic structure showed three populations—two in the Eastern Pacific (Cabo San Lucas and Eastern Pacific) and one in the Central Pacific (Hawaii). Future projections suggest a loss of suitable habitat and potential range contractions for most scenarios, while fishing pressure decreased population connectivity. Our results suggest that future climate change scenarios and fishing pressure will affect the genomic structure and genotypic composition of S. rivoliana and lead to loss of genomic diversity in populations distributed in the eastern-central Pacific Ocean, which could have profound effects on fisheries that depend on this resource.     

Integrating local plant resources and habitat management

The use of wild plant resources as a source of basic needs is an important aspect of multiple-use of land in much of Africa, ranging from vegetation with a low species diversity, high biomass production and resilience to harvesting (e.g.Phragmites reedbeds,Cymbopogon grasslands) through to vegetation with a high diversity of species (and plant life-forms), with a multiple of uses and often low resilience to resource harvesting (e.g. medicinal plants, pole cutting, fuelwood from Afro-montane forest). Complexity and costs of managing sustainable use of wild populations increase markedly with an increasing number of uses and resource users. It is suggested that if the primary objective of core conservation areas is accepted to be the maintenance of habitat and species diversity, then the limited money and manpower available for management of core conservation areas in southern Africa limits sustainable harvesting use of plant resources to low diversity, low conservation priority vegetation types or encroaching species. For high diversity, high conservation priority sites such as Afro-montane or Coastal evergreen forest, the emphasis must be on providing alternative sources of supply to resource users outside of core areas. Botanical gardens staff, with their horticultural experience, can play a very important role through bulking up material for supply to small farmers, herbalists and introducing additional species for agro-forestry. Botanical gardens can play a greater role in the establishment of field gene banks andex situ conservation of vulnerable species. They can also provide a valuable educational and research role on mass production techniques to boost local stocks of threatened and commercially valuable species for cultivation by local people, whether farmers or specialist users (e.g. herbalists) in the country of origin as a means of generating employment and restoring local self-sufficiency.     

Effect of Biodiversity Changes in Disease Risk: Exploring Disease Emergence in a Plant-Virus System

The effect of biodiversity on the ability of parasites to infect their host and cause disease (i.e. disease risk) is a major question in pathology, which is central to understand the emergence of infectious diseases, and to develop strategies for their management. Two hypotheses, which can be considered as extremes of a continuum, relate biodiversity to disease risk: One states that biodiversity is positively correlated with disease risk (Amplification Effect), and the second predicts a negative correlation between biodiversity and disease risk (Dilution Effect). Which of them applies better to different host-parasite systems is still a source of debate, due to limited experimental or empirical data. This is especially the case for viral diseases of plants. To address this subject, we have monitored for three years the prevalence of several viruses, and virus-associated symptoms, in populations of wild pepper (chiltepin) under different levels of human management. For each population, we also measured the habitat species diversity, host plant genetic diversity and host plant density. Results indicate that disease and infection risk increased with the level of human management, which was associated with decreased species diversity and host genetic diversity, and with increased host plant density. Importantly, species diversity of the habitat was the primary predictor of disease risk for wild chiltepin populations. This changed in managed populations where host genetic diversity was the primary predictor. Host density was generally a poorer predictor of disease and infection risk. These results support the dilution effect hypothesis, and underline the relevance of different ecological factors in determining disease/infection risk in host plant populations under different levels of anthropic influence. These results are relevant for managing plant diseases and for establishing conservation policies for endangered plant species.     

