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1.
On Australia's south‐east coast, hybridization between estuary‐restricted black bream Acanthopagrus butcheri Munro and its migratory coastal congener yellowfin bream Acanthopagrus australis (Günther) has led to estuarine populations largely composed of hybrids that are most genetically similar to A. butcheri. The fertilization success achieved when ova of estuary‐caught A. butcheri were fertilized with the cryogenically preserved sperm of either ocean‐caught A. australis or estuary‐caught A. butcheri‐like was compared. The experimental crosses, which by chance included both pure parental and hybrid bream, revealed no evidence that gametic incompatibility provides a barrier to fertilization among both pure species and their hybrids.  相似文献   

2.
Populations of obligately estuarine taxa are potentially small and isolated and may lack genetic variation and display regional differentiation as a result of drift and inbreeding. Hybridization with a wide‐ranging marine congener should introduce genetic variation and reduce the effects of inbreeding depression and genetic drift. However, high levels of hybridization can cause demographic and genetic swamping. In southeastern Australia hybridization occurs between obligately estuarine Black bream (Acanthopagrus butcheri) and migratory marine Yellowfin bream (Acanthopagrus australis). Here, we surveyed genetic variation at eight microsatellite loci and the mitochondrial control region of juvenile fish from five coastal lagoons (including temporal replication in two lagoons) (total n = 970) to determine the frequency and persistence of hybridization, and its likely consequence for the estuarine restricted A. butcheri. Of 688 juvenile fish genotyped 95% were either A. australis (347) or hybrids (309); only 5% (32) were A. butcheri. Most hybrids were later generation hybrids or A. butcheri backcrosses, which are likely multi‐generational residents within lagoons. Far greater proportions of hybrid juveniles were found within two lagoons that are generally closed to the ocean (>90% hybrid fish within generally closed lagoons vs. 12–27% in permanently or intermittently open lagoons). In both lagoons, this was consistent across multiple cohorts of fish [79–97% hybrid fish (n = 282)]. Hybridization and introgression represent a major threat to the persistence of A. butcheri and have yet to be investigated for large numbers of estuarine taxa.  相似文献   

3.
We have compared the species composition and diversity of the diets of black bream, Acanthopagrus butcheri, in three normally closed estuaries on the central south coast of Western Australia, which vary markedly in the extents to which they become hypersaline during dry periods. Although black bream was caught seasonally in Stokes Inlet during this 3-year study, it was obtained from the Hamersley and Culham inlets only during the earlier seasons because salinities in those two estuaries subsequently rose to levels that caused massive mortalities of this sparid. Although a wide range of taxa, including macrophytes, polychaetes, molluscs, crustaceans, insects and teleosts, were ingested by A. butcheri in each estuary, the frequencies of ingestion and volumetric dietary contributions of these taxa varied greatly among the fish in these three estuaries. Thus, for example, in comparison with other estuaries, relatively greater contributions were made to the diet by polychaetes and crustaceans in Stokes Inlet, by macrophytes in Hamersley Inlet, and by insects (mainly chironomid larvae) in Culham Inlet. The relatively greater contribution of teleosts to the diets of black bream in the Hamersley and Culham inlets than in Stokes Inlet, and also differences in the main teleost species ingested in the first two estuaries, are consistent with differences in the densities of fish overall and of the main fish species in those estuaries. The diversity of the diet was far greater in Stokes Inlet than in the other two far more variably saline estuaries, presumably reflecting a greater diversity of food. The dietary compositions of black bream in upstream pools in the tributary of Culham Inlet, which offer refuge when salinities increase markedly in the main body of the estuary, differ from those in those downstream regions, further emphasising the opportunistic nature of the feeding behaviour of black bream. The dietary compositions of black bream underwent size-related changes, but the taxa contributing most to those changes varied greatly among estuaries. Size-related changes would be particularly beneficial in reducing intraspecific competition for food in the two estuaries that vary greatly in salinity and would thus be likely to contain a less diverse range of prey.  相似文献   

