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.     

Lethal and non-lethal effects of multiple indigenous predators on the invasive golden apple snail (Pomacea canaliculata)

1. We investigated the individual and combined effects of two predators (the climbing perch, Anabas testudineus, and the wetland crab, Esanthelphusa nimoafi) indigenous to wetlands in Laos, on the behaviour and survival of the invasive South American golden apple snail (Pomacea canaliculata). The snail is considered a pest, consuming large amounts of rice and other aquatic vegetation in the region. 2. Snail avoidance reactions to released predator chemical cues were investigated in aquaria while the effects of predators on a mixed snail population were studied in field enclosures that contained native aquatic plants (Salvinia cucullata, Ludwigia adscendens and Ipomoea aquatica). 3. In the aquaria experiment, neonate (2–3 mm) and medium‐sized snails (8–10 mm) responded to fish chemical cues by going to the surface, whereas adult snails (35–40 mm) went to the bottom. In contrast, no size class of snails reacted to chemical cues released by crabs. 4. In the field experiment, fish reduced the abundance of neonate snails, and crabs reduced the abundance of all size classes. The effect of the combined predators could not be predicted from the mortality rate observed in single predator treatments. The survival of neonate and medium‐sized snails was greater and of adults less than expected. The presence of predators did not affect egg production. Snails consumed significant amounts of plants despite the presence of predators. 5. Our findings suggest that some indigenous Asian predators have lethal and sublethal effects on P. canaliculata that depend on snail size and predator type. When in the presence of several predators the response of snails to one predator may either increase or decrease the vulnerability of snails to the others.     

Size-dependent effects of an invasive herbivorous snail (Pomacea canaliculata) on macrophytes and periphyton in Asian wetlands

1. The invasive golden apple snail (Pomacea canaliculata), native to South America, is a serious pest on rice seedlings in south‐east Asia and has also been shown to consume large amounts of macrophytes in natural wetlands, with large effects on ecosystem functioning. Earlier studies suggest that the snail undergoes an ontogenetic diet shift, feeding on algae and detritus as juveniles and shifting to aquatic macrophytes as adults. 2. Here, we study the effects of snail populations with a size‐structure typical of either populations at an invasive front or the size‐structure of established populations. In an enclosure experiment performed in a wetland in Laos, we compared treatments with small snails only (3 mm; invasive treatment) to treatments with small, medium sized (10 mm) and adult (>25 mm) snails (established treatment). The effects of snail grazing on three aquatic macrophyte species and periphytic algae were quantified. 3. We found that snails of all sizes had a strong negative effect on the biomass of all macrophyte species and periphytic algae. There was no evidence of an ontogenetic diet change, i.e. snails in both the invasive and established treatments affected macrophyte biomass. Foraging was size‐dependent in that small snails had higher relative foraging capacity (g plant consumed per g of snail) compared with medium and adult snails. Small snails, therefore, depressed growth of medium snails at increasing densities through exploitative competition for preferred resources, while adult snails did not grow at all in the presence of small snails. 4. Density dependence is common in freshwater invertebrates, including gastropod populations, but differences in size dependent foraging‐ and competitive‐ability have rarely been demonstrated in this group of organisms. Knowledge about intra‐specific differences in ecological performance may, however, both deepen our understanding of the processes that underlie population dynamics in invertebrates such as gastropods, and help develop control strategies for invasive golden apple snails.     

