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1.
Recently, a highly invasive alien species, the zebra mussel Dreissena polymorpha, has colonized Italian aquatic ecosystems that had been previously colonized by another highly invasive alien species—the North American crayfish Procambarus clarkii. This was the first time and world region where these two species met. To evaluate the relative importance of their interactions, we studied prey selection according to its length, as well as prey choice by P. clarkii preying on D. polymorpha in the presence and absence of alternative prey—the freshwater snail Physella acuta. We followed an optimability-based approach, by first estimating the most profitable length of each mollusc and then by conducting a prey choice experiment, on which both species were provided simultaneously to a crayfish with different size combinations. Prey selection was dependent on prey length, handling time and crayfish length for both molluscs. According to our profitability estimates, snails should be more profitable than mussels in the length range 7–10 mm, while for lengths over 11.0 mm, mussels should be more profitable. The results of the prey choice experiment indicated that D. polymorpha length, P. acuta length, and the difference in profitability between the offered individuals were all relevant for the choice of one species over the other by P. clarkii. However, the overall tendency was the choice of the smallest prey, regardless of species, and the estimates of prey profitability were not useful to predict prey choice. Our study shows that D. polymorpha represents a novel prey resource for P. clarkii, even in the presence of an alternative prey, and that zebra mussels may be a preferred prey, especially small-sized individuals (5–10 mm).  相似文献   

2.
Impacts of invasive species may manifest most strongly if these organisms are highly distinct functionally from the native species they often replace. Yet, should we expect functional differences between native and invasive species of generalist organisms like freshwater crayfish? Some existing evidence has pointed to native and invasive crayfish species as ecologically equivalent. We contribute to this literature by comparing the trophic niches of the globally invasive crayfishes Pacifastacus leniusculus and Procambarus clarkii, by applying carbon and nitrogen stable isotope analyses to replicated allopatric (alone) and sympatric (together) lake populations in western Washington State, USA, where P. clarkii has been recently introduced and P. leniusculus is presumed native. Our study corrected for potential inherent differences in lake food webs as a consequence of lake abiotic or biotic characteristics using random effects in linear mixed effects models. We found that although overall trophic niche size or area of these species was not significantly different, P. leniusculus was significantly higher in trophic position than P. clarkii when also accounting for the effects of body size, sex, and lakes as random effects. This pattern of increased trophic position of P. leniusculus over P. clarkii was conserved over time in one sympatric lake for which we had data over multiple years. Cumulatively, our findings point to trophic differences between the globally cosmopolitan crayfishes P. leniusculus and P. clarkii, particularly when accounting for the ways that ecosystem context can affect food web structure of communities and the trophic resources available to these consumers.  相似文献   

3.
Red swamp crayfish, Procambarus clarkii, presently is an important aquatic commercial species in China. The crayfish is a hot area of research focus, and its genetic improvement is quite urgent for the crayfish aquaculture in China. However, the knowledge of its genomic landscape is limited. In this study, a survey of P. clarkii genome was investigated based on Illumina’s Solexa sequencing platform. Meanwhile, its genome size was estimated using flow cytometry. Interestingly, the genome size estimated is about 8.50 Gb by flow cytometry and 1.86 Gb with genome survey sequencing. Based on the assembled genome sequences, total of 136,962 genes and 152,268 exons were predicted, and the predicted genes ranged from 150 to 12,807 bp in length. The survey sequences could help accelerate the progress of gene discovery involved in genetic diversity and evolutionary analysis, even though it could not successfully applied for estimation of P. clarkii genome size.  相似文献   

4.
To improve the knowledge on the Antarctic fur seal foraging behavior, evaluate whether changes in its diet are associated with changes in prey availability, and evaluate whether fisheries had negative impacts on Antarctic fur seal populations, a total of 1359 scats of non-breeding males were collected in ten consecutive sampling periods between 1994 and 2003 at Laurie Island, South Orkney Islands. Antarctic krill was the most numerous prey throughout the sampling period followed in importance by fish prey. Antarctic krill also predominated by reconstructed mass, except during 1994 and 1998 when penguins were the most important prey, and during 1995 when fish dominated. Among fish, demersal-benthic species (mainly Gobionotothen gibberifrons, Chaenocephalus aceratus and Chionodraco rastrospinosus) dominated the diet except in 1997, 1998, 1999 and 2002 when myctophids (mainly Gymnoscopelus nicholsi and Electrona antarctica) were the most important fish prey. In seasons when fish dominated the diet, the diet was primarily comprised of demersal species. Although our results are in overall agreement with previous studies, the contribution to the diet of the main preys changed throughout the sampling period, probably in parallel with changes in prey abundance. According to historical results on fish consumption, the fisheries carried out in the past have had negative impacts on seal populations from the South Orkney Islands.  相似文献   