4.
Acanthopagrus butcheri completes its entire life history within estuaries and coastal lakes of southern Australia, although adults occasionally move between estuaries via the sea. Consequently, it is expected that populations of A. butcheri in different estuaries will be genetically distinct, with the magnitude of genetic divergence increasing with geographic isolation. However, previous genetic studies of A. butcheri from southeast Australia yielded conflicting results; allozyme variation exhibited minimal spatial structuring (θ = 0.012), whereas mitochondrial DNA distinguished the majority of populations analyzed (θ = 0.263) and genetic divergence was positively correlated with geographic isolation. This discrepancy could reflect high male gene flow, which impacts nuclear but not mitochondrial markers. Here we estimated allele frequencies at five nuclear microsatellite loci across 11 southeast Australian populations (595 individuals). Overall structuring of microsatellite variation was weaker (θ = 0.088) than that observed for mitochondrial DNA, but was able to distinguish a greater number of populations and was positively correlated with geographic distance. Therefore, we reject high male gene flow and invoke a stepping-stone model of infrequent gene flow among estuaries for both sexes. Likewise, management of A. butcheri within the study range should be conducted at the scale of individual or geographically proximate estuaries for both sexes. The lack of allozyme structuring in southeast Australia reflects either the large variance in structuring expected among loci under neutral conditions and the low number of allozymes surveyed or a recent colonization of estuaries such that some but not all nuclear loci have approached migration-drift equilibrium.  相似文献   

5.
Understanding challenges posed by climate change to estuaries and their faunas remains a high priority for managing these systems and their communities. Freshwater discharge into a range of estuary types in south‐western Australia between 1990 and 2015 is shown to be related to rainfall. This largely accounts for decreases in discharge in this microtidal region being more pronounced on the west coast than south coast, where rainfall decline was less. Results of an oxygen‐balance model imply that, as demonstrated by empirical data for the Swan River Estuary, declines in discharge into a range of estuary types would be accompanied by increases in the extent of hypoxia. In 2013–15, growth and body condition of the teleost Acanthopagrus butcheri varied markedly among three permanently open, one intermittently‐open, one seasonally‐closed and one normally‐closed estuary, with average time taken by females to reach the minimum legal length (MLL) of 250 mm ranging from 3.6 to 17.7 years. It is proposed that, in a given restricted period, these inter‐estuary variations in biological characteristics are related more to differences in factors, such as food resources and density, than to temperature and salinity. The biological characteristics of A. butcheri in the four estuaries, for which there are historical data, changed markedly between 1993–96 and 2013–15. Growth of both sexes, and also body condition in all but the normally‐closed estuary, declined, with females taking between 1.7 and 2.9 times longer to attain the MLL. Irrespective of period, body condition, and growth are positively related. Age at maturity typically increased between periods, but length at maturity declined only in the estuary in which growth was greatest. The plasticity of the biological characteristics of A. butcheri, allied with confinement to its natal estuary and ability to tolerate a wide range of environmental conditions, makes this sparid and comparable species excellent subjects for assessing estuarine “health.”  相似文献   

6.
The dietary compositions were determined for Acanthopagrus butcheri in four estuaries and a saline, coastal lake, which vary in the extent, if any, of their connection to the sea and amongst which, in spring and summer, their salinities ranged from 2 to 7‰ in an intermittently open estuary to >40‰ in a normally closed estuary. The dietary compositions of A. butcheri in each of the five water bodies were significantly different, which reflected differences in the abundance of different components of the biota in those systems. Biotic differences amongst water bodies thus accounted for the far larger contributions made to the volume of the stomach contents by the macroalgae Cladophora sp. in the Moore River Estuary on the lower west coast of Australia, by amphipods and decapods in the Swan River Estuary 85 km further south and by polychaetes in the landlocked Lake Clifton a further 85 km further south. The diet of A. butcheri in the Nornalup/Walpole Estuary on the south coast of Western Australia contained atypically large volumes of the seagrass Ruppia megacarpa and teleosts, whereas that in the normally closed Wellstead Estuary, 260 km to the east was characterised by large volumes of the macroalgae Chaetomorpha sp. and a tube‐dwelling amphipod. However, there is evidence that A. butcheri selects certain prey, when two or more of the typical prey of A. butcheri are present in the environment, and that it prefers to feed on or above the substratum, rather than within the substratum. The dietary composition of A. butcheri underwent pronounced ontogenetic changes in each water body, these being progressive in estuaries on the lower west coast and abrupt in those on the south coast. Within the upper Swan Estuary, the dietary composition changed in an upstream direction, reflecting changes in the relative abundance of certain benthic macroinvertebrate prey, but did not undergo conspicuous seasonal changes, which is consistent with the lack of any clear cut seasonal changes in the abundance of their major prey.  相似文献   

