Increasing variation in taxonomic distinctness reveals clusters of specialists in the deep sea

Changes in biodiversity with latitude or along a given environmental gradient have been described in many studies, including for marine ecosystems. Currently there is no scientific consensus, however, regarding macroecological patterns of diversity vs depth. Here, we describe variation in the biodiversity of fishes along a depth gradient from 0 to 2000 m in the region of the Norfolk Ridge and Lord Howe Rise (Western Pacific), using data obtained during the NORFANZ voyage. We modelled α diversity (richness), β diversity (using Jaccard's coefficient), evenness, taxonomic distinctness and taxonomic resemblances among fish communities. Although α diversity did not change appreciably with depth, β diversity decreased significantly in deeper strata. Both taxonomic resemblances and Jaccard similarities diminished with depth, indicating convergence in community structure. In addition, average taxonomic distinctness showed no clear pattern with depth, but taxonomic trees constructed among species within deeper samples had more variable path‐lengths than those in shallower samples. The presence of taxonomically distinct clusters of highly related species at depth indicates specialised niches that have developed in a relatively extreme (dark, pressurized) yet stable environment. We propose that reduced β diversity and increased variation in taxonomic distinctness might serve as indicators of ecological communities living in harsh environments – a hypothesis that should be tested in other systems, such as deserts, high altitudes or latitudes.     

Linking ecology with parasite diversity in Neotropical fishes

A comparative analysis was performed to seek large-scale patterns in the relationships between a set of fish species traits (body size, type of environment, trophic level, schooling behaviour, depth range, mean habitat temperature, geographical range, ability to enter brackish waters and capability of migration) and the diversity of their metazoan parasite assemblages among 651 Neotropical fish species. Two measurements of parasite diversity are used: the species richness and the taxonomic distinctness of a fish's parasite assemblage, including all metazoan parasites, ectoparasites only, or endoparasites only. The results showed that, on this scale, the average taxonomic distinctness of parasite assemblages was clearly more sensitive to the influence of host traits than parasite species richness. Differences in the taxonomic diversification of the parasite assemblages of different fish species were mainly related to the fish's environment (higher values in benthic–demersal species), trophic level (positive correlation with increasing level), temperature (positive correlation with temperature in marine ectoparasites, negative in endoparasites; positive for all groups of parasites in freshwater fishes) and oceanic distribution (higher values in fish species from the Pacific Ocean than those of the Atlantic). The results suggest that, among Neotropical fish species, only certain key host traits have influenced the processes causing the taxonomic diversification of parasite assemblages.     

Land‐use effects on the functional distinctness of arthropod communities

Land‐use change is a major driver of the global loss of biodiversity, but it is unclear to what extent this also results in a loss of ecological traits. Therefore, a better understanding of how land‐use change affects ecological traits is crucial for efforts to sustain functional diversity. To this end we tested whether higher species richness or taxonomic distinctness generally leads to increased functional distinctness and whether intensive land use leads to functionally more narrow arthropod communities. We compiled species composition and trait data for 350 species of terrestrial arthropods (Araneae, Carabidae and Heteroptera) in different land‐use types (forests, grasslands and arable fields) of low and high land‐use intensity. We calculated the average functional and taxonomic distinctness and the rarified trait richness for each community. These measures reflect the range of traits, taxonomic relatedness and number of traits that are observed in local communities. Average functional distinctness only increased significantly with species richness in Carabidae communities. Functional distinctness increased significantly with taxonomic distinctness in communities of all analyzed taxa suggesting a high functional redundancy of taxonomically closely related species. Araneae and Heteroptera communities had the expected lower functional distinctness at sites with higher land‐use intensity. More frequently disturbed land‐use types such as managed grasslands or arable fields were characterized by species with smaller body sizes and higher dispersal abilities and communities with lower functional distinctness or trait richness. Simple recommendations about the conservation of functional distinctness of arthropod communities in the face of future land‐use intensification and species loss are not possible. Our study shows that these relationships depend on the studied taxa and land‐use type. However, for some arthropod groups functional distinctness is threatened by intensification and conversion from less to more frequently disturbed land‐uses.     