5.
Understanding predator-prey dynamics is an important component of management strategy development for wildlife populations that are directly affected by predation. Ungulates often serve as a significant source of prey for many large mammal predators, and patterns of predation are known to influence population dynamics. Although black bear and wolf diets have been investigated extensively, prey preference has been less commonly examined, especially in analyses that take into account age class (i.e., juvenile and adult) of the ungulate prey. We examined black bear (Ursus americanus), wolf (Canis lupus), and hybrids (Canis spp.) prey preference in Ontario based on the availability of three ungulate species—elk (Cervus elaphus), moose (Alces alces), and white-tailed deer (Odocoileus virginianus). We analyzed the presence of prey items in black bear and wolf scats collected over 3 years by examining prey hair cuticular scale patterns. We applied correction factors to frequencies of occurrence of prey items found in predator scat and related diet composition to the availability of ungulates, determined by fecal pellet transect surveys. In addition, non-ungulate diet items were identified to obtain full diet composition profiles. We found that black bear diet consisted of more than 87% vegetation, and they were opportunistic, not selecting for any particular ungulate species in either adult or juvenile age class. Wolf diet was comprised mainly of ungulates (~?73.2%), muskrat (Ondatra zibethica; ~?8.5%), and beaver (Castor canadensis; ~?14.6%), and although moose were at least 1.5 times more abundant then each of the other ungulate prey species in the study area, wolves preferred elk, using moose less than expected. Although we found black bear diet to be opportunistic during the summer, wolves in our study heavily utilized both juvenile and adult ungulates, however, among ungulate species, displayed preference for elk. The preference displayed by wolves provides insight that wildlife managers can use to guide further investigation and assist with the development of strategies to ensure continued elk reintroduction success, and moose and white-tailed deer population sustainability.  相似文献   

6.
Species distribution models combining environmental and spatial components are increasingly used to understand and forecast species invasions. However, modelling distributions of invasive species inhabiting stream networks requires due consideration of their dendritic spatial structure, which may strongly constrain dispersal and colonization pathways. Here we evaluate the application of novel geostatistical tools to species distribution modelling in dendritic networks, using as case study two invasive crayfish (Procambarus clarkii and Pacifastacus leniusculus) in a Mediterranean watershed. Specifically, we used logistic mixed models to relate the probability of occurrence of each crayfish to environmental variables, while specifying three spatial autocorrelation components in random errors. These components described spatial dependencies between sites as a function of (1) straight-line distances (Euclidean model) between sites, (2) hydrologic (along the waterlines) distances between flow-connected sites (tail-up model), and (3) hydrologic distances irrespective of flow connection (tail-down model). We found a positive effect of stream order on P. clarkii, indicating an association with the lower and mid reaches of larger streams, while P. leniusculus was affected by an interaction between stream order and elevation, indicating an association with larger streams at higher altitude. For both species, models including environmental and spatial components far outperformed the pure environmental models, with the tail-up and the Euclidean components being the most important for P. clarkii and P. leniusculus, respectively. Overall, our study highlighted the value of geostatistical tools to model the distribution of riverine and aquatic invasive species, and stress the need to specify spatial dependencies representing the dendritic network structure of stream ecosystems.  相似文献   

7.
Many amphibian population declines have been associated with the introduction of alien aquatic predators. Here, we explore the vulnerability of tadpoles of two sympatric Japanese species [Pelophylax nigromaculatus (PN) and Rhacophorus schlegelii (RS)] to the invasive crayfish Procambarus clarkii. We first examined the behavioral responses of the tadpoles to the cues of caged, fed crayfish predator in a controlled laboratory experiment, and subsequently tested their survival when together in the presence of free-ranging predator in outdoor mesocosms that simulated natural ponds. Only PN reduced activity level to the cues of the predator, but this apparent behavioral defense recorded in the laboratory did not result into higher survival in outdoor mesocosms. In mesocosms, PN exhibited higher biomass increment but experienced higher mortality in predator environments. The mechanism mediating mortality remains unclear though. These results indicate that sympatric prey may differentially respond and be disproportionally vulnerable to novel predators. Our study illustrates the possible contribution of a life-history trait influencing risk of predation in newly invaded systems.  相似文献   

