Effects of macrophytes on feeding and life-history traits of the invasive apple snail Pomacea canaliculata

1. Biological invasions have become a serious threat to ecosystems worldwide. Various factors can contribute to the success of biological invasion. We examined how different macrophyte food affected feeding and life‐history traits of the invasive herbivorous snail Pomacea canaliculata, and whether differences in snail life‐history traits could explain its successful infestation of agricultural and non‐agricultural wetlands in Asia. 2. We tested five cultivated and five wild semi‐aquatic macrophytes. Snail daily feeding rate varied substantially with plant species, ranging from 1.3% to 22% of its body mass. Snails fed with four (Amaranthus gangeticus, Apium graveolens dulce, Ipomoea aquatica and Nasturtium officinale) of the five cultivated macrophyte species exhibited high survivorship, fast growth and high fecundity. Snails fed with Colocasia esculenta, however, grew poorly, did not reproduce and eventually died. 3. Of the five wild species (Eichhornia crassipes, Ludwigia adscendens, Murdannia nudiflora, Myriophyllum aquaticum and Polygonum hydropiper), M. nudiflora supported a high snail survival, but snails had slower growth and lower fecundity than those reared on the four palatable cultivated species. Snails fed with L. adscendens grew substantially slower than those fed with M. nudiflora, and produced only a small clutch of eggs. Snails fed with E. crassipes, M. aquaticum and P. hydropiper had very low survivorship, grew very little and did not reproduce. 4. We determined six plant properties and their correlation with the feeding, growth and reproduction of the apple snails. Cultivated macrophytes in general had a higher nutritional value and lower physical and chemical defences. Phenolic content was negatively correlated with snail feeding rate, while plant nitrogen and phosphorus contents were positively correlated with snail egg production and growth, respectively. 5. These results indicate that, due to their higher nutritional value and lower chemical and physical defences, cultivated macrophytes are in general desirable for the apple snail which may partly explain its successful invasion into wet agricultural areas in Asia. This snail may also selectively graze poorly defended wild macrophytes in non‐agricultural wetlands, leading to changes in floral diversity and wetland functioning. Management of this and other apple snails with similar life‐history traits should thus focus on the prevention of their further spread.     

香港东北角吉澳群岛入侵植物调查

什么样的生态系统容易入侵是入侵生态学关心的核心问题之一。最近,我们对香港东北角吉澳群岛中的鸦洲湾、小鸦洲、鸦洲、拦船排、虎王洲、娥眉洲、往湾洲、吉澳等8个岛屿上的所有外来植物及本土植物的种类、数量和盖度进行了调查,结果表明:具有高钙低硅土壤基质的鸦洲、小鸦洲、鸦洲湾等三个小岛上分布有大量外来入侵植物的种类和数量,而具有高硅低钙土壤基质的拦船排、虎王洲、娥眉洲、往湾洲等岛屿上外来植物的种类和数量稀少,吉澳岛虽然外来植物的种类多,但外来植物的个体数量少。分析说明:钙离子可能是影响外来植物的入侵能力或生态系统的可入侵性的关键因子。此外,文章还讨论及介绍了外来植物在石灰岩生境、盐碱地、海岸沙滩、温带广布的各种钙土、地中海气候影响下形成的干旱土等钙离子丰富的生态系统中入侵情况。     

Patterns of density dependence in growth,reproduction and survival in the invasive freshwater snail Pomacea canaliculata in Japanese rice fields

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The effects of nutrients and their ratios on phytoplankton abundance in Junk Bay,Hong Kong

Eutrophication has been considered to be undoubtedly one of the key factors stimulating phytoplankton growth, since it involves the enrichment of a water mass with both inorganic and organic nutrients supporting plant growth. Nutrient enrichment as a result of anthropogenic activity occurs in estuaries and coastal waters as well as in lakes and freshwater impoundments, and blooms of phytoplankton are one of the effects of such an accelerated process of nutrient enrichment. This paper presents the results of a two-year survey of the nutrients and phytoplankton at 3 stations in Junk Bay, Hong Kong, carried out from 1997 to 1998. The relationships between nitrogen, phosphorus, and their ratio, with phytoplankton abundance have been studied. The results show that the highest nitrogen concentration was in Station 2 which is close to a sewage input, whereas the highest phosphorus concentration was in Station 1 which is close to a landfill area. The mean N:P ratios at the three stations were between 8 and 14. The diatoms were the dominant group during most of the year but it seems that diatoms were more sensitive than dinoflagellates and other algal groups to the increase in nutrients.     