Taxonomic distinctness in a linear system: a test using a tropical freshwater fish assemblage

Taxonomic distinctness, a measure of diversity that captures the phylogenetic relatedness of an assemblage, has been successfully applied in several marine communities and appears to have considerable potential for environmental assessment and conservation. This approach has, however, not hitherto been used in structured freshwater systems. We evaluated this technique in the context of vulnerable tropical river drainages in India. We found that the inherent physical structure of river systems and longitudinal gradients in habitat characteristics influenced the pattern of taxonomic distinctness. Downstream locations were in general more taxonomically diverse than upstream ones. This in turn means that highly perturbed sites in the lower reaches of rivers would not display a reduction in taxonomic distinctness relative to a random expectation. The utility of the approach for detecting habitat and other disturbances on stream fish communities is thus limited. Implications of these unique properties of river ecosystems and their communities to conservation are discussed.     

鄱阳湖鱼类多样性的时空变化特征研究

研究基于2010年4月、7月和2018年8月、2019年5月在鄱阳湖9个区域的鱼类多样性调查数据,分析鱼类物种多样性、功能多样性和分类差异指数的时空变化以及其与环境之间的关系,了解鄱阳湖近十年来鱼类群落多样性的时空变化特征。结果显示, 2010年和2018—2019年分别调查到鱼类74种和93种,群落结构差异显著(P<0.05),差异贡献率最高的物种为短颌鲚、似鳊、鲫、光泽黄颡鱼和鲤;水温、总悬浮物和叶绿素等环境因素具有显著的年际和季节差异(P<0.05)。与2010年相比, 2018—2019年鱼类物种多样性和功能多样性指数有一定增加,分类差异指数没有显著变化。分类差异指数的随机检验显示(Randomization test),与2010年相比, 2018—2019年位于95%概率置信范围下方的区域增加。水温、总悬浮物和叶绿素等环境因素对物种多样性、功能多样性和分类差异指数有显著的影响(P<0.05)。结果表明,近十年来,鄱阳湖鱼类群落结构发生明显改变,但是小型鱼类依旧是优势种,鱼类群落小型化明显,主要原因可能是过度捕捞的影响。同时,鄱阳湖的人类活动干扰增大,鄱阳县...     

The Evolution of Taxonomic Diversity in Helminth Assemblages of Mammalian Hosts

Several studies have searched for the key forces behind the diversification of parasite assemblages over evolutionary time. All of these studies have used parasite species richness as their measure of diversity, thus ignoring the relatedness among parasite species and the taxonomic structure of the assemblages. This information is essential, however, if we want to elucidate which processes have caused an assemblage of parasites to acquire new species. Here, we performed a comparative analysis across 110 species of mammalian hosts in which we evaluated the effects of four host traits (body mass, population density, geographic range, and basal metabolic rate) on the diversity of their assemblages of helminth endoparasites. As measures of diversity, we used parasite species richness, as well as the average taxonomic distinctness of the assemblage and its variance; the latter measures are based on the taxonomic distance between two parasite species, computed across all possible species pairs in an assemblage. Unlike parasite species richness, both the average taxonomic distinctness and its variance were unaffected by the number of hosts examined. These two measures of parasite diversity also proved highly repeatable among host populations of the same mammalian species; in contrast, parasite species richness was unreliable as a species character, as it varied as much within a host species than among different host species. Using phylogenetically independent contrasts, and correcting for potential confounding variables, we found that host population density correlated positively with parasite species richness. There were, however, no other relationships between any of the four host traits investigated and either of our measures of parasite diversity. The processes facilitating the taxonomic diversification of parasite assemblages thus remain unclear, but their elucidation will be necessary if we are to fully understand parasite evolution.     

江湖阻隔对长江中下游湖泊鱼类群落分类多样性的影响

基于1950s以来的长江中下游湖泊鱼类调查数据,分析通江湖泊与阻隔湖泊的鱼类分类多样性差异,以及通江和阻隔湖泊鱼类分类多样性的时间序列变化,探讨江湖阻隔对鱼类多样性的影响。结果显示,阻隔湖泊鱼类物种数、平均分类差异指数(Δ⁺)和分类差异变异指数(Λ⁺)平均值分别为48.47±14.64、74.02±3.09和736.89±33.80;通江湖泊为76.22±14.40、78.31±0.98和697.31±25.53。阻隔湖泊物种数和Δ⁺值显著低于通江湖泊(P<0.001),而阻隔湖泊Λ⁺值显著高于通江湖泊(P=0.002),表明阻隔湖泊物种间亲缘关系更近,均匀度下降,即物种分类单元减少,且集中分布于某几个分类阶元,稳定性变差。典型通江与阻隔湖泊鱼类群落分类多样性的时间变化分析发现,两种类型湖泊的鱼类物种数和Δ⁺值均随时间推移整体呈现下降趋势,Λ⁺值整体呈现升高趋势;并且阻隔湖泊的Λ+值随阻隔时间增加而大幅上升,Δ⁺和Λ^+...     