8.
Environmental DNA (eDNA) is a powerful method for assessing the presence and distribution of invasive aquatic species. We used this tool to detect and monitor several invasive crayfishes Procambarus clarkii, Orconectes limosus and Pacifastacus leniusculus present in, or likely to invade, the ponds of the Brenne Regional Natural Park. A previous study showed that the eDNA method was not very efficient in detecting P. clarkii. In the present study, we explored new improvements in the detection of invasive crayfish. We designed specific primers for each crayfish species, and set up an experimental mesocosm approach to confirm the specificity of the primers and the sampling protocol. We analysed samples taken from ponds in 2014 and 2015. We compared two qPCR protocols involving either SybrGreen or TaqMan assays. Using these same primers, we were able to detect crayfish eDNA with both assays during the mesocosm experiment. However, crayfish from field samples could only be detected by performing qPCR with a SybrGreen assay. We successfully monitored the presence of three invasive species of crayfish using eDNA. This method is a powerful tool for establishing the presence or absence of invasive species in various freshwater environments.  相似文献   

9.
The invasive clam Corbicula fluminea causes severe environmental and economic impacts in invaded sites and fouled water-dependent industries. The biological control of invasive species has potential as an effective, safe and low-cost tool. The potential of using direct (clam consumption) and indirect approaches (predator-avoidance behaviour) to control the pest was evaluated, aiming at establishing a proof-of-principle regarding the suitability of such strategies in confined settings (e.g. pipelines, channels, raw water tanks and other water-dependent facilities). Invertebrate (Procambarus clarkii) and vertebrate (Lepomis gibbosus, Luciobarbus bocagei) predators were examined for their control potential. In feeding experiments, barbels foraged upon C. fluminea but ingested very few clams, and clam consumption by crayfish was only observed in the smallest size class (<1 cm). Additional experiments linked these results to the protective role of the clam’s shell. In practical terms, the potential of the fish species as C. fluminea predators is limited, but crayfish can be important predators of small clams. In a second set of experiments, putative predator avoidance by clams was assessed in dual-choice aquaria. Refuge availability, predator diet and light conditions were taken into account. Clam avoidance was observed under particular conditions, but the amplitude of dislocation was limited.  相似文献   

10.
Freshwater ecosystems are particularly vulnerable to biological invasions; the aquatic animal pet trade has been recognized as a significant pathway of introductions. Crayfish are considered a model group of traded organisms with a series of highly successful species already established in the wild, having the potential to negatively influence both indigenous crayfish species (ICS) as well as alter occupied ecosystems. Eastern Europe includes the entire native ranges of indigenous Astacus leptodactylus sensu lato and A. pachypus. This region has been largely overlooked and considered relatively safe from the adverse impacts of non-indigenous crayfish species (NICS). In this study, we evaluated the crayfish pet trade in the Lower Volga Region with special emphasis on Astrakhan, the biggest city of Russian Federation in the region located just in the delta of Volga River, thus being a potential gateway of introductions to the Caspian Sea and adjacent freshwaters. The local pet trade involves 12 NICS. Considering their origin, availability, probability of establishment, invasiveness and further aspects, Procambarus fallax f. virginalis, P. clarkii and Cherax destructor are considered potentially the most problematic, including transmission of diseases like crayfish plague caused by Aphanomyces astaci or white spot syndrome virus. Taking this information as a whole, the availability of NICS with a high probability of overlapping the entire range of European ICS means that attention is warranted. Further research is needed to corroborate the abilities of NICS and their associated diseases to withstand specific conditions of the Caspian Sea as well as the adjacent Black and Azov Seas, all possessing different degrees of elevated salinity.  相似文献   