7.
Hypoxia is a widespread environmental stressor that affects marine, estuarine and freshwater systems worldwide. Investigating the effects of hypoxia on aquatic animals in the natural environment is difficult and expensive. Laboratory experiments provide an alternative that allows manipulation of environmental variables and simulation of altered conditions that are expected in the future. However, controlling dissolved oxygen (DO) levels precisely in laboratory test situations is challenging and generally costly and time intensive. In this paper, we describe a novel chamber design that is capable of maintaining low DO levels precisely for a period of at least 4 days with minimal re-adjustment and nitrogen gas requirement. The system is simple, inexpensive and easy to use, and offers the additional benefit of being portable. The utility of this design is demonstrated with an experiment to test the effects of hypoxia and high salinity on hatch rates and yolk-sac larvae survival in the estuarine fish black bream Acanthopagrus butcheri (Munro, 1949). The results show that early life-stage black bream are sensitive to hypoxia and increased salinity, indicating that deterioration in estuarine environmental conditions may have negative effects on spawning success and recruitment to fish populations. This new laboratory technique will be useful for testing future scenarios for the estuaries of southern Australia as predicted by climate change models. Other potential applications and modifications for further development of this new technique are discussed.  相似文献   

8.
Movement patterns and habitat utilization by black bream Acanthopagrus butcheri (Sparidae), an estuarine resident species, were investigated using acoustic telemetry in a small estuary on the east coast of Tasmania, Australia. Thirty‐four adult A. butcheri were tracked for periods of up to 187 days between August 2005 and January 2006. Although able to tolerate a wide range of salinities, the fish spent most of the time within the upper and middle regions of the estuary, where brackish conditions dominated. The species exhibited extensive movements linked to tidal cycles, with small‐scale upstream movements during incoming tides and downstream movements during out going tides. The extent of these movements was positively correlated with the tidal height difference between consecutive tidal peaks and troughs. Freshwater inflows and resultant changes in salinity also significantly influenced distribution and movement patterns. Fish moved downstream during the periods of heavy inflows, returning upstream as salinities increased to c. >10. During the peak of spawning period (November to December) fish moved into the upper region of the estuary, where they aggregated to spawn. Periodic increases in freshwater discharge, however, resulted in fish leaving the spawning grounds and moving downstream. Towards the end of the spawning season (January), the fish became more dispersed throughout the entire estuarine system.  相似文献   

9.
Estuaries are particularly susceptible to climate change and drought resulting in atypical changes to freshwater flows. How such changes in flow impact on the ecology of estuarine fishes may depend on how a species moves in response to changing flow conditions. Acoustic telemetry was used to interpret fine-scale movements of two co-inhabiting estuarine fish species, black bream, Acanthopagrus butcheri and estuary perch, Macquaria colonorum in relation to freshwater flows, season and moon phase. We found black bream to be highly mobile, regularly travelling the length of the estuary and into the neighbouring estuaries. In contrast, estuary perch had particular home ranges and made occasional, upstream or downstream movements. Possibly influenced by freshwater flows, estuary perch moved at greater rates in the Tambo compared to fish in the Mitchell. Black bream resided in the upper estuary during winter and spring and the lower estuary during summer and autumn, whereas estuary perch remained in the upper estuary throughout the year, with occasional downstream movements in winter and spring. This study revealed 1) significantly large increases in freshwater flows result in mass downstream movements in both species, 2) fish moved upstream during full moons and 3) there are contrasting spatio-temporal patterns in movement between species. The results from this study highlight that estuarine fishes are likely to show differential sensitivity to the impacts of drought and climate change and illustrate how acoustic telemetry methods can be used to determine the environmental needs of fishes and help efforts to conserve and manage estuaries worldwide.  相似文献   