Predicting the similarity of parasite communities in freshwater fishes using the phylogeny,ecology and proximity of hosts

Parasite communities tend to be dissimilar in hosts that are geographically, phylogenetically, ecologically and developmentally distant from one another. The decay of community similarity is a powerful and increasingly common method of studying parasite beta diversity, but most studies have examined only a single type of distance. Here, we evaluate distances based on the phylogeny, ecology, spatial proximity and size of hosts, as predictors of the similarity of parasite communities in individual hosts, host populations and host species. We surveyed parasites in six species of fish collected simultaneously from six localities in the St. Lawrence River, Canada, and species in a common group of larval parasites were discriminated using DNA sequences from barcode region of cytochrome c oxidase I. Distances based on the habitat use patterns of host species were good predictors of short‐term, ecological similarity of parasite communities, such as that operating at the scale of the individual host. The genetic distance between host species was associated with almost all types of similarity at all scales, particularly qualitative and phylogenetic similarity of parasite communities at the level of populations and meta‐populations of hosts. The trophic level, diet, spatial proximity and size of hosts were poor predictors of parasite community similarity. The increased taxonomic resolution provided by molecular data increased the explanatory power of regression models, and different factors were implicated when parasite species were distinguished with DNA barcodes than when larval parasites were lumped into morphospecies, as is commonly practiced.     

Habitat dynamics,marine reserve status,and the decline and recovery of coral reef fish communities

Severe climatic disturbance events often have major impacts on coral reef communities, generating cycles of decline and recovery, and in some extreme cases, community‐level phase shifts from coral‐ to algal‐dominated states. Benthic habitat changes directly affect reef fish communities, with low coral cover usually associated with low fish diversity and abundance. No‐take marine reserves (NTRs) are widely advocated for conserving biodiversity and enhancing the sustainability of exploited fish populations. Numerous studies have documented positive ecological and socio‐economic benefits of NTRs; however, the ability of NTRs to ameliorate the effects of acute disturbances on coral reefs has seldom been investigated. Here, we test these factors by tracking the dynamics of benthic and fish communities, including the important fishery species, coral trout (Plectropomus spp.), over 8 years in both NTRs and fished areas in the Keppel Island group, Great Barrier Reef, Australia. Two major disturbances impacted the reefs during the monitoring period, a coral bleaching event in 2006 and a freshwater flood plume in 2011. Both disturbances generated significant declines in coral cover and habitat complexity, with subsequent declines in fish abundance and diversity, and pronounced shifts in fish assemblage structure. Coral trout density also declined in response to the loss of live coral, however, the approximately 2:1 density ratio between NTRs and fished zones was maintained over time. The only post‐disturbance refuges for coral trout spawning stocks were within the NTRs that escaped the worst effects of the disturbances. Although NTRs had little discernible effect on the temporal dynamics of benthic or fish communities, it was evident that the post‐disturbance refuges for coral trout spawning stocks within some NTRs may be critically important to regional‐scale population persistence and recovery.     

The decay of similarity with geographical distance in parasite communities of vertebrate hosts

Aim The rate at which similarity in species composition decays with increasing distance was investigated among communities of parasitic helminths in different populations of the same host species. Rates of distance decay in similarity of parasite communities were compared between populations of fish and mammal hosts, which differ with respect to their vagility and potential to disperse parasite species over large distances. Location Data on helminth communities were compiled for several populations of three mammalian host species (Ondatra zibethicus, Procyon lotor and Canis latrans) and three fish host species (Perca flavescens, Catostomus commersoni and Esox lucius) from continental North America. Methods Distances between localities and similarity in the composition of helminth communities, the latter computed using the Jaccard index, were calculated for all possible pairs of host populations within each host species. Similarity values were then regressed against distance to see if they decayed at exponential rates, as reported for plant communities; the significance of the regressions was assessed using randomization tests. Results The number of hosts examined per population did not correlate with the number of helminth species found per population, and thus sampling effort is unlikely to have confounded the results. In four (two mammals and two fish) of the six host species, similarity in helminth communities decayed exponentially with distance. When the log of similarity is plotted against untransformed distance, the slopes obtained for the two fish species are lower than those obtained for the two mammalian host species. Main conclusions Similarity in the composition of parasite communities appears to decay exponentially with increasing distance in some host species, but not in all host species. The rate of decay is not necessarily associated with the vagility of the host. Although distance decay of similarity is generally occurring, it seems that other ecological processes, related either to the host or its habitat, can obscure it.     