Genomic investigation of the Chinese alligator reveals wild-extinct genetic diversity and genomic consequences of their continuous decline

Critically endangered species are usually restricted to small and isolated populations. High inbreeding without gene flow among populations further aggravates their threatened condition and reduces the likelihood of their long-term survival. Chinese alligator (Alligator sinensis) is one of the most endangered crocodiles in the world and has experienced a continuous decline over the past c. 1 million years. In order to identify the genetic status of the remaining populations and aid conservation efforts, we assembled the first high-quality chromosome-level genome of Chinese alligator and explored the genomic characteristics of three extant breeding populations. Our analyses revealed the existence of at least three genetically distinct populations, comprising two breeding populations in China (Changxing and Xuancheng) and one breeding population in an American wildlife refuge. The American population does not belong to the last two populations of its native range (Xuancheng and Changxing), thus representing genetic diversity extinct in the wild and provides future opportunities for genetic rescue. Moreover, the effective population size of these three populations has been continuously declining over the past 20 ka. Consistent with this decline, the species shows extremely low genetic diversity, a large proportion of long runs of homozygous fragments, and mutational load across the genome. Finally, to provide genomic insights for future breeding management and conservation, we assessed the feasibility of mixing extant populations based on the likelihood of introducing new deleterious alleles and signatures of local adaptation. Overall, this study provides a valuable genomic resource and important genomic insights into the ecology, evolution, and conservation of critically endangered alligators.     

Effect of new founders on retention of gene diversity in captive populations: A formalization of the nucleus population concept

A nucleus population is a small captive population genetically supported by periodic importation of wild caught animals. Periodic importation will allow nucleus populations to maintain the same amount of gene diversity as larger captive populations that do not import wild caught animals. The function of nucleus populations as envisioned by the IUCN/SSC Captive Breeding Specialist Group (CBSG) is to make additional captive space available for endangered taxa not currently maintained in captivity. In this article, mathematical models are developed to assess the effectiveness of the nucleus population concept in reducing the population sizes necessary to maintain appreciable amounts of gene diversity in captive populations. It is shown that the Nucleus I population concept, as defined and promoted by the CBSG, requires an importation rate 10–20 times greater than they have indicated. Whereas nucleus populations are not appropriate for maintenance of significant amounts of gene diversity in long-term breeding programs, small populations can be valuable for research, education, and reintroduction projects with short-term goals. Decisions have to be made on which of the many endangered taxa will be maintained and for what purposes, if captive breeding is to be an effective component of species conservation. © 1993 Wiley-Liss, Inc.     

A multi-faceted framework of diversity for prioritizing the conservation of fish assemblages

Floodplain waterbodies and their biodiversity are increasingly threatened by human activities. Given the limited resources available to protect them, methods to identify the most valuable areas for biodiversity conservation are urgently needed. In this study, we used freshwater fish assemblages in floodplain waterbodies to propose an innovative method for selecting priority areas based on four aspects of their diversity: taxonomic (i.e. according to species classification), functional (i.e. relationship between species and ecosystem processes), natural heritage (i.e. species threat level), and socio-economic (i.e. species interest to anglers and fishermen) diversity. To quantitatively evaluate those aspects, we selected nine indices derived either from metrics computed at the species level and then combined for each assemblage (species rarity, origin, biodiversity conservation concern, functional uniqueness, functional originality, fishing interest), or from metrics directly computed at the assemblage level (species richness, assemblage rarity, diversity of biological traits). Each of these indices belongs to one of the four aspects of diversity. A synthetic index defined as the sum of the standardized aspects of diversity was used to assess the multi-faceted diversity of fish assemblages. We also investigated whether the two main environmental gradients at the catchment (distance from the sea) and at the floodplain (lateral connectivity of the waterbodies) scales influenced the diversity of fish assemblages, and consequently their potential conservation value. Finally, we propose that the floodplain waterbodies that should be conserved as a priority are those located in the downstream part of the catchment and which have a substantial lateral connectivity with the main channel.     