Regeneration of excised shell by the invasive apple snail Pomacea canaliculata

After excision of shell from Pomacea canaliculata, survival rates, repair process, haemocyte response, and calcium content were observed and quantified for three weeks. The following experimental treatments were used: black-colored individuals with large (BL-group) or small (BS-group) shell heights, yellow-colored individuals with large (YL-group) or small (YS-group) shell heights. The survival rates in BL-, YL-, BS-, and YS-groups after making partial excisions of snail shells were, respectively, 93.59, 94.87, 92.31, and 96.15%. Freshly regenerated shell could be seen at 1 day among the four groups after induction of shell regeneration. Regeneration shell filled the wound after 5–10 days. The area of regenerated shell in yellow-colored individuals was larger than that of black-colored individuals, but the thickness of regenerated shell was not significantly different. Multiple (four times) inductions of shell regeneration significantly enhanced the thickness of newly formed shells. Two types of circulating haemocytes (hyalinocytes and granulocytes) were identified under light microscopy based on Wright’s staining. Artificial induction of shell regeneration rendered a significant increase in the number of total circulating haemocytes which was followed by a decrease to pre-wounding levels. This peak value of total circulating haemocytes was 2.86, 2.43, 1.69, and 1.71-fold higher than pre-wounding levels in group BL, YL, BS, and YS, respectively. The number of total circulating haemocytes in yellow-colored individuals was significantly higher than in black-colored. Calcium concentrations in the mantles of snails are favorable for calcium nucleation. It is postulated that calcium content increased in the mantle increase at the start of the experiment and then returned to pre-wounding levels at the end of the experiment. This study shows that apple snails have effective shell regeneration abilities and that haemocytes and calcium transportation appear to play an important role in shell growth and regeneration.     

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

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常用萃取剂提取福寿螺卵中类胡萝卜素的初步研究

【背景】福寿螺为世界性恶性入侵水生动物,也是我国公布的第一批外来人侵物种之一。福寿螺大量啃食为害水稻、茭白、白莲等重要农作物,对我国南方各省农业生产造成了巨大威胁,因此,预防和控制福寿螺灾害显得尤为重要。合理利用福寿螺能有效控制福寿螺的数量和危害,是生物防治的一个重要部分。福寿螺卵中含丰富的类胡萝卜素,充分利用螺卵中的类胡萝卜素能拓展福寿螺的利用途径和方法。【方法】为探寻福寿螺卵中类胡萝卜素的提取方法,本研究采用甲醇、无水乙醇、丙酮、乙酸乙酯、二氯甲烷、石油醚等6种常用萃取剂提取福寿螺卵中类胡萝卜素,用紫外可见光分光光度计测定其含量。【结果】结果显示,不同萃取剂中类胡萝卜素的提取量不同,醇类为较适合的提取液（甲醇〉无水乙醇〉丙酮）。【结论与意义】本研究对福寿螺卵中类胡萝卜素的提取方法进行了探索和研究,找出了合适的提取液,为拓展福寿螺的利用途径,以及福寿螺的综合防治提供了参考。     

福寿螺的寄主植物及其对福寿螺体重的影响

为了明确福寿螺对寄主植物的选择性,进一步掌握其可能的危害及入侵暴发机制,对稻田水域及周边的24科43种植物进行了室外取食试验,分别在取食后3、5d对受试样品及成螺称重,比较其无选择条件下的取食偏好性及其对成螺增长率的影响。结果表明:福寿螺对受试的24科43种植物(样品)的根、茎、叶、果实以及块根(茎)均能取食,对水生(湿生)和陆生植物没有偏好性;在取食过程中,优先取食细嫩的植株和部分,硬老部分需经水浸泡软化后才能被取食;辣椒、苦瓜等果实的辣苦味对福寿螺无驱避作用;取食率与体重增长率呈显著正相关,取食越多生长越快;但体重与植物的总酚含量呈负相关。福寿螺的广食性、取食量、植物总酚含量、植物丰盛度等种群和生态系统特征是影响其快速扩散的重要原因,同时可以借助福寿螺对植物的不同利用类型防范其扩散和危害。     

Spatial variation in adult sex ratio across multiple scales in the invasive golden apple snail,Pomacea canaliculata