贝类壳-体质量比和静水沉降特性的相关性

软体动物贝壳表型是适应性进化的结果,为探究贝壳形态特性与软体动物随水流扩散功能的相关性,实验选取9种具代表性的腹足纲和双壳纲贝类作为研究对象,研究了其成体的壳-体质量比与静水沉降特性及二者的相关性。结果表明：体质量的变异系数（C.V.=1.11）大于壳长（C.V.=0.67）、壳宽（C.V.=0.54）的变异系数;壳-体质量比具较强的种属特异性,种内变异系数小（C.V.=0.20）,而种间差异极显著（P < 0.01）。受试贝类的平均壳-体质量比为0.32±0.13,陆生贝类的壳-体质量比显著小于水生贝类（P < 0.01）,壳-体质量比最大的河蚬（0.54±0.06）是最小的非洲大蜗牛（0.17±0.04）的3倍,福寿螺的壳-体质量比为0.50±0.06。双壳类和螺类的沉降行为差异明显,只有中华园田螺、铜锈环棱螺和河蚬不能在静水中漂浮;沉降速度最大的河蚬（（24.99±4.22）cm/s）是最小的椭圆萝卜螺（（4.13±0.96）cm/s）的6倍,入侵种非洲大蜗牛（（18.30±3.64）cm/s）和福寿螺（（21.77±5.23）cm/s）与土著种铜锈环棱螺（（19.48±3.14）cm/s）和中华园田螺（（21.44±3.92）cm/s）在沉降速度上无显著差异（P < 0.01）。成体贝类的壳-体质量比与沉降速度（R²=0.28）、沉降加速度（R²=0.39）之间存在一定的相关性。在此基础上,贝类沉降特性随生活史变化的研究将进一步揭示贝类进化与种群扩散中的生态水力学作用。     

Biological control of invasive apple snails by two species of carp: Effects on non-target species matter

Molluscivorous fish, especially carp, have been adopted as bio-control agents of the invasive apple snail Pomacea canaliculata, but previous studies have focused on their effectiveness, with little attention paid to their undesirable effects on non-target plants and animals. We conducted an 8-week mesocosm study to compare the effectiveness of two indigenous fish, common carp (Cyprinus carpio) and black carp (Mylopharyngodon piceus), in removing P. canaliculata, and their potential side effects on macrophytes and non-target mollusks in a freshwater wetland. Three species of macrophytes and a community of mollusks in the wetland sediment were enclosed in 1 × 1 × 1 m enclosures either with apple snails (AS), with apple snails and common carp (AS + CC), with apple snails and black carp (AS + BC), or without apple snails and fish. Both species of carp were effective predators of P. canaliculata, removing most of the individuals in the enclosures except a few that were too big to fit into their mouth. By reducing apple snail population, black carp reduced grazing of apple snail on macrophytes. In contrast, although common carp controlled apple snail population, it did not reduce overall loss in plant biomass as the fish might also fed on macrophytes. Both species of carp preyed on non-target mollusks. Application of bio-control agents in invasive species management needs to consider their effects on both the pest and non-target plants and animals. Adoption of common and/or black carp to control apple snail populations thus depends on the weight given to their effectiveness and subtle different effects on non-target organisms by wetland management authority.     

广西北海西村港互花米草对红树林湿地大型底栖动物群落的影响

为了解互花米草(Spartina alterniflora)入侵红树林的生态影响, 作者对位于北海市西村港的红树林湿地以及周边互花米草盐沼的大型底栖动物群落多样性和群落结构进行了研究。2012年10月至2013年9月连续4次取样, 按照取样时间研究大型底栖动物的种类、物种组成、生物量和生物多样性等群落特征的差异, 探讨互花米草入侵红树林湿地对大型底栖动物的影响。本研究共采集底栖动物16种, 隶属于5门7纲15科, 其中互花米草群落10种, 红树林湿地12种。研究发现互花米草入侵后中国绿螂(Glauconome chinensis)个体数量剧增, 导致不同采样时间互花米草盐沼的大型底栖动物生物量均显著高于红树林湿地; 除个别月份外, 红树林湿地大型底栖动物的Margalef丰富度指数、Shannon-Wiener多样性指数、Simpson多样性指数和Pielou均匀度指数均显著高于互花米草群落。基于生境-采样时间的双因素方差分析结果表明, Shannon-Wiener多样性指数和Simpson指数在两种生境间差异显著; 两种生境的Margalef丰富度和Pielou均匀度指数在不同采样时间差异显著; 大型底栖动物生物量和物种数量在两种生境间和不同采样时间差异均显著。基于多元回归分析的研究结果表明, 互花米草密度是影响大型底栖动物生物量的关键因子, 而互花米草株高可以解释物种个体数量、Shannon-Wiener多样性指数和Simpson指数在两种生境的变化。对不同采样时间大型底栖动物群落结构的非度量多维度(non-metric multidimensional scaling, NMDS)分析结果表明, 红树林与互花米草群落的大型底栖动物群落相似性很低。总而言之, 在西村港地区, 互花米草入侵虽然增加了大型底栖动物的生物量, 但由于优势物种的凸显, 显著降低了大型底栖动物群落的多样性, 且种类组成与群落结构与红树林群落相比已有差异。由此可见, 互花米草入侵红树林对当地的大型底栖动物群落多样性造成影响。     

