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Relative Importance of Propagule Size and Propagule Number for Establishment of Non-Indigenous Species: a Stochastic Simulation Study

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Relative Importance of Propagule Size and Propagule Number for Establishment of Non-Indigenous Species: a Stochastic Simulation Study Aquatic Invasions (2016) Volume 11, Issue 1: 101–110 DOI: http://dx.doi.org/10.3391/ai.2016.11.1.11 Open Access © 2016 The Author(s). Journal compilation © 2016 REABIC Research Article Relative importance of propagule size and propagule number for establishment of non-indigenous species: a stochastic simulation study 1,2 3 David Drolet * and Andrea Locke 1Department of Health Management, Atlantic Veterinary College, University of Prince Edward Island, 550 University Avenue, Charlottetown, Prince Edward Island, C1A 4P3 Canada 2Current address: Fisheries and Oceans Canada, Institut Maurice-Lamontagne, 850 route de la Mer, Mont-Joli, Québec, G5H 3Z4 Canada 3Department of Fisheries and Oceans Canada, Gulf Fisheries Centre, P.O. Box 5030, Moncton, New Brunswick, E1C 9B6 Canada E-mail: [email protected] (DD), [email protected] (AL) *Corresponding author Received: 11 February 2015 / Accepted: 23 September 2015 / Published online: 8 December 2015 Handling editor: Charles W. Martin Abstract Propagule pressure is emerging as the most consistent predictor of establishment in non-indigenous species. Increasing propagule size (the number of individuals arriving in a novel environment at one time) is thought to increase probability of establishment by counteracting demographic stochasticity and Allee effects. Increasing propagule number (the number of introduction events) is thought to increase probability of establishment by counteracting environmental stochasticity. However, the relative importance of these effects and the conditions under which one effect may become predominant is largely unexplored. We first used stochastic population simulations, with a constant number of immigrants distributed over varying numbers of introduction events, to determine the relative importance of propagule size and number on the probability of establishment. Simulations were conducted under different environmental stochasticity and Allee effects regimes with parameters set to be representative of species with high risk of invasion. In absence of Allee effects, there was a slight increase in probability of establishment when going from very low to low propagule number. However, when Allee effects were included, probability of establishment consistently decreased with increasing propagule number. This pattern was consistent across a wide range of parameter values, and tended to be more pronounced when other model parameters resulted in low probability of establishment. This suggests that, within a range of conditions representative of non-indigenous species at high risk of invasion, large infrequent introductions are riskier than smaller more frequent ones. This is particularly true when Allee effects are likely to influence population dynamics at the early stages of the invasion process. Key words: propagule pressure, invasive species, colonization, stochastic population models, Allee effects Introduction population will persist (e.g., Cassey et al. 2004; Memmott et al. 2005; Korsu and Huusko 2009; A major goal of invasion biology is to predict Blackburn et al 2013; Britton and Gozlan 2013). establishment and spread of a newcomer following In addition, propagule pressure may confound its arrival in a novel environment. Initially, species traits (Simberloff 2009; Colautti et al. 2006), research focused on identifying the species-level overwhelm community characteristics (Von Holle traits of successful invaders (e.g., Rejmanek and and Simberloff 2005), and overcome unfavourable Richardson 1996) and the characteristics making abiotic conditions (Warren et al. 2012) in explaining a community prone to invasions (e.g., Levine et establishment. al. 2004). However, it is becoming increasingly Propagule pressure can be decomposed into two clear that propagule pressure (the total number of parts: propagule size (the number of individuals individuals introduced to a novel environment arriving during an introduction event) and propagule within a given time interval) is the most consistent number (the number of introduction events) predictor of establishment (Lockwood et al. 2005; (Simberloff 2009). These two components are Simberloff 2009). Larger numbers of immigrants thought to influence probability of establish- increase the probability that an introduced ment through different mechanisms. Propagule 101 D. Drolet and A. Locke size can influence the severity of Allee effects: a propagule size and number for the probability of positive density-dependence in population growth establishment of sexually reproducing species, where rates (Courchamp et al. 1999). Allee effects were Allee effects are likely, is still unknown from detected in experimental introductions (Grevstad empirical studies. 