11.
The feeding habits and trophic ecology of Mustelus lunulatus and Mustelus henlei in the central coast of the Colombian Pacific were evaluated and compared to determine whether there was trophic niche overlap or resource partitioning between these two sympatric shark species. A total of 59 prey items were identified and grouped into 10 taxonomic categories. Mustelus lunulatus fed in large proportion on Stomatopoda and Brachyura, whereas M. henlei fed almost exclusively on Teleostei. Dendrobranchiata, Cephalopoda, Anomura and Polychaeta complemented the diets of both species. There were significant differences in diet between the two species, and there was an ontogenetic diet shift in Mustelus lunulatus, with Stomatopoda being the main prey item of juveniles and Brachyura the main prey item of adults. Dietary overlap (by sex and size) was low between species, but it was high for each species, with significant overlap between juveniles and adults of M. lunulatus and M. henlei, and between males and females of M. henlei. There were differences in the trophic levels of the species, sexes and sizes. This study suggests there is food resource partitioning, and differing ecological function of the two Mustelus species in the food web of the study area.  相似文献   

12.
The introduction of some crustacean species has produced alterations of freshwater environments and declines of native species worldwide. The red swamp crayfish, Procambarus clarkii Girard, was introduced in the Southwest Iberian Peninsula in the 1970’s, producing severe impacts on rice agriculture and on native biota such as macrophytes, gastropods, native crayfish and amphibians. We studied the distribution of P. clarkii in two areas of SW Iberian Peninsula: the Sado River basin (SW Portugal), an area colonized by this species around 1990, and the Parque Natural del Entorno de Doñana (SW Spain), colonized soon after its introduction in the Iberian Peninsula, in the 1970’s. Our main goal was to determine which factors limit crayfish distribution, which could help to identify the most effective management practices to contain its spread. Procambarus clarkii was found in most types of water bodies, including small and shallow ones. Distance to a crayfish source was the single predictor variable explaining crayfish occurrence in most types of habitats and in both areas. The only exception was for the Sado permanent stream points, where crayfish presence was negatively affected by an interaction between elevation and flow velocity. Other habitat characteristics have apparently little or no importance for its successful colonization. Moreover, this study indicated that overland dispersal is apparently a frequent phenomenon in this species. Our findings can be used to determine which habitats are most likely to be colonized by the crayfish and to develop practical measures which may limit its spread and minimize its impacts.  相似文献   

13.
Protracted drought in southern Australia has degraded the system function of the Coorong estuary and intensified resource competition among forage fish species. The present study investigates the gut content, prey composition and dietary overlap of three forage fishes: smallmouth hardyhead (Atherinosoma microstoma), Tamar River goby (Afurcagobius tamarensis) and sandy sprat (Hyperlophus vittatus) influenced by environmental variation in the Murray Estuary and Coorong. The prey species identified in fish stomachs were dominated by crustaceans (amphipods, ostracods and harpacticoids), but nematodes and acanthocephalans were also common in all forage fishes. The diet of the sandy sprat and Tamar River goby highly overlapped (α = 0.8) in the Murray Estuary and all three forage fishes showed potential diet overlap (α ≥ 0.6) in the North Lagoon. Spatiotemporal variation of prey diversity was observed in smallmouth hardyhead while temporal variation of prey diversity was observed in sandy sprat and Tamar River goby. Overall, the prey abundance was temporally variable and predominantly regulated by salinity, pH, dissolved oxygen, water transparency and chlorophyll a in the Murray Estuary and Coorong. This study adds to our knowledge on dietary overlap and resource partitioning among small-bodied forage fishes mediated by environmental factors in the Murray Estuary and Coorong.  相似文献   

14.
Carbon dioxide has been found to produce various negative consequences for a number of aquatic species and is projected to rise in the future for many aquatic ecosystems. Crayfish act as indicators of water quality and function as keystone species in aquatic food webs; however, there is a paucity of research on how crayfish may respond to elevated CO2. This study quantified how shelter-seeking behavior in freshwater crayfish (e.g., family Cambaridae), a behavior critical for survival and fitness, may change following exposure to elevated pCO2. Red Swamp crayfish (Procambarus clarkii, Girard, 1852) were exposed to one of three potential levels of dissolved CO2 that could be found in freshwater basins currently or under future climate change conditions: Control (2 levels as the respective initial exposure. The treatment aquarium contained a shelter and was divided into three equal sections based on proximity to that shelter. Crayfish proximity to the shelter (defined by the tank sections) in the treatment aquarium was monitored every 5 s for a 2-min trial. Crayfish spent differing amounts of time in differing zones of the experiment and had different levels of activity, depending on their pCO2 exposure; crayfish acclimated to High pCO2 increased their time spent hiding and decreased their overall activity when compared to the Low pCO2 and Control treatments. Augmented shelter-seeking behavior may affect crayfish social hierarchies, feeding, mating, and mortality, which could generate cascading effects on the ecology of many freshwater ecosystems.  相似文献   