10.
Climate change can affect marine and estuarine fish via alterations to their distributions, abundances, sizes, physiology and ecological interactions, threatening the provision of ecosystem goods and services. While we have an emerging understanding of such ecological impacts to fish, we know little about the potential influence of climate change on the provision of nutritional seafood to sustain human populations. In particular, the quantity, quality and/or taste of seafood may be altered by future environmental changes with implications for the economic viability of fisheries. In an orthogonal mesocosm experiment, we tested the influence of near‐future ocean warming and acidification on the growth, health and seafood quality of a recreationally and commercially important fish, yellowfin bream (Acanthopagrus australis). The growth of yellowfin bream significantly increased under near‐future temperature conditions (but not acidification), with little change in health (blood glucose and haematocrit) or tissue biochemistry and nutritional properties (fatty acids, lipids, macro‐ and micronutrients, moisture, ash and total N). Yellowfin bream appear to be highly resilient to predicted near‐future ocean climate change, which might be facilitated by their wide spatio‐temporal distribution across habitats and broad diet. Moreover, an increase in growth, but little change in tissue quality, suggests that near‐future ocean conditions will benefit fisheries and fishers that target yellowfin bream. The data reiterate the inherent resilience of yellowfin bream as an evolutionary consequence of their euryhaline status in often environmentally challenging habitats and imply their sustainable and viable fisheries into the future. We contend that widely distributed species that span large geographic areas and habitats can be “climate winners” by being resilient to the negative direct impacts of near‐future oceanic and estuarine climate change.  相似文献   

11.
Niche variation between hybrid taxa and their parental species has been deemed imperative to the persistence of hybrid populations in nature. However, the ecological factors promoting hybrid establishment remain poorly understood. Through the application of a multidisciplinary approach integrating genetics, morphometry, life‐history, and trophic ecology, we studied the hybrids of roach (Rutilus rutilus L.) and bream (Abramis brama L.), and their parental species inhabiting an Irish lake. The roach × bream hybrid exhibited a body shape intermediate of that of the parental species. Diet analyses depicted the hybrid as a generalist, feeding on all prey items consumed by either parental species. Stable isotope data confirm the trophic niche breadth of hybrids. A significant correlation between body shape and diet was detected, suggesting that the intermediate phenotype of hybrids might play a role in their feeding abilities, resulting in the utilization of a broader trophic spectrum than the parental species. Growth and age class structure analyses also yielded a scenario that is consistent with the ecological success of hybrids. Genetic analyses suggest that the majority of hybrids result from first‐generation crosses between the parental species; however, a potentially significant proportion of back‐crosses with bream were also detected. The recent introduction of roach and bream into Irish waters, as well as the climatic and ecological features of the colonized habitats, can explain the remarkable success of the roach × bream hybrid in Ireland. The adaptive significance of hybridization and its demographic consequences for the parental species are discussed. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 768–783.  相似文献   

12.
The bream (Abramis brama L.) population of Tjeukemeer was studied for three successive years. The growth, condition, gonad development, diet and feeding conditions of the fish are described. Food competition for two years with the then abundant young planktivorous smelt (Osmerus eperlanus) and perch (Perca fluviatilis) resulted in a decreased growth rate, condition and gonad development of the larger bream. The biomass of chironomids was too small in relation to that of the zooplankton to be important for the bream population, although they were eaten more efficiently.  相似文献   

13.
There is an increasing desire for researchers to use the elemental concentrations in fish otoliths to reconstruct environmental histories of fish. These reconstructions may be plausible due to the unique incorporation of elements into discrete layers of otolith material that correspond to daily growth, and because environmental variables of temperature, salinity, and water chemistry can influence otolith chemistry. However, it is essential to establish exactly how temperature, salinity, and the ambient concentration of elements influence otolith chemistry in order to interpret environmental histories of fish. Using a controlled laboratory experiment we tested the relative and interactive effects of temperature, salinity, and ambient concentration of strontium (Sr) and barium (Ba) on the resulting concentration of Sr and Ba in otoliths of black bream Acanthopagrus butcheri (Munro 1949). Salinity and concentration, and temperature and concentration interacted to affect the elemental concentration of Sr:Ca and Ba:Ca in otoliths. Regression analysis revealed that temperature and ambient concentration contributed most to the trend in otolith chemistry for both elements. Importantly, this is the first experiment to combine three environmental variables and assess their effect on otolith chemistry. Based on these results, it should be possible to use changes in the elemental concentration in otoliths to better reconstruct previous environments of temperature, salinity, and ambient water chemistry, which is especially useful when determining occupancy in habitats such as estuaries that display variable environmental characteristics.  相似文献   