Impact of invasive apple snails on the functioning and services of natural and managed wetlands

At least 14 species of apple snail (Ampullariidae) have been released to water bodies outside their native ranges; however, less than half of these species have become widespread or caused appreciable impacts. We review evidence for the impact of apple snails on natural and managed wetlands focusing on those studies that have elucidated impact mechanisms. Significant changes in wetland ecosystems have been noted in regions where the snails are established: Two species in particular (Pomacea canaliculata and Pomacea maculata) have become major pests of aquatic crops, including rice, and caused enormous increases in molluscicide use. Invasive apple snails have also altered macrophyte community structure in natural and managed wetlands through selective herbivory and certain apple snail species can potentially shift the balance of freshwater ecosystems from clear water (macrophyte dominated) to turbid (plankton dominated) states by depleting densities of native aquatic plants. Furthermore, the introductions of some apple snail species have altered benthic community structure either directly, through predation, or indirectly, through exploitation competition or as a result of management actions. To date much of the evidence for these impacts has been based on correlations, with few manipulative field or mesocosm experiments. Greater attention to impact monitoring is required, and, for Asia in particular, a landscape approach to impact management that includes both natural and managed-rice wetlands is recommended.     

Impact of summer warming on the thermal characteristics of a polymictic lake and consequences for oxygen, nutrients and phytoplankton

1. The impact of long thermal stratification events on some key properties in a polymictic lake was studied by determining the mixing regime of Müggelsee, Germany, using water temperature profiles taken hourly over 4 years. The period included two exceptional summer heatwaves. 2. Long thermal stratification events lasted from about 1 week to 2 months, and exhibited a high variability in thermocline depth and stratification intensity within and between events. 3. During stratification events, hypolimnetic oxygen concentrations strongly decreased while hypolimnetic SRP accumulation increased, depending on the duration and intensity of stratification and on hypolimnetic water temperature. 4. The impact of stratification on the functional phytoplankton composition increased with increasing stratification duration, but was rather different for the heatwaves. 5. Stratification events were followed by strong nutrient pulses into the euphotic zone and intense phytoplankton growth, particularly after the heatwaves. Hence, the influence of the climate extremes counteracted effects of reduced external nutrient loading.     

Cold tolerance of invasive freshwater snails,Pomacea canaliculata,P. maculata,and their hybrids helps explain their different distributions

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福寿螺和田螺消化酶活性比较

为比较福寿螺(Pomacea canaliculata(Lamarck,1828))和当地中国圆田螺(Cipangopaludina chinensis(Gray,1832))消化能力的差异,探索福寿螺成功入侵的机制,以田螺为对照,测定了1—4龄的福寿螺和田螺的胃和肝脏的消化酶——纤维素酶(羧甲基纤维素法)、淀粉酶(3,5-二硝基水杨酸法)和脂肪酶(滴定法)的活性。结果表明:1)相同年龄的福寿螺胃和肝脏中的消化酶活性明显高于田螺。其中,纤维素酶活性分别高出1.00—2.11倍、1.66—2.84倍;淀粉酶活性分别高出1.53—3.47倍、1.47—1.80倍;脂肪酶活性分别高出2.07—4.73倍、6.13—9.93倍。2)在生长发育过程中,福寿螺胃和肝脏中的消化酶活性变化幅度(51.2%—131.2%)明显高于田螺(23.3%—47.1%)。3)福寿螺的各种消化酶之间存在协同作用。如福寿螺的淀粉酶活性与脂肪酶活性呈极显著正相关(胃中r=0.736**、肝脏中r=0.867**)。此外,胃中的淀粉酶活性还与纤维素酶活性呈显著正相关关系(r=0.696*)。相应地,田螺胃中的淀粉酶和脂肪酶之间也存在显著的正相关关系(r=0.706*),而肝脏中的纤维素酶与脂肪酶活性呈显著负相关(r=-0.593*)。4)福寿螺对纤维素类和淀粉类物质都有较强的消化能力,且能较好地消化脂肪类物质,而田螺能消化纤维素类和淀粉类物质,对脂肪的消化能力却很弱。福寿螺的纤维素酶和淀粉酶活性分别是田螺的2.42和1.88倍,脂肪酶活性达到了5.66倍。可见,福寿螺具有较高的消化酶活性,且各消化酶之间存在正协同性。这可能是导致福寿螺食量大、食性杂,使其能快速生长和成功入侵的重要原因之一。     