Adult sex ratio (ASR) has critical effects on behavior and life history and has implications for population demography, including the invasiveness of introduced species. ASR exhibits immense variation in nature, yet the scale dependence of this variation is rarely analyzed. In this study, using the generalized multilevel models, we investigated the variation in ASR across multiple nested spatial scales and analyzed the underlying causes for an invasive species, the golden apple snail Pomacea canaliculata. We partitioned the variance in ASR to describe the variations at different scales and then included the explanatory variables at the individual and group levels to analyze the potential causes driving the variation in ASR. We firstly determined there is a significant female‐biased ASR for this species when accounting for the spatial and temporal autocorrelations of sampling. We found that, counter to nearly equal distributed variation at plot, habitat and region levels, ASR showed little variation at the town level. Temperature and precipitation at the region level were significantly positively associated with ASR, whereas the individual weight, the density characteristic, and sampling time were not significant factors influencing ASR. Our study suggests that offspring sex ratio of this species may shape the general pattern of ASR in the population level while the environmental variables at the region level translate the unbiased offspring sex ratio to the female‐biased ASR. Future research should consider the implications of climate warming on the female‐biased ASR of this invasive species and thus on invasion pattern.     

广东省福寿螺形态性状变异及其与体质量的关系

【背景】福寿螺是危害极其严重的入侵我国的水生生物,目前利用相关分析、通径分析对福寿螺形态性状变异的研究较少。【方法】随机采集7个地区的福寿螺(雌雄比例差别很大),测量其壳高、壳宽、口宽、层高4个形态性状和体质量,采用相关分析、通径分析方法计算福寿螺的各形态指标变异系数、相关系数和通径系数,剔除对体质量影响不显著的指标,确定每个地区与体质量最相关的指标。【结果】除惠州之外,其余地区福寿螺的壳高和壳宽与体质量的相关性较高,且口宽与壳高共同作用对体质量的直接影响很大。【结论与意义】此结果为研究不同生境、不同温度下福寿螺的形态性状变异程度奠定了基础,同时为福寿螺形态鉴定、分类及其灾害预测预报提供了基础数据。     

Survivorship of geographic Pomacea canaliculata populations in responses to cold acclimation

1. Pomacea canaliculata, a freshwater snail from South America, has rapidly established natural populations from south to north subtropical region in China, since its original introductions in the 1980s. Low temperature in winter is a limiting factor in the geographic expansion and successfully establishment for apple snail populations. There have been some studies on population level of low temperature tolerance for P. canaliculata, yet little is quantified about its life‐history traits in responses to cold temperatures. Whether these responses vary with the acclimation location is also unclear. We investigated the survivorship and longevity of P. canaliculata in responses to cold temperatures and examine whether these responses vary with the location and snail size. We hypothesized that survival of the snails depends on their shell height and the level of low temperature, and P. canaliculata population from the mid-subtropical zone may exhibit the highest viability over the cold thermal range.
2. We sampled P. canaliculata populations from five latitude and longitude ranges of subtropical China: Guangzhou population in southernmost (SM‐GZ), three populations of Yingtan (MR‐YT), Ningbo (MR‐NB), Ya'an (MR‐YA) in midrange, and Huanggang population in northernmost (NM‐HG) subtropical zone. For each P. canaliculata population, survival and longevity at six cold acclimation temperature levels (12, 9, 6, 3, 0, and ?3°C) were quantified, and the effects of location and shell height were examined.
3. The MR‐YA population from mid-subtropical zone of China exhibited the highest survival rate and prolonged survival time regardless of the temperature acclimation treatments, whereas the SM‐GZ population from southern subtropical was the most sensitive to cold temperatures, particular temperatures below 9°C. No individuals of the SM‐GZ population could survive after stressed for 30 days (3°C), 5 days (0°C) and 2 days (?3°C), respectively. For each experimental P. canaliculata population held at 3, 0, and ?3°C, individuals with intermediate shell height of 15.0–25.0 mm had significantly higher survivals.
4. The results highlight a request of a more thorough investigation on acclimation responses in each of the life table demographic parameters for P. canaliculata, and pose the question of whether natural selection or some genetic changes may have facilitated adaptation in invasive locations.