14.
Eight pairs of simple sequence repeat markers were developed from random amplified polymorphic DNA product in black sea bream Acanthopagrus schlegeli. Twenty microsatellites were selected for designing microsatellite primers, of which eight gave working primer pairs. They had between three and seven alleles. Observed and expected heterozygosities varied from 0.65 to 0.90, and from 0.58 to 0.82, respectively. Eight additional fish species assessed for cross‐species amplification revealed between two and five positive amplifications and between zero and three polymorphic loci per species.  相似文献   

15.
Aim The seaweed Fucus ceranoides is restricted to spatially discrete estuarine habitats and lacks planktonic dispersal phases; it is therefore expected to exhibit strong population differentiation. Its cold‐temperate affinities and mtDNA variation imply that the northern part of the species’ range, where F. ceranoides is now ubiquitous, was recently colonized after the onset of the last deglaciation, potentially resulting in areas with greater genetic homogeneity. Here we examine the population structure of F. ceranoides to test these predictions, emphasizing the contrasting genetic signatures of limited dispersal in refugial versus recently colonized regions. Location North‐eastern Atlantic estuaries from Portugal to Norway. Methods A total of 504 individuals from 21 estuarine sites spanning the entire range of F. ceranoides were sampled and genotyped for nine microsatellite loci. Population structure was inferred from several genotypic and allele‐frequency analyses. Geographical patterns of genetic diversity were used to reconstruct the historical biogeography of the species. Results Genetic diversity and regional population differentiation showed a consistent decline with increasing latitude. Southernmost populations harboured most of the endemic variation, whereas the northern populations (> 55° N) were almost fixed for the same alleles across loci. In southern and central regions of its distribution, F. ceranoides showed striking population subdivision, with many of the sampled estuaries corresponding to coherent genetic units that were easily discriminated from one another with standard clustering methods. Main conclusions The geographical pattern of genetic diversity supports the long‐term refugial status of Iberia and a post‐glacial range expansion of F. ceranoides into previously glaciated latitudes. Despite the species’ capacity to colonize newly available habitats, the genetic structure of F. ceranoides outside the recently (re)colonized range reveals that gene flow between populations is extremely low. This study provides a remarkable example of how infrequent and spatially limited dispersal can have contrasting effects at the scales of meta‐population (connectivity) versus range dynamics (habitat tracking), and of how dispersal restrictions can result in either genetic divergence or homogeneity depending on the maturity and demographic conditions of the populations.  相似文献   

16.
A laboratory experiment was conducted to determine the interactive effects of temperature and diet on condition indices of juvenile black bream Acanthopagrus butcheri, reared for time periods ranging from 2 to 42 days. After fish were reared for varying periods, growth, morphometric (Fulton's K) and biochemical [RNA:DNA (R:D) ratios] indices were measured. Fulton's K responded primarily to temperature, with progressive decrease in condition over time for fish reared at high temperatures. In contrast, R:D ratios were primarily affected by diet composition, with the highest values observed for fish reared on fish‐based diets as opposed to vegetable‐based diets. Significant effects of rearing time were also observed for Fulton's K and R:D ratios, as were some interactive treatment effects. In addition, Fulton's K and R:D ratios were not significantly correlated, perhaps due to the different periods of time integrated by each index or their relative sensitivity to lipid and protein deposition. These results highlight the complex responses of these condition indices to environmental variables and nutritional status.  相似文献   

17.
A new sparid species, Acanthopagrus akazakii, is described on the basis of 12 types collected from Noumèa, New Caledonia. Acanthopagrus akazakii is most similar to Acanthopagrus berda in overall appearance but differs from the latter in having 41/2 scale rows between the 5th dorsal fin spine base and lateral line, anteriormost head scales broadly rounded without small scales anteriorly, upward- and downward-oriented portions present on anteriormost part of upper lip, a downward-oriented portion on anteriormost part of lower lip in specimens over ca. 160 mm in standard length, and upper head profile gently convex from snout tip to above eye throughout growth (vs. 31/2 scale rows between 5th dorsal fin spine base and lateral line, anteriormost head scales rounded with small scales anteriorly, no upward and downward portions in anteriormost upper and lower lips, and upper head profile becoming concave from snout tip to above eye with growth). Furthermore, A. berda develops a strong concavity of the ventral edge of the first two infraorbitals above the posterior part of upper jaw with growth, whereas A. akazakii has a generally straight series throughout growth. Nominal species in A. berda are reviewed, with notes on nominal species of Acanthopagrus.  相似文献   