Compensation effect of phytoplankton on nutrients from a sewage outfall in summer

This paper investigates the relationship between nutrients, chlorophyll a and physical variables in the upper Saronikos Gulf, an oligotrophic marine environment south of the Greater Athens Metropolitan Area. Phosphate, silicate, ammonia, nitrite, nitrate, temperature, salinity and dissolved oxygen were determined at eight stations on 9 occasions during summer 1982. A thermocline led to the stratification of the water column and the pycnocline was related to the thermocline. The values of oxygen were more or less normal. The Eutrophication of the seawater in the vicinity of the sewage outfall was demonstrated by surface levels of chlorophyll a being forty to two hundred times above background. This parameter provides evidence for a high phytoplankton standing stock. However, there were no appreciable differences between the nutrients in the outfall area and those in the background. This suggests rapid uptake of nutrients and/or effective dispersal from the outfall.     

Ecological studies of phytoplankton in Tai Tam Bay,Hong Kong

Of the 141 phytoplankton species found during the sampling period, the Bacillariophyceae was the most important group and included 70% of the total number of species; the Dinophyceae comprised a smaller percentage (28%) of the taxa; and only 3 chrysophyte species occurred throughout the study period. In terms of numbers of individuals, the Bacillariophyceae remained as the dominant group and contained 97% of the total numbers whereas the Dinophyceae represented only 3% of the total phytoplankton population.The most dominant diatom species was Thalassiosira allenii Takano which constituted 35% of the Bacillariophyceae group whereas the most abundant dinoflagellate was Prorocentrum gracile Schutt which represented 40% of the total Dinophyceae numbers.Chlorophyll a levels varied from 0.40 to 32.31 mg m^–3 at the surface and from 0.33 to 33.91 mg m^–3 at the bottom.Seasonal trends of phytoplankton abundance and chlorophyll a concentration were observed in the survey area with generally high peaks in summer and low values during winter months. Such variations can be attributed to the interactions between, and changes of, various environmental parameters, such as temperature, salinity and nutrient availability (particularly silicate).     

Effects of invasive zebra mussels on phytoplankton,turbidity, and dissolved nutrients in reservoirs

Few experiments have quantified the effects of invasive zebra mussels (Dreissena polymorpha) on man-made reservoirs relative to other aquatic habitats. Reservoirs, however, are the dominate water body type in many of the states that are at the current front of the zebra mussel invasion into the western United States. The objective of this research, therefore, was to determine how zebra mussels affected phytoplankton, turbidity, and dissolved nutrients in water that was collected from three Kansas reservoirs that varied in trophic state (mesotrophic to hypereutrophic), but all experienced frequent cyanobacterial blooms. Laboratory mesocosm experiments were conducted to document the effects of zebra mussels on cyanobacteria and general water quality characteristics in the reservoir water. Zebra mussels significantly reduced algal biomass, and the total biovolume of cyanobacteria (communities were dominated by Anabaena) in each reservoir experiment. The effects of zebra mussels on other major algal groups (diatoms, flagellates, and green algae) and algal diversity were less consistent and varied between the three reservoir experiments. Similarly, the effects of zebra mussels on nutrient concentrations varied between experiments. Zebra mussels increased dissolved phosphorus concentrations in two of the reservoir experiments, but there was no effect of zebra mussels on dissolved phosphorus in the mesotrophic reservoir experiment. Combined, our results strongly suggest that zebra mussels have the potential to significantly impact reservoirs as they continue to expand throughout the western United States. Moreover, the magnitude of these effects may be context dependent and vary depending on the trophic state and/or resident phytoplankton communities of individual reservoirs as has similarly been reported for natural lakes.     