Phylogenetic and environmental DNA insights into emerging aquatic parasites: implications for risk management

Species translocation leads to disease emergence in native species of considerable economic importance. Generalist parasites are more likely to be transported, become established and infect new hosts, thus their risk needs to be evaluated. Freshwater systems are particularly at risk from parasite introductions due to the frequency of fish movements, lack of international legislative controls for non-listed pathogens and inherent difficulties with monitoring disease introductions in wild fish populations. Here we used one of the world’s most invasive freshwater fish, the topmouth gudgeon, Pseudorasbora parva, to demonstrate the risk posed by an emergent generalist parasite, Sphaerothecum destruens. Pseudorasbora parva has spread to 32 countries from its native range in China through the aquaculture trade and has introduced S. destruens to at least five of these. We systematically investigated the spread of S. destruens through Great Britain and its establishment in native fish communities through a combination of phylogenetic studies of the host and parasite and a novel environmental DNA detection assay. Molecular approaches confirmed that S. destruens is present in 50% of the P. parva communities tested and was also detected in resident native fish communities but in the absence of notable histopathological changes. We identified specific P. parva haplotypes associated with S. destruens and evaluated the risk of disease emergence from this cryptic fish parasite. We provide a framework that can be applied to any aquatic pathogen to enhance detection and help mitigate future disease risks in wild fish populations.     

Founding population size of an aquatic invasive species

Non-native species of fish threaten native fishes throughout North America, and in the Rocky Mountains, introduced populations of lake trout threaten native populations of bull trout. Effective management of lake trout and other exotic species require understanding the dynamics of invasion in order to either suppress non-native populations or to prevent their spread. In this study, we used microsatellite genetic data to estimate the number of lake trout that invaded a population of bull trout in Swan Lake, MT. Examination of genetic diversity and allele frequencies within the Swan Lake populations showed that most of the genes in the lake trout population are descended from two founders. This emphasizes the importance of preventing even a few lake trout from colonizing new territory.     

Food borne parasites as indicators of trophic segregation between Arctic charr and brown trout

The habitat and diet choice and the infection (prevalence and abundance) of trophically transmitted parasites were compared in Arctic charr and brown trout living sympatrically in two lakes in northern Norway. Arctic charr were found in all main lake habitats, whereas the brown trout were almost exclusively found in the littoral zone. In both lakes the parasite fauna reflected the niche segregation between trout and charr. Surface insects were most common in the diet of trout, but transmit few parasites, and accordingly the brown trout had a relatively low diversity and abundance of parasites. Parasites transmitted by benthic prey such as Gammarus and insect larva, were common in both salmonid host species. Copepod transmitted parasites were much more common in Arctic charr, as brown trout did not include zooplankton in their diets. Parasite species that may use small fish as transport hosts, were far more abundant in piscivorous fish, especially brown trout. The seasonal dynamics in parasite infection were also consistent with the developments in the diet throughout the year. The study demonstrates that the structure of parasite communities of charr and the trout is highly dependent on shifts in habitat and diet of their hosts both on an annual base and through the ontogeny, in addition to the observed niche segregation between the two salmonid species.     