Estimation of survival rate and extinction probability for stage-structured populations with overlapping life stages

The development of methods providing reliable estimates of demographic parameters (e.g., survival rates, fecundity) for wild populations is essential to better understand the ecology and conservation requirements of individual species. A number of methods exist for estimating the demographics of stage-structured populations, but inherent mathematical complexity often limits their uptake by conservation practitioners. Estimating survival rates for pond-breeding amphibians is further complicated by their complex migratory and reproductive behaviours, often resulting in nonobservable states and successive cohorts of eggs and tadpoles. Here we used comprehensive data on 11 distinct breeding toad populations (Bufo calamita) to clarify and assess the suitability of a relatively simple method [the Kiritani–Nakasuji–Manly (KNM) method] to estimate the survival rates of stage-structured populations with overlapping life stages. The study shows that the KNM method is robust and provides realistic estimates of amphibian egg and larval survival rates for species in which breeding can occur as a single pulse or over a period of several weeks. The study also provides estimates of fecundity for seven distinct toad populations and indicates that it is essential to use reliable estimates of fecundity to limit the risk of under- or overestimating the survival rates when using the KNM method. Survival and fecundity rates for B. calamita populations were then used to define population matrices and make a limited exploration of their growth and viability. The findings of the study recently led to the implementation of practical conservation measures at the sites where populations were most vulnerable to extinction.     

Biogeography of Parasitic Nematode Communities in the Galápagos Giant Tortoise: Implications for Conservation Management

The Galápagos giant tortoise is an icon of the unique, endemic biodiversity of Galápagos, but little is known of its parasitic fauna. We assessed the diversity of parasitic nematode communities and their spatial distributions within four wild tortoise populations comprising three species across three Galápagos islands, and consider their implication for Galápagos tortoise conservation programmes. Coprological examinations revealed nematode eggs to be common, with more than 80% of tortoises infected within each wild population. Faecal samples from tortoises within captive breeding centres on Santa Cruz, Isabela and San Cristobal islands also were examined. Five different nematode egg types were identified: oxyuroid, ascarid, trichurid and two types of strongyle. Sequencing of the 18S small-subunit ribosomal RNA gene from adult nematodes passed with faeces identified novel sequences indicative of rhabditid and ascaridid species. In the wild, the composition of nematode communities varied according to tortoise species, which co-varied with island, but nematode diversity and abundance were reduced or altered in captive-reared animals. Evolutionary and ecological factors are likely responsible for the variation in nematode distributions in the wild. This possible species/island-parasite co-evolution has not been considered previously for Galápagos tortoises. We recommend that conservation efforts, such as the current Galápagos tortoise captive breeding/rearing and release programme, be managed with respect to parasite biogeography and host-parasite co-evolutionary processes in addition to the biogeography of the host.     

Comparative genetic diversity of wild and hatchery-produced Pacific oyster (Crassostrea gigas) populations in Korea using multiplex PCR assays with nine polymorphic microsatellite markers

The Pacific oyster, Crassostrea gigas, is the most important and valuable commercial fishery species in Korea. Its farming started 20 years ago and is still rapid expansion in Korea. In this study, to maintain the genetic diversity of this valuable marine resource, possible genetic similarity and differences between the wild population and hatchery population in Tongyeong, Korea were accessed using multiplex assays with nine highly polymorphic microsatellite loci. A total of 250 different alleles were found over all loci. Despite a long history of hatchery practices, very high levels of polymorphism (mean alleles = 22.89 and mean heterozygosity = 0.92) were detected between the two populations. No statistically significant reductions were found in heterozygosity or allelic diversity in the hatchery population compared with the wild population. However, significant genetic heterogeneity was found between two populations. These results provide no evidence to show that hatchery practice of Pacific oyster in Korea has significantly affected the genetic variability of the hatchery stock. Although further studies are needed for comprehensive determinations of the hatchery and wild populations with increased number of Pacific oyster sample collections, information on the genetic variation and differentiation obtained in this study can be applied for genetic monitoring of aquaculture stocks, genetic improvement by selective breeding and designing of more efficient conservation management guidelines for these valuable genetic materials.     