Ambiguous climate impacts on competition between submerged macrophytes and phytoplankton in shallow lakes

1. Shallow lakes may switch from a state dominated by submerged macrophytes to a phytoplankton‐dominated state when a critical nutrient concentration is exceeded. We explore how climate change may affect this critical nutrient concentration by linking a graphical model to data from 83 lakes along a large climate gradient in South America. 2. The data indicate that in warmer climates, submerged macrophytes may tolerate more underwater shade than in cooler lakes. By contrast, the relationship between phytoplankton biomass [approximated by chlorophyll‐a (chl‐a) or biovolume] and nutrient concentrations did not change consistently along the climate gradient. In warmer climates, the correlation between phytoplankton biomass and nutrient concentrations was overall weak, especially at low total phosphorus (TP) concentrations where the chl‐a/ TP ratio could be either low or high. 3. Although the enhanced shade tolerance of submerged plants in warmer lakes might promote the stability of their dominance, the potentially high phytoplankton biomass at low nutrient concentrations suggests an overall low predictability of climate effects. 4. We found that near‐bottom oxygen concentrations are lower in warm lakes than in cooler lakes, implying that anoxic P release from eutrophic sediment in warm lakes likely causes higher TP concentrations in the water column. Subsequently, this may lead to a higher phytoplankton biomass in warmer lakes than in cooler lakes with similar external nutrient loadings. 5. Our results indicate that climate effects on the competitive balance between submerged macrophytes and phytoplankton are not straightforward.     

Nutrient and faecal contamination and retention in wetland enclosures (gei wais) in the Mai Po Marshes,Hong Kong

Nutrient and faecal contamination is an increasing problem to the shrimp productivity and wildlife conservation at the internationally important wetland ecosystems of the Mai Po Marshes (Hong Kong, P.R. China). The present study examined the nutrient status and faecal bacteria loading and potential retention capacity of contaminants of two wetland enclosures. Water in the wetland enclosures was eutrophicated with high concentrations of dissolved inorganic nitrogen (inorg–N_diss= 15.0 mg l^–1) and orthophosphate phosphorus (o-P = 1.89 mg l^–1) and was loaded with high levels of faecal coliforms (172 ×10³ cfu in 100 ml) and faecal streptococci (1.94 ×10³ cfu in 100 ml). The pattern of nutrient enrichment of two wetland enclosures is related to a north-to-south pollution gradient from the Shenzhen River to the wetlands. By retaining tidal water in the wetland for an 8-day period, water quality was greatly improved; NH₄–N was removed by 83%, o-P by 45% and faecal bacteria by 100%. This implies a self-purification capability of the wetland enclosures and a potentiality of using them as an alternative sewage treatment.     

江苏沿江地区出口产品仓储昆虫群落结构数量特征研究

对江苏沿江地区出口产品仓储昆虫进行调查,分析了5种出口产品仓储昆虫群落的优势种.利用群落物种的丰富度、生态优势度、多样性和均匀度等群落特征指数,研究了5种类型出口产品仓储昆虫群落结构的数量特征,并分析了它们的相似性.通过系统聚类分析将5种群落分为4类,草柳藤制品和羽绒制品仓储昆虫群落同属一类,其它3种产品仓储昆虫群落各属一类.除木制品群落结构相对合理外,其它群落结构都不合理,皮毛制品群落结构尤其不合理.     

Interactive effects of light and nutrients on phytoplankton stoichiometry

The stoichiometric composition of autotrophs can vary greatly in response to variation in light and nutrient availability, and can mediate ecological processes such as C sequestration, growth of herbivores, and nutrient cycling. We investigated light and nutrient effects on phytoplankton stoichiometry, employing five experiments on intact phytoplankton assemblages from three lakes varying in productivity and species composition. Each experiment employed two nutrient and eight irradiance levels in a fully factorial design. Light and nutrients interactively affected phytoplankton stoichiometry. Thus, phytoplankton C:N, C:P, and N:P ratios increased with irradiance, and slopes of the stoichiometric ratio versus irradiance relationships were steeper with ambient nutrients than with nutrients added. Our results support the light–nutrient hypothesis, which predicts that phytoplankton C:nutrient ratios are functions of the ratio of available light and nutrients; however, we observed considerable variation among lakes in the expression of this relationship. Phytoplankton species diversity was positively correlated with the slopes of the C:N and C:P versus irradiance relationships, suggesting that diverse assemblages may exhibit greater flexibility in the response of phytoplankton nutrient stoichiometry to light and nutrients. The interactive nature of light and nutrient effects may render it difficult to generate predictive models of stoichiometric responses to these two factors. Our results point to the need for future studies that examine stoichiometric responses across a wide range of phytoplankton communities.