1999a; Fauvergue and Hopper 2009) and theoretically Stochastic population models have been used shown to reduce probability of establishment widely in conservation biology. For example, they (Grevstad 1999b; Drake and Lodge 2006), i.e., can be used to evaluate the probability that an introductions based on small propagule size were endangered population will persist over a certain more likely to fail when Allee effects were present. time span (population viability analyses; Akçakaya Propagule size is also thought to affect the and Sjögren-Gulve 2000) and the smallest population influence of demographic stochasticity. Small size that keeps the probability of extinction below propagule sizes are more likely to fail in establishing a specified level (minimum viable population; populations because random variation in reproduction Traill et al. 2007). Establishment, the reverse of and survival induces relatively more pronounced extinction, can be used to investigate colonization fluctuations in small than large populations (Lande processes (e.g., Drake and Lodge 2006). Such models 1993; Simberloff 2009). Conversely, propagule number have been used by Grevstad (1999b) to inform is expected to decrease the effect of environmental best release strategies for biocontrol agents when stochasticity; more frequent introductions temporally a fixed number of individuals is available. She spread the risk and make it less likely that short- found that, at high levels of environmental term occurrence of poor conditions will eliminate stochasticity, the probability that at least one popu- all immigrants arriving over the time period lation persisted was greater if fewer individuals (Grevstad 1999b; Simberloff 2009). were released at multiple independent sites. Three empirical studies have attempted to Conversely, in the presence of Allee effects, larger evaluate the relative importance of propagule introductions at fewer independent sites were size and number on the probability of population more effective. In presence of both, an optimal establishment, and none have addressed the release strategy could be determined, depending influence of Allee effects. Drake and Lodge (2005) on the severity of both Allee effects and varied propagule size and number in laboratory environmental stochasticity. In a similar study, populations of the parthenogenetic freshwater Wittmann et al. (2014) investigated time taken by a zooplankter Daphnia magna Straus, 1820. They population to reach a certain size under various found that population size at the end of the scenarios of propagule size and number, while experiment was positively influenced by propagule keeping propagule pressure constant. They found number, with more frequent introductions resulting that, in easy conditions (i.e. positive population in larger populations. However, D. magna being growth rate at all population sizes) populations parthenogenetic eliminates the potential role of grew faster with smaller, frequent introductions, reproductive Allee effects. Hedge et al. (2012) whereas the converse was true in difficult conditions varied propagule size and number (keeping the (i.e. negative population growth rate at some total number of individuals constant) using population sizes). larvae of the oyster Crassostrea gigas (Thunberg, Here we developed stochastic population models 1793) in a field experiment. They found that to investigate the relative importance of propagule settlement was higher with more frequent smaller size and number on the probability of establish- introductions, which was attributed to the effect ment in a situation where introductions are non- of environmental stochasticity and/or facilitation independent, i.e. immigrants influence local among settlers of different ages. However, the population dynamics. This was first done under experiment terminated while the populations different regimes of environmental stochasticity were still only composed of juveniles, making and Allee effects, with parameters chosen to be potential reproductive Allee effects undetectable. representative of species posing a high invasion Finally, Britton and Gozlan (2013) manipulated risk. We then conducted a sensitivity analysis to propagule size and number (keeping the total evaluate the consistency of the patterns when a number of individuals constant) using the invasive much wider range of conditions is modeled. Our fish Pseudorasbora parva (Temminck and Schlegels, study should be helpful from a management 1846) in experimental ponds over a period of perspective as it allows understanding the vectors three years. This design would allow detection of of introduction more likely to result in invasions. Allee effects, but all replicates established a These vectors can then be prioritized for management population. Therefore, the
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