SOS!濒临极限的生物多样性

本文论述了生物多样性的价值、多样性丧失的严重性和后果,以及多样性保护与持续利用的对策。文中强调指出,生物界中的每片基因,每一个物种,每一类生态系统对人类的持久生存都是无价之宝,任何多样性的丧失都是不可逆、不可再生的,因而对人类的损失是难以估量的。然而目前由于人类剧烈的活动干扰如滥砍滥伐、滥捕滥猎、环境污染、火灾、垦荒等,生物多样性丧失的速率怵目惊心！若不赶快行动起来,人类赖以生存的生物多样性将所剩无几,人类生存的危机也将难以避免。从保护与利用及其协同发展出发,本文呼吁要大力开展生物多样性的研究和开发工作,目的一方面在于进一步加强生物多样性的基础调查和研究工作,另一方面在于使生物多样性资源更好地为人类造福。为减缓目前生物多样性所承受的压力及促进其恢复,本文建议对完全依赖野生生物资源的传统产业征收“资源更新税”;而对开发和利用生物多样性资源,不仅不影响野生资源,而且能替代它,或减缓其压力,或促进其恢复的高新技术产业,在税收上给予特别优惠。     

Macrogeographic variation of a pond predator's top-down effects in a common garden environment

Geographic variation in species behavior and life history has been well documented in biology. Species with wide geographic distributions (i.e., across a continent) but small home ranges (i.e., <1 km²) likely experience wide variability in abiotic environments across the entirety of their range, possibly exhibiting strong local adaptation. Understanding variation across a large geographic scale is especially important when considering species that have strong ecological importance, such as keystone species. Yet, few studies have compared the potential cascading ecological effects of a predator with a keystone role in at least part of its range. To understand how keystone ability in pond food webs can vary across a large geographic range, we conducted an artificial pond experiment with a known keystone predator in at least part of its range, the marbled salamander (Ambystoma opacum). To do so, we collected size-matched salamander larvae from three geographically distant populations (>650 km apart) in Ohio, Mississippi, and North Carolina and placed them in mesocosms with a suite of spring breeding amphibian prey species. We observed differential survival of some prey species leading to differences in spring-breeding amphibian diversity among the three predator populations, indicating that keystone predation may vary at a geographic scale. Prey diversity was lowest with predators from northern (Ohio) populations of salamanders. Further understanding of large-scale variability in ecologically important predators and the potential effects of translocating wide-ranging ambystomatid species is needed to direct future conservation efforts and preserve biodiversity.     

Traditional home-garden conserving genetic diversity: a case study of Acacia pennata in southwest China

Conserving biodiversity in human-dominated systems requires research into mechanisms that can maintain biodiversity in fragmented landscapes. Home-garden as traditional agroforestry system in many regions has shown great value in maintaining a wide range of species. Here we show that home-garden populations are also capable of maintaining high level of genetic variation. Using six polymorphic microsatellite DNA markers, we have genotyped 260 individuals of Acacia pennata, a popular wild vegetable in the tropical region of southeast Asia. Samples were collected from home-gardens and wild populations in Xishuangbanna, southwest China. Microsatellite DNA diversity in planted populations were compared with that in geographically nearby wild populations with similar population size. Over 90?% of microsatellite genetic variation in wild populations was also present in planted populations. Pairwise comparison of planted and adjacent wild population showed no significant difference in allelic diversity and heterozygosity. Analysis revealed no significant genetic differences between wild and planted populations, while four home-garden populations showed sign of bottleneck. We conclude that home-gardens show great promise in maintaining genetic diversity, and that these managed patches could be of significant conservation value in tropical regions.     