15.
The Chinese mitten crab (Eriocheir sinensis) is an invasive crustacean with a widespread distribution, sometimes overlapping with the red swamp crayfish (Procambarus clarkii). E. sinensis performs long river migrations, often moving overland to cross obstacles, but the factors affecting its movement are not well known. We studied its movement speed in and out of water, the survival time out of water and also its interaction with the red swamp crayfish during dispersal. At 16°C the LT50 (time to death of 50% of the individuals) was 70 h, with a mean weight loss of 20%, and at 24°C the LT50 was 31 h, with a mean 22% weight loss. In and out of the water, these crabs moved with mean velocities of 0.074 and 0.016 m s?1, respectively. P. clarkii did not affect the dispersion of E. sinensis. This crab was able to disperse under various environmental conditions and its speed was affected by density, temperature, relative humidity, carapace length, and condition of the individuals (Fulton index). Moreover, we found that this species has a great capacity to resist desiccation. These characteristics give them all the requirements to successfully invade and adapt to new areas.  相似文献   

16.
Stomach contents from 1221 Pacific spiny dogfish (Squalus suckleyi) were collected from the Gulf of Alaska over 2004–2006 and analyzed to determine predominant prey species. Pacific spiny dogfish in the Gulf of Alaska have a variable diet suggesting that they are generalist feeders. Of the 68.30% of stomachs with prey contents, the most important prey groups based on the percent prey-specific index of relative importance (%PSIRI) were shrimp (27.06%), cephalopods (17.16%), and forage fish (17.11%). Remaining components of the diet were inconsistent. Commercially valuable species, such as salmon, Oncorhynchus spp., and rockfish, Sebastes spp. constituted only 2.37% and 1.51% of the diet of Pacific spiny dogfish, respectively. Diet diversity was not significantly influenced by spatial, temporal or ontogenetic factors; however, trophic level of prey and average prey item weight varied both interannually and with ontogeny. With increasing size, Pacific spiny dogfish incorporate larger prey items into their diet, and diet composition may be driven more by seasonal availability and prey size than any other factors.  相似文献   

17.
Syngnathus abaster is an euryhaline species, which has penetrated freshwaters in Europe and its range is expanding. The goal was to investigate the feeding patterns of this species in an invaded freshwater habitat, where it first appeared at the beginning of 1990s, with implications for assessing its possible effects on invaded ecosystems. In total, 36 prey items were identified in the diet of S. abaster. The most important prey were zooplanktonic organisms, among which copepods predominated significantly. The second most important prey were cladocerans, represented by 20 species/genera. The diet composition of this species showed clear seasonal dynamics but copepods remained the predominant prey item throughout the year. Significant positive relationships between the abundance of zooplanktonic organisms in the gut contents of S. abaster and their abundance in zooplankton samples were found for copepods and Ceriodaphnia sp. S. abaster showed a significant positive electivity for copepods throughout all months. It also positively selected some large cladocerans, which probably shared the same habitat. S. abaster also consumed fish items including gobiid larvae and its own juveniles, when they were abundant in May and especially in June. The possible effect of S. abaster on native ecosystems may include the shaping of zooplanktonic communities through selective predation on copepods. This species is unlikely to be a serious food competitor for native fish inhabiting European freshwaters, the majority of which are cyprinids. Nevertheless, it can have effects on some native fish through predation on their juveniles, which might share the same habitats.  相似文献   