     

The salinity tolerance of the invasive golden apple snail (Pomacea canaliculata)

We exposed snails of an invasive species of golden apple snail (Pomacea canaliculata) to five artificial sea water treatments at salinity levels of 0, 5, 10, 15 or 20 parts per thousand (ppt) to assess their salinity tolerance. We observed the behaviour, heart rate, total haemocyte counts, haemolymph ionic concentration and Na⁺/K⁺-ATPase activity in the mantle at 0, 12, 24, 48, 72 and 96 h post salinity exposures. The heart rate declined with increasing salinity, while Na⁺/K⁺-ATPase activity in the mantle presented a reverse trend, possibly to maintain normal osmolality. A trend of rising total haemocyte count was observed from 0 ppt and 5 ppt to 10 ppt salinities, while a sudden increase in the count was observed at 15 ppt and 20 ppt salinity groups. Furthermore, haemolymph Cl^?, Na⁺ and K⁺ concentrations increased directly with elevated salinity. An additional trial was performed to assess the growth performance of the snails under exposure to low salinities. During a 1 month trial, snails grew better at 5 ppt salinity treatment. Taken together, our results demonstrate that P. canaliculata can tolerate salt stress to some extent. The finding also obviously implies a possible invasive risk to estuaries.     

Toward the Molecular Deciphering of Pomacea canaliculata Immunity: First Proteomic Analysis of Circulating Hemocytes

Pomacea canaliculata is a freshwater snail with interesting biological features that include invasiveness, human parasite hosting, and adult regeneration. Its immune system may represent the target for strategies aimed at controlling the spread of the snail population and its hosting of the human parasite Angiostrongylus cantonensis. Moreover, immune functions likely have a role in the snail's ability to wound heal and regenerate. Despite its importance in multiple processes, very little is known about the molecular basis of P. canaliculata immunity. Aiming to contribute to filling this gap, the ultrastructure of circulating hemocytes in healthy snails is studied and the first proteomic analysis of these cells is performed, evidencing 83 unique proteins, 96% of which have identifiable homologs in other species. Fifteen proteins are retrieved as potentially involved in immune‐related signaling pathways, such as hemocyanin, C1q‐like protein, and HSP90 together with cytoskeleton and cytoskeleton‐related proteins involved in cell motility and membrane dynamics. This first proteome study on non‐stimulated hemocytes provides a valid reference for future investigations on the molecular changes under stressful circumstances, like pathogen exposure, wounding, or environmental changes.     

Biological control of the exotic invasive snail Pomacea canaliculata with the indigenous medicinal leech Whitmania pigra

The apple snail Pomacea canaliculata has been an invasive species in China for decades and causes enormous losses to agriculture. The predation capability of the leech Whitmania pigra on P. canaliculata was studied for its economic benefit and potential application. In the present study, the leech W. pigra exhibited a strong predatory capacity in controlling P. canaliculata under both laboratory and field conditions, and it showed no bias towards consuming smaller snails during the experiments. More than 80% of the 80 snails (of which half had reached maturity) were preyed upon by 20 developing leeches (2–7?g) in miniature rice fields over a 15-day period, and the number of rice seedlings damaged by P. canaliculata was decreased in the presence of W. pigra. In a separate experiment, 15 developing snails were exposed to caged leeches and conspecific snails for four weeks. The food intake and growth of P. canaliculata were inhibited under the predation risk imposed by W. pigra, but the feeding rate, food conversion efficiency and survival of P. canaliculata were not conspicuously influenced.     