The taxonomic distinctness of coastal bottom-dwelling fish communities of the North-east Atlantic

1. New techniques for identifying the average taxonomic range of species assemblages were applied to an extensive dataset of bottom-dwelling fish in the coastal waters of NW Europe. These taxonomic distinctness indices provided much greater resolution than traditional diversity indices as they incorporated information on taxonomic relationships into an index which measures species dominance. Unlike standard measures of species richness and diversity, the mean value of these statistics is independent of sampling effort, and this allows objective comparisons to be made between samples from studies where sampling effort is not standardized.
2. A reduction in the average taxonomic range between the fauna of western waters of the UK and that of the southern North Sea was consistent with the general decline in species richness observed between these regions, and suggests that these two factors may be spatially positively correlated. Indices calculated for individual samples of fish on a local scale, however, did not all fit this trend.
3. Much of the variability in taxonomic diversity within the coastal waters of NW Europe was caused by the variable geographical distribution of the elasmobranchs. Of all the families which comprise the fish communities, this group has life-history characteristics which make it most susceptible to impact by commercial trawl fisheries.
4. The use of taxonomic distinctness measures provided additional insights, of relevance to biodiversity assessment, suggesting that they might usefully be applied to other aquatic and terrestrial fauna.     

Has the introduction of brown trout altered disease patterns in native New Zealand fish?

1. It is well recognised that non-indigenous species (NIS) can affect native communities via the 'spillover' of introduced parasites. However, two other potentially important processes, the 'spillback' of native parasites from a competent NIS host, where the latter acts as a reservoir leading to amplified infection in native hosts, and the 'dilution' of parasitism by a NIS host acting as a sink for native parasites, have either not been tested or largely overlooked.
2. We surveyed the helminth parasite fauna of native New Zealand fish in Otago streams that varied in the abundance of introduced brown trout Salmo trutta , to look for evidence of spillback and/or dilution. Spillover is not an issue in this system, with trout introduced as parasite-free eggs.
3. Seven native parasite species were present across 12 sites; significant inverse relationships with an index of trout abundance (i.e. dilution) were documented for three species infecting the native upland bully Gobiomorphus breviceps , and one species infecting the native roundhead galaxias Galaxias anomalus .
4. An inverse relationship between bully energy status and infection intensity of one parasite species suggests that parasite dilution could have positive effects on bully populations. Our failure to detect similar relationships for the other parasites does not preclude the possibility that dilution is beneficial to native fish, since parasites may have subtle or unmeasured impacts.
5. The parasite dilution patterns reported are compelling in that they occurred across several native host and parasite species; as such they have important implications for invasion ecology, providing an interesting contrast to the largely negative impacts reported for NIS. Mechanisms potentially responsible for the patterns observed are discussed.     

Geographical variation in host specificity of fleas (Siphonaptera) parasitic on small mammals: the influence of phylogeny and local environmental conditions

The evolution of host specificity remains a central issue in the study of host‐parasite relationships. Here we tackle three basic questions about host specificity using data on host use by fleas (Siphonaptera) from 21 geographical regions. First, are the host species exploited by a flea species no more than a random draw from the locally available host species, or do they form a taxonomically distinct subset? Using randomization tests, we showed that in the majority of cases, the taxonomic distinctness (measured as the average taxonomic distances among host species) of the hosts exploited by a flea is no different from that of random subsets of hosts taken from the regional pool. In the several cases where a difference was found, the taxonomic distinctness of the hosts used by a flea was almost always lower than that of the random subsets, suggesting that the parasites use hosts within a narrower taxonomic spectrum than what is available to them. Second, given the variation in host specificity among populations of the same flea species, is host specificity truly a species character? We found that host specificity measures are repeatable among different populations of the same flea species: host specificity varies significantly more among flea species than within flea species. This was true for both measures of host specificity used in the analyses: the number of host species exploited, and the index measuring the average taxonomic distinctness of the host species and its variance. Third, what causes geographical variation in host specificity among populations of the same flea species? In the vast majority of flea species, neither of our two measures of host specificity correlated with either the regional number of potential host species or their taxonomic distinctness, or the distance between the sampled region and the center of the flea's geographical range. However, in most flea species host specificity correlated with measures of the deviation in climatic conditions (precipitation and temperature) between the sampled region and the average conditions computed across the flea's entire range. Overall, these results suggest that host specificity in fleas is to a large extent phylogenetically constrained, while still strongly influenced by local environmental conditions.     