18.
We investigated the dietary habits of the polkadot skate, Dipturus chinensis (Japanese name: Gangi-ei), in the East China Sea by analyzing the stomach contents of 529 individuals. Of these, 18 (3.40%) had empty stomachs. The mean and maximum fullness indices (FIs) were 0.67% and 6.12%, respectively. The mean FI of female skates > 500 mm in total length (TL) was higher than that of males and smaller females. The most dominant prey category was crustaceans, which included shrimps, crabs, and anomurans, followed by fishes. Cephalopods were additionally observed. Dipturus chinensis fed on a wide variety of crustacean taxa, and among them, Caridea and Galatheidae were particularly common. Fishes that could be identified were mostly demersal fishes, such as Callionymidae, Triglidae, and Rajidae. Some juvenile D. chinensis were also observed, indicating that cannibalism occurs in this species. Even though dental sexual dimorphism was observed in large size classes (mainly > 500 mm TL), the diet of males and females significantly overlapped. Size-related dietary shifts from small crustaceans, such as shrimps and anomurans, to relatively fast-swimming prey, such as fishes, crabs, and cephalopods, were observed in all seasons. The composition of these prey categories varied among seasons. Shannon–Wiener diversity indices calculated for each size class ranged from 2.01 to 2.51, indicating D. chinensis are generalist predators. As size increased, trophic level increased from 3.58 to 3.85, indicating that larger specimens are at a slightly higher trophic level than smaller specimens. These results suggest that D. chinensis are middle-trophic-level generalist predators that utilize diverse prey depending on their size and the season, and thus play an influential role in the demersal ecosystem of the East China Sea.  相似文献   

19.
Silver Carp Hypophthalmicthys molitrix (Valenciennes, 1844) are an invasive species in the Mississippi River Basin and their current role in this novel ecosystem is not fully understood. Juvenile Silver Carp can and do occur in great numbers after a successful spawn. These massive schools of small Silver Carp seem to be an obvious prey source for the common predators of the Mississippi River system. The level to which native piscivores are consuming this novel prey item is unclear. Therefore, the goal of this research was to assess the diets of native piscivores collected in Pool 26 and the Open River reach of the Mississippi River. Using diet contents and catch rates of small fishes, selection or avoidance could be determined for predator prey interactions. Then a controlled laboratory experiment was conducted to determine if common predators [White Bass Morone chrysops and Largemouth Bass Micropterus salmoides (Lacepede, 1802)] would select for or against this novel prey in the presence of two native prey fish [Gizzard Shad Dorosoma cepedianum (Lesueur, 1818) and Emerald Shiner Notropis atherinoides]. Understanding how predator-prey interactions occur in a controlled laboratory experiment may provide insight to trends observed in the field. The majority of native piscivores appeared to be avoiding Silver Carp in both reaches and selecting native prey fishes. The order in which prey fish were consumed by both predators showed significant avoidance of Silver Carp. It appears that the trends observed in the field were supported by our controlled laboratory experiment. The effectiveness of different native piscivores to consume Silver Carp may have impacts on future management decisions.  相似文献   

20.
Examining the functional response of predators can provide insight into the role of predation in structuring prey populations and ecological communities. This study explored feeding behaviour and functional responses of planktivorous damselfishes when offered captive reared larvae of crown-of-thorns starfish, Acanthaster sp., with the aim of determining whether these predators could ever play a role in moderating outbreaks of Acanthaster sp. We examined predatory behaviour of 11 species of planktivorous damselfish, testing: (1) the relationship between predator size and predation rate, both within and among fish species; (2) consumption rates on larvae of Acanthaster sp. versus larvae of a common, co-occurring coral reef asteroid Linckia laevigata; (3) maximal feeding rates upon both Acanthaster sp. and L. laevigata; and (4) functional responses of planktivorous fishes to increasing densities of Acanthaster sp. Consumption rates of crown-of-thorns larvae by damselfishes were independent of predator size; however, there was a significant negative relationship between predator size and consumption rate of L. laevigata, when pooling across all predatory species. Some damselfishes, including Acanthochromis polyacanthus and Amblyglyphidodon curacao, consumed larval Acanthaster sp. at a greater rate than for L. laevigata. Most predatory species (all except A. curacao and Pomacentrus amboinensis) exhibited a Type II functional response whereby the increasing feeding rate decelerated with increasing prey density. In addition to revealing that a wide range of planktivorous fishes can prey upon larvae of Acanthaster sp., these data suggest that planktivorous damselfishes may have the capacity to buffer against population fluctuations of Acanthaster sp. Importantly, predators with Type II functional responses often contribute to stability of prey populations, though planktivorous fishes may be swamped by an abnormally high influx of larvae, potentially contributing to the characteristic population fluctuations of Acanthaster sp.  相似文献   

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