Ecological impacts of non-native Pacific oysters (<Emphasis Type="Italic">Crassostrea gigas</Emphasis>) and management measures for protected areas in Europe

Pacific oysters are now one of the most ‘globalised’ marine invertebrates. They dominate bivalve aquaculture production in many regions and wild populations are increasingly becoming established, with potential to displace native species and modify habitats and ecosystems. While some fishing communities may benefit from wild populations, there is now a tension between the continued production of Pacific oysters and risk to biodiversity, which is of particular concern within protected sites. The issue of the Pacific oyster therefore locates at the intersection between two policy areas: one concerning the conservation of protected habitats, the other relating to livelihoods and the socio-economics of coastal aquaculture and fishing communities. To help provide an informed basis for management decisions, we first summarise evidence for ecological impacts of wild Pacific oysters in representative coastal habitats. At local scales, it is clear that establishment of Pacific oysters can significantly alter diversity, community structure and ecosystem processes, with effects varying among habitats and locations and with the density of oysters. Less evidence is available to evaluate regional-scale impacts. A range of management measures have been applied to mitigate negative impacts of wild Pacific oysters and we develop recommendations which are consistent with the scientific evidence and believe compatible with multiple interests. We conclude that all stakeholders must engage in regional decision making to help minimise negative environmental impacts, and promote sustainable industry development.     

From Sea to Sea: Canada's Three Oceans of Biodiversity

Evaluating and understanding biodiversity in marine ecosystems are both necessary and challenging for conservation. This paper compiles and summarizes current knowledge of the diversity of marine taxa in Canada''s three oceans while recognizing that this compilation is incomplete and will change in the future. That Canada has the longest coastline in the world and incorporates distinctly different biogeographic provinces and ecoregions (e.g., temperate through ice-covered areas) constrains this analysis. The taxonomic groups presented here include microbes, phytoplankton, macroalgae, zooplankton, benthic infauna, fishes, and marine mammals. The minimum number of species or taxa compiled here is 15,988 for the three Canadian oceans. However, this number clearly underestimates in several ways the total number of taxa present. First, there are significant gaps in the published literature. Second, the diversity of many habitats has not been compiled for all taxonomic groups (e.g., intertidal rocky shores, deep sea), and data compilations are based on short-term, directed research programs or longer-term monitoring activities with limited spatial resolution. Third, the biodiversity of large organisms is well known, but this is not true of smaller organisms. Finally, the greatest constraint on this summary is the willingness and capacity of those who collected the data to make it available to those interested in biodiversity meta-analyses. Confirmation of identities and intercomparison of studies are also constrained by the disturbing rate of decline in the number of taxonomists and systematists specializing on marine taxa in Canada. This decline is mostly the result of retirements of current specialists and to a lack of training and employment opportunities for new ones. Considering the difficulties encountered in compiling an overview of biogeographic data and the diversity of species or taxa in Canada''s three oceans, this synthesis is intended to serve as a biodiversity baseline for a new program on marine biodiversity, the Canadian Healthy Ocean Network. A major effort needs to be undertaken to establish a complete baseline of Canadian marine biodiversity of all taxonomic groups, especially if we are to understand and conserve this part of Canada''s natural heritage.     

Protected and threatened components of fish biodiversity in the Mediterranean sea

The Mediterranean Sea (0.82% of the global oceanic surface) holds 4%-18% of all known marine species (~17,000), with a high proportion of endemism [1, 2]. This exceptional biodiversity is under severe threats [1] but benefits from a system of 100 marine protected areas (MPAs). Surprisingly, the spatial congruence of fish biodiversity hot spots with this MPA system and the areas of high fishing pressure has not been assessed. Moreover, evolutionary and functional breadth of species assemblages [3] has been largely overlooked in marine systems. Here we adopted a multifaceted approach to biodiversity by considering the species richness of total, endemic, and threatened coastal fish assemblages as well as their functional and phylogenetic diversity. We show that these fish biodiversity components are spatially mismatched. The MPA system covers a small surface of the Mediterranean (0.4%) and is spatially congruent with the hot spots of all taxonomic components of fish diversity. However, it misses hot spots of functional and phylogenetic diversity. In addition, hot spots of endemic species richness and phylogenetic diversity are spatially congruent with hot spots of fishery impact. Our results highlight that future conservation strategies and assessment efficiency of current reserve systems will need to be revisited after deconstructing the different components of biodiversity.     