Wetland Restoration and Invasive Species: Apple snail (Pomacea insularum) Feeding on Native and Invasive Aquatic Plants

The apple snail Pomacea insularum is an aquatic invasive gastropod native to South America that has the potential to cause harm to aquatic ecosystems, wetland restoration, and agriculture. To predict the potential impact of this snail on aquatic ecosystems, we tested the feeding rate of P. insularum , under laboratory nonchoice experiments, for 3 species of invasive macrophytes and 13 species of native aquatic plants that are important for wetland restoration and health. High levels of consumption were recorded for four native species ( Ceratophyllum demersum , Hymenocallis liriosme , Ruppia maritima , and Sagittaria lancifolia ) and three invasive species ( Colocasia esculenta , Alternanthera philoxeroides , and Eichhornia crassipes ). In contrast, less than 10% of the biomass of Spartina alterniflora , Scirpus californicus , Thalia dealbata , and Typha latifolia was consumed by P. insularum over the test period. The palatability of macrophytes was negatively correlated with dry matter content, making our results generalizable to all regions where this invader may be present. Based on our results, wetland restoration in areas invaded by P. insularum should focus on emergent structural species with low palatability. Apple snails should not be considered as agents of biocontrol for invasive plants; although apple snails fed on invasive plants at a high rate, their consumption of many native species was even greater.     

Changes in chemical components in the freshwater apple snail, Pomacea canaliculata (Gastropoda: Ampullariidae), in relation to the development of its cold hardiness

The apple snail, Pomacea canaliculata, is an invasive freshwater snail. It increases its cold hardiness before winter. However, the physiological mechanism of cold hardiness in molluscs is poorly understood, especially in freshwater molluscs. In this study, we examined the changes in low molecular weight compounds, glycogen and lipids, in the body of P. canaliculata in association with the development of cold hardiness. When snails without cold hardiness were experimentally cold-acclimated, the amount of glycerol, glutamine, and carnosine increased, while glycogen and phenylalanine decreased. Overwintering cold-tolerant snails collected from a drained paddy field in November also showed increased glycerol in their bodies with decreasing glycogen concentration, compared to summer snails collected from a submerged field. Water content also decreased during the cold acclimation, although the water loss was minimal. These results indicate that the freshwater snail, P. canaliculata enhances cold hardiness by accumulation of some kinds of low molecular weight compounds in its body as some insects do. However, the actual function of each low molecular compound is still unknown.     

Exceeding its own limits: range expansion in Argentina of the globally invasive apple snail Pomacea canaliculata

Hydrobiologia - Pomacea canaliculata is a freshwater snail native to southern South America. The aims of this work are to update its distribution in Argentina and to analyze through niche models...     

Field observations of the alarm response to crushed conspecifics in the freshwater snail Pomacea canaliculata: effects of habitat,vegetation, and body size

The freshwater snail Pomacea canaliculata shows alarm responses to chemical cues released from injured conspecifics, but its behavioural responses in the field are unknown. We investigated effects of habitat (canals or paddy fields), vegetation, and body size on alarm responses in the field. Snails responded to crushed conspecifics within 4 min by burying themselves, but the proportions of self-buried snails were generally lower (0–28% depending on experiments) than those reported in the laboratory. Snails not only showed the alarm response, but also frequently fed on crushed conspecifics. There were no influences of habitat or body size on the proportions of individuals showing the alarm response. Nevertheless, in paddy fields with high-density vegetation a higher proportion of snails showed the alarm response than in paddy fields with low-density vegetation.     

入侵生物金苹果螺在滇池流域的首次记（英文）

2004年10月,中国科学院昆明动物研究所首次发现金苹果螺(Pomacea canaliculata)入侵重要的水源保护区嵩明白邑黑龙潭。金苹果螺起源于中南美洲,在亚洲,它通过有意或无意的传播而逐渐扩散到菲律宾、越南、泰国、老挝、柬埔寨、马来西亚、印尼、巴布几内亚、韩国、日本和中国的南部。金苹果螺已成为水稻产区的最大害虫,给农业生产带来巨大的损失。为防止金苹果螺在云南扩散,目前已经实施了严格的预防、控制措施,同时开展了公众保护教育宣传活动。