18.
This study examined inter‐period changes over two to three decades in the fish fauna of an urbanized estuary experiencing rapid population growth and a drying climate (Swan–Canning Estuary, Western Australia). Responses were compared at the fish community level (species composition; 1978–2009 in the shallows and 1993–2009 in deeper waters) and at the population and individual levels of an estuarine indicator species, black bream Acanthopagrus butcheri (biomass–abundance and per capita mass at age, respectively; 1993–2009). All three levels showed distinct shifts from earlier to later periods, but their patterns, sensitivity and breadth differed. Community composition changed markedly in the shallows of the lower‐middle estuary between the late 1970s and all later periods and moderately between more disparate periods from 1995 to 2009. Several species trends could be linked to the increasing salinity of the estuary or declining dissolved oxygen levels in its middle–upper reaches. Community changes were, however, small or insignificant in the shallow and deeper waters of the upper estuary and deeper waters of the middle estuary, where environmental perturbations are often most pronounced. This may reflect the resilience of the limited suite of species that typify those reaches and thus their lack of sensitivity in reflecting longer‐term change at the coarser level of mean abundance. One such species, the selected indicator, A. butcheri, did, however, show marked temporal changes at both the population and individual levels. Biomass decreased markedly in deeper waters while increasing in the shallows from earlier to later periods, presumably reflecting an onshore movement of fish, and per capita body mass in the 2+, 3+ and 4+ year classes fell steadily over time. Such changes probably indicate deteriorating habitat quality in the deeper waters. The study outcomes provide support for a multifaceted approach to the biomonitoring of estuaries using fishes and highlight the need for complementary monitoring of relevant stressors to better disentangle cause–effect pathways.  相似文献   

19.
In a two-year-study, the fish community of the Loosdrecht Lakes area was characterized with regard to the professional fishery and the forage base. The lakes are shallow (mean depth 2 m) and eutrophic to hypertrophic. The bream,Abramis brama, dominates the fish biomass. The pikeperch,Stizostedion lucioperca, is the main predator.Bream up to 30 cm have a slow growth rate and are in a bad condition. A faster growth rate and a better condition are shown by bream of 30 cm and more. The small bream feeds on chironomid larvae, benthic cladocerans and zooplankton. The better condition of bream over 30 cm is explained by the more efficient feeding of larger bream onChironomus plumosus larvae. Pikeperch show a fast growth rate and a good condition. Recruitment is limited by the low densities of smelt,Osmerus eperlanus, leaving cannibalism as the most important way for the 0+ pikeperch to become piscivorous and to manifest a fast growth.The impact of the professional gillnet fishery on bream and pikeperch is small because the mesh sizes in use are as large as 75–100 mm bar mesh.The planktivorous 0+ pikeperch consumes mainly the carnivorous zooplanktersLeptodora kindtii and cyclopoids. The zooplankton community lacks large herbivorous species likeDaphnia hyalina, capable of consuming bluegreens. A possible experiment in biomanipulation with a view to find out whether the development ofD.hyalina is depressed by the small planktivorous cyprinids, is predator enhancement with the aid of a stocking programme for indoor-raised 0+ pikeperch in early summer.  相似文献   

20.
For free-spawning estuarine taxa, gene flow among estuaries may occur via hybridization with mobile congeners. This phenomenon has rarely been investigated, but is probably susceptible to anthropogenic disturbance. In eastern Australia, the estuarine Black Bream Acanthopagrus butcheri and marine Yellowfin Bream Acanthopagrus australis have overlapping distributions and the potential to hybridize. We used surveys of microsatellite and mtDNA variation in 565 adults from 25 estuaries spanning their distributional range to characterize the species and their putative hybrids. Hybrids were widespread (68% of estuaries) and hybrid frequencies varied greatly among estuaries (0-58%). Most (88%) were classed as advanced generation backcrosses with A. butcheri and displayed A. butcheri mtDNA haplotypes. We found most hybrids in the three estuaries within the zone of sympatry (57%). Our study highlights the underemphasized importance of estuaries as sites of hybridization and suggests that hybridization is driven both by opportunity for contact and human activity.  相似文献   

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