Perspectives of genomics for genetic conservation of livestock

Genomics provides new opportunities for conservation genetics. Conservation genetics in livestock is based on estimating diversity by pedigree relatedness and managing diversity by choosing those animals that maximize genetic diversity. Animals can be chosen as parents for the next generation, as donors of material to a gene bank, or as breeds for targeting conservation efforts. Genomics provides opportunities to estimate diversity for specific parts of the genome, such as neutral and adaptive diversity and genetic diversity underlying specific traits. This enables us to choose candidates for conservation based on specific genetic diversity (e.g. diversity of traits or adaptive diversity) or to monitor the loss of diversity without conservation. In wild animals direct genetic management, by choosing candidates for conservation as in livestock, is generally not practiced. With dense marker maps opportunities exist for monitoring relatedness and genetic diversity in wild populations, thus enabling a more active management of diversity.     

The potential for large carnivores to act as biodiversity surrogates in southern Africa

Biodiversity in southern Africa is globally extraordinary but threatened by human activities. Although there are considerable biodiversity conservation initiatives within the region, no one has yet assessed the potential use of large carnivores in such actions. Surrogate approaches have often been suggested as one such way of capitalizing on large carnivores. Here we review the suitability of the large carnivore guild (i.e., brown hyaena Hyaena hyaena, spotted hyaena Crocuta crocutta, cheetah Acinonyx jubatus, leopard Panthera pardus, lion Panthea leo and African wild dog Lycaon pictus) to act as surrogate species for biodiversity conservation in southern Africa. We suggest that the guild must be complete for the large carnivores to fully provide their role as ecological keystones. The potential for large carnivores to act as umbrella and indicator species seems limited. However, self-sustaining populations of large carnivores may be useful indicators of unfragmented landscapes. Moreover, diversity within the large carnivore guild may reflect overall biodiversity. Although the global appeal of the large African carnivores makes them important international flagships, we stress that international conservation funding must be linked to local communities for them to be important also locally. In summary, we suggest that the flagship value of these large carnivores should be used to promote biodiversity conservation in the region, and that the suggested relationship between large carnivore diversity and overall biodiversity is empirically tested. Finally we suggest that direct conservation activities should focus on enhancing the keystone values of large carnivores through complete guild conservation and restoration.     

A review of conservation status of freshwater fish diversity in China

China harbors a high species diversity of freshwater fishes not shared with any of its neighboring nations. Freshwater fish diversity in the country has been under severe threat from human activities over the past decades, thus conservation freshwater fishes and ecosystems is urgently needed. To accumulate baseline data for guiding protection actions, the third red list assessment of Chinese freshwater fishes was carried out. Among Chinese freshwater fishes assessed, there are 355 at-risk species (22.3% of the total), including 69 ranked as Critically Endangered, 97 as Endangered, and 189 as Vulnerable. Two species are classified as Extinct and one as Regionally Extinct. China's threat level seems to be lower than the known average level found in the IUCN's global assessment of freshwater fishes, but this is an artifact of a high rate of species classified as Data Deficient. Conservation of freshwater fishes is presently facing a grim situation in China. Imperilment of Chinese freshwater fishes is primarily attributed to habitat loss and degradation arising from human perturbations, particularly river damming. Despite the adoption of protected areas setting up, captive breeding and release, and a fishing moratorium, conservation efforts for freshwater fishes are compromised by disproportional attention in China's biodiversity conservation, baseline data deficiency, insufficiently designed protection networks, and inefficient or inadequate implementation of conservation strategies. To achieve the objectives of Chinese freshwater fish conservation, it is proposed to conduct a national-scale survey of fish diversity and reassess their at-risk status, develop systematic conservation planning of freshwater fish diversity and ecosystems, prioritize strategies for protected areas development, perform genetic-based captive breeding for releasing in concert with other protection actions, and implement flexible fishing moratorium strategies in different water bodies.