Invasive species are less parasitized than native competitors, but for how long? The case of the round goby in the Great Lakes-St. Lawrence Basin

There is increasing evidence that parasitism represents an unpredictable dimension of the ecological impacts of biological invasions. In addition to the risk of exotic pathogen transmission, other mechanisms such as parasite-release, could contribute to shaping the relationship between introduced species and native communities. In this study, we used the Eurasian round goby (Neogobius menalostomus) in the Great Lakes-St. Lawrence River ecosystem to further explore these ideas. As predicted by the parasite-release hypothesis, recently established populations of round goby were parasitized by a depauperate community of generalist helminths (8 taxa), all commonly found in the St. Lawrence River. In comparison, two native species, the logperch (Percina caprodes) and spottail shiner (Notropis hudsonius), were the hosts of 25 and 24 taxa respectively. Round gobies from each of 3 sampled localities were also less heavily infected than both indigenous species. This is in contrast to what is observed in round goby’s native range where the species is often the most parasitized among gobid competitors. This relative difference in parasite pressure could enhance its competitiveness in the introduced range. However, our study of an older population of round goby in Lake St. Clair suggests that this advantage over native species could be of short duration. Within 15 years, the parasite abundance and richness in the round goby has more than doubled whereas the number of parasite species per fish has increased to levels of those typical of fish indigenous to the St. Lawrence-Great Lakes watershed.     

Utilization of species checklist data in revealing the spatial distribution of fish diversity

Marine fish species checklists from six Chinese coastal waters were combined for the analysis of taxonomic diversity. The Genus-Family index (G-F index) ranged between 0.39 and 0.84, which generally indicated a decreasing trend with increasing latitude, with the exception of the southernmost area. Average taxonomic distinctness showed a slight increasing trend from northern to central study areas, but whether the taxonomic distinctness indices represent a latitudinal gradient of biodiversity requires further study. The multivariate analysis revealed a distinct latitudinal variation in fish assemblages. These results indicate that species checklist data are helpful in understanding the diversity distribution of fish species in the coastal zone. The potential of a species inventory should be exploited to fully understand biodiversity.     

Crying wolf, crying foul, or crying shame: alien salmonids and a biodiversity crisis in the southern cool-temperate galaxioid fishes?

The galaxioid fishes are the dominant, most speciose group of freshwater fishes (with >50 species) in the lands of the cool southern hemisphere, with representatives in western and eastern Australia, Tasmania, New Caledonia, Lord Howe Island, New Zealand, the Chatham, Auckland and Campbell Islands, Patagonian South America (Chile, Argentina), the Falkland Islands and South Africa. The group is most diverse in Australia and New Zealand. Lepidogalaxiidae is found only in Australia, Retropinnidae in Australia and New Zealand, and Galaxiidae across the entire range of the group. Many species are in serious conservation crisis for a diversity of reasons, including habitat deterioration and possibly fisheries exploitation, but there is enduring and pervasive information that shows that the group has been seriously impacted by the acclimatisation of salmonid fishes originating in the cool-temperate northern hemisphere, particularly brown and rainbow trout. With few exceptions, where these trout have been introduced there has been major decline in the galaxioids, especially Galaxiidae, as a result of a complexly interacting series of adverse impacts from these introduced fishes. In some places, centrarchids and cichlids may also have adverse impacts. In addition, there appear to have been adverse impacts from the translocation of galaxioids into communities where they do not naturally occur. In many instances it appears that displacement of the galaxioids has led to a situation where galaxioids and salmonids no longer co-occur, owing either to displacement or predation, leading to fish communities in which there is no explicit evidence for displacement. These effects are resulting in the galaxioid fishes being amongst the most seriously threatened fishes known.     

Orb-weaving spider diversity in the Iberá Marshlands, Argentina

The Iberá Marshlands RAMSAR reserve, in the northeast of Argentina, is one of the largest and most important wetlands of America. In this study we assess orb-weaving spider (Araneae: Orbiculariae) diversity in this reserve, analyzing different facets of local diversity (species richness, diversity, evenness and taxonomic distinctness), and the contribution of species differentiation (beta diversity) among localities and habitat types to the composition of regional diversity. We found 1657 individuals of 59 orb-weaving spider species/morphospecies. Local diversity differs among the three sampled localities. At the habitat level, the different facets of biodiversity followed a clear pattern, where woodlands have higher species richness, diversity, evenness and taxonomic distinctness than savannas. Savanna sites shared a common spider species composition, while woodland communities have high values of complementarity. Thus, beta diversity has a very high contribution to the regional diversity of the orb-weaving spiders in the Iberá Marshlands. We suggest that conservation management in the reserve should be directed towards promoting natural spatial heterogeneity, giving special protection to habitat mosaics in different localities.