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Behaviour and Ecology of Pomacea Canaliculata from Southern Pampas (Argentina)

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Behaviour and Ecology of Pomacea Canaliculata from Southern Pampas (Argentina) Behaviour and ecology of Pomacea canaliculata from Southern Pampas (Argentina) Pablo R. Martín, María E. Seuffert, Nicolás E. Tamburi, Silvana Burela and Lucía Saveanu Laboratorio de Ecología, Departamento de Biología Bioquímica y Farmacia, Universidad Nacional del Sur, Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Argentina. E-mail:[email protected], [email protected], [email protected], [email protected], lsaveanu@ conicet.gov.ar Abstract Pomacea canaliculata is in many respects the best known species of apple snails (family Ampullariidae), although the available information is both fragmentary and geographically biased. Most studies in its non-native range have focused on applied aspects in managed or artificial wetlands in various countries in SoutheastAsia. In its natural range the emphasis has been on basic studies of its reproductive biology, ecology and behaviour in populations from small streams at the southernmost extreme of its distribution (Southern Pampas, Argentina). The extreme geographic position and the lotic nature of these populations may have biased some conclusions about the behavioural and ecological traits of P. canaliculata; contemporary evolution and genetic exchange may also have diversified these traits in the non-native range. Even though the ecological information from native populations may not be directly applicable elsewhere, it nevertheless remains as a necessary reference to understand the full potential of adaptation and spread of P. canaliculata to new environments around the world. Surprisingly enough, comparative studies of native and non-native populations of Pomacea spp. are almost lacking. This short review focuses on the distribution, thermal biology, aerial respiration, feeding, reproduction, phenotypic plasticity and shell shape of Pomacea canaliculata in its native range in Argentina. Additional keywords: Ampullariidae, distribution, feeding, life-history, Mollusca, phenotypic plasticity, respiration, thermal biology, shell shape 241 Introduction In many respects Pomacea canaliculata is the best known species in the species-rich family Ampullariidae. Most of the interest in this species in recent decades has been spurred by its fast spread and major ecological and agricultural impacts outside its native range (Cowie, 2002; Joshi & Sebastian, 2006; Horgan et al., 2014). The IUCN Invasive Species Specialist Group listed P. canaliculata among 100 of the World’s Worst Invasive Alien Species (Lowe et al., 2000) because of its serious impacts, but it is also as an illuminating example of patterns and processes in biological invasions. The publication of ecological studies on this species in its non-native range have flourished in the 21st century; however, several of these studies erroneously or dubiously identified P. canaliculata as their object of study, probably because of the unnoticed presence of other alien Pomacea spp. and a mostly shell-based taxonomy (Cowie et al., 2006; Hayes et al., 2012, 2015). Fig. 1. A section of the Pigué stream, one of the long streams inhabited by Pomacea canaliculata in the plains of the Encadenadas del Oeste basin (Buenos Aires province, Argentina). (Photo: S. Burela) 242 BIOLOGY AND MANAGEMENT OF INVASIVE APPLE SNAILS Most studies in the non-native range have focused on applied aspects of the ecology of P. canaliculata and other Pomacea spp. in managed or artificial wetlands. In the natural range of P. canaliculata (Río de la Plata basin, South America; Hayes et al., 2012), in contrast, the emphasis has been on basic studies of its reproductive biology, ecology and behaviour in natural habitats. Ecological studies of apple snails in the Pampas region, Argentina, from which apparently most Pomacea invaders have originated (Hayes et al., 2008, 2012; Lv et al., 2013), have concentrated on P. canaliculata (reviewed by Estebenet & Martín, 2002). The aim of this contribution is to review the research done since around 2002 on the behaviour and ecology of P. canaliculata in Southern Pampas and to highlight those aspects that may be useful to understand, predict and control the invasions as well as to understand the ecological and agricultural impacts in the non-native range. Determinants of distribution The natural range of P. canaliculata extends from the Río de la Plata basin southwards to the Encadenadas del Oeste basin in Southern Pampas (37°S, Buenos Aires Province, Argentina), making it the southernmost apple snail in the world (Martín et al., 2001). Factors determining the presence and abundance of P. canaliculata in lotic and lentic waterbodies at its southernmost limit have been studied at different spatial scales. It is generally found in shallow, quiet and turbid sites with low Na+/(K++ + Mg++) ratios (Martín et al., 2001). The arid conditions in western Buenos Aires Province and the high salinity and temporary nature of most waterbodies probably limit its expansion westwards. On the other hand, the main factor that limits its southward spread has been the hydrological barrier imposed by the Tandilia and Ventania Mountains of southern Buenos Aires Province and not temperature (Martín et al., 2001; Martín & De Francesco, 2006). The upper reaches of streams (altitudes and slopes higher than 230 m above sea level and 3.2 %, respectively) arising from the Tandilia and Ventania Mountains are mostly uninhabited by P. canaliculata (Martín et al., 2001, Seuffert & Martín, 2013a). Heavy rains in the steeper headwater sections result in floods that sometimes eradicate P. canaliculata populations, making headwaters unfavourable habitats for their mid and long term persistence. In the plains of the Encadenadas del Oeste basin, P. canaliculata is generally found in the long streams arising from the Ventania Mountains (Fig. 1), while the short streams BEHAVIOUR AND ECOLOGY OF POMACEA CANALICULATA FROM SOUTHERN PAMPAS (ARGENTINA) 243 that arise from springs in the plains are rarely inhabited (Seuffert & Martín, 2013a). In long streams, P. canaliculata is most frequently recorded in places close to the shore, with low current velocities, fine sediments rich in organic matter and abundance of submersed macrophytes. The association with microhabitats in proximity to the shore or with available emergent substrata is due in part to the need to ventilate the lung periodically (Seuffert & Martín, 2009, 2010). Short streams are characterized by higher conductivities, more alkaline pH, very low or zero current velocities and the absence of trees on the margins (Seuffert & Martín 2013a); during hot summer days water temperatures are high enough to induce snail inactivation (Seuffert et al., 2010). Evidence from two sink populations showed that P. canaliculata can survive and reproduce in the short streams, although these streams are not suitable for long term persistence of the populations. Knowledge of the determinants of the distribution of P. canaliculata in its natural realm is necessary to predict its potential habitat range in invaded areas and for an efficient design of barriers to its spread. However, unpredictable variation in climatic and hydrological conditions generates stochastic patterns of extinction-colonization that blur the relationships between snail abundance and environmental factors (Seuffert & Martín, 2013a). A metapopulational approach seems necessary to fully understand and predict the distribution of this snail in its natural and invaded ranges. Thermal biology The activity of P. canaliculata decreases sharply below water temperatures of 15 °C, and the snails are totally inactive at 10 °C. Activity levels also decrease above 30 °C, especially in poorly aerated or fouled waters (Seuffert & Martín, 2010). Crawling speed increases linearly with temperature, but the time spent crawling is nearly constant. The time spent feeding increases with temperature between 10 and 25 °C but drops at higher temperatures (Seuffert et al., 2010). Juvenile growth rates increase with temperature from 15 to 25 °C but do not show a further increase up to 35 °C, probably because of the decline in feeding time and the increase in metabolic rates (Seuffert & Martín, 2013b). The duration of incubation of the aerial egg masses of P. canaliculata is highly dependent on air temperature (Pizani et al., 2005). The lower threshold for embryonic development is around 16 °C, while the cumulative degree-days until hatching range between 120 and 132 (Seuffert et al., 2012). The egg masses of P. canaliculata have 244 BIOLOGY AND MANAGEMENT OF INVASIVE APPLE SNAILS been the target of various chemical and mechanical control methods (Wang et al., 2012). Degree-day models of egg development coupled with field temperature data can be used to predict time required to hatch and thereby to optimize the frequency and timing of the application of control methods to improve their effectiveness (Seuffert et al., 2012). At the southernmost limit of its distribution (37ºS), P. canaliculata remains inactive during the five or six colder months but does not fall into deep dormancy, as the snails soon become active if water temperature rises (Seuffert et al., 2010). Even though P. canaliculata can survive in somewhat colder regions (Colorado river, 39.5°S; Martín et al., 2001), low temperatures are probably one of the main impediments to its expansion to higher latitudes in invaded areas. Behaviour in flowing water Pomacea canaliculata has traditionally been considered a lentic dweller but it also thrives in lotic systems (Martín et al.,
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