A test for Allee effects in the self‐incompatible wasp‐pollinated milkweed Gomphocarpus physocarpus

It has been suggested that plants that are good colonizers will generally have either an ability to self‐fertilize or a generalist pollination system. This prediction is based on the idea that these reproductive traits should confer resistance to Allee effects in founder populations and was tested using Gomphocarpus physocarpus (Asclepiadoideae: Apocynaceae), a species native to South Africa that is invasive in other parts of the world. We found no significant relationships between the size of G. physocarpus populations and various measures of pollination success (pollen deposition, pollen removal and pollen transfer efficiency) and fruit set. A breeding system experiment showed that plants in a South African population are genetically self‐incompatible and thus obligate outcrossers. Outcrossing is further enhanced by mechanical reconfiguration of removed pollinaria before the pollinia can be deposited. Self‐pollination is reduced when such reconfiguration exceeds the average duration of pollinator visits to a plant. Observations suggest that a wide variety of wasp species in the genera Belonogaster and Polistes (Vespidae) are the primary pollinators. We conclude that efficient pollination of plants in small founding populations, resulting from their generalist wasp‐pollination system, contributes in part to the colonizing success of G. physocarpus. The presence of similar wasps in other parts of the world has evidently facilitated the expansion of the range of this milkweed.     

Pollen limitation and Allee effect related to population size and sex ratio in the endangered Ottelia acuminata (Hydrocharitaceae): implications for conservation and reintroduction

Small populations may suffer more severe pollen limitation and result in Allee effects. Sex ratio may also affect pollination and reproduction success in dioecious species, which is always overlooked when performing conservation and reintroduction tasks. In this study, we investigated whether and how population size and sex ratio affected pollen limitation and reproduction in the endangered Ottelia acuminata, a dioecious submerged species. We established experimental plots with increasing population size and male sex ratio. We observed insect visitation, estimated pollen limitation by hand‐pollinations and counted fruit set and seed production per fruit. Fruit set and seed production decreased significantly in small populations due to pollinator scarcity and thus suffered more severe pollen limitation. Although frequently visited, female‐biased larger populations also suffered severe pollen limitation due to few effective visits and insufficient pollen availability. Rising male ratio enhanced pollination service and hence reproduction. Unexpectedly, pollinator preferences did not cause reduced reproduction in male‐biased populations because of high pollen availability. However, reproductive outputs showed more variability in severe male‐biased populations. Our results revealed two component Allee effects in fruit set and seed production, mediated by pollen limitation in O. acuminata. Moreover, reproduction decreased significantly in larger female‐biased populations, increasing the risk of an Allee effect.     

Pollen limitation in invasive populations of Solanum rostratum and its relationship to population size

Aims Small plant populations may be more likely to suffer more severe pollen limitation due to the lower number of potential mates or suitable pollinators. For invasive species, this phenomenon may be more common when an invading population colonizes a new habitat. Here, we investigated whether pollen limitation occurs in invasive populations of Solanum rostratum during its invasion from North America to China and evaluated the patterns between pollen limitation and population size.Methods Pollen addition experiments were performed on six invasive populations of S. rostratum. By comparing fruit set and seed production with open pollination treatment, we calculated the index of pollen limitation and regressed it to population size and density.Important findings Among the six sampled invasive populations of S. rostratum, the fruit set and seed production per fruit were 0.346±0.014 and 52.38±9.29, respectively, with open pollination treatment and 0.572±0.022 and 56.28±10.79, respectively, with pollen addition treatment. Compared with open pollination, pollen addition significantly increased fruit set and seed production by 65.3 and 7.4%, respectively. The standardized index of pollen limitation ranged from 0.022 to 0.125, with an average of 0.065, suggesting that invasive populations of S. rostratum do suffer from pollen limitation. The index of pollen limitation was negatively correlated with population size, which is consistent with the pattern that smaller populations suffer from more severe pollen limitation.     

Density-dependent seed set in the Haleakala silversword: evidence for an Allee effect

Plant species may be subject to Allee effects if individuals experience a reduction in pollination services when populations are small or sparse. I examined temporal variation in reproductive success of the monocarpic Haleakala silversword (Argyroxiphium sandwicense subsp. macrocephalum) over five years, to determine if plants flowering out of synchrony with most of the population (i.e., in low flowering years) exhibited lower percent seed set than synchronously-flowering plants (i.e., those flowering in high flowering years). Through two pollination experiments conducted over multiple years, I also measured pollen limitation and self-incompatibility in this species. The number of flowering plants varied greatly among years, as did reproductive success. Percent seed set was significantly correlated with the number of plants flowering annually, such that plants flowering in high flowering years (1997 and 2001) exhibited significantly higher percent seed set than did plants flowering in low flowering years (1998-2000). In the 3-year pollen limitation study, plants flowering asynchronously were pollen-limited, whereas plants flowering synchronously were not. This species is strongly self-incompatible. Results of this study demonstrate that the Haleakala silversword experiences reduced reproductive success in low flowering years, and suggest that this Allee effect is pollinator-mediated. Allee effects in plants are an understudied yet potentially important force with implications for the population dynamics and conservation of rare species.     

When bigger is not better: intraspecific competition for pollination increases with population size in invasive milkweeds

One of the essential requirements for an introduced plant species to become invasive is an ability to reproduce outside the native range, particularly when initial populations are small. If a reproductive Allee effect is operating, plants in small populations will have reduced reproductive success relative to plants in larger populations. Alternatively, if plants in small populations experience less competition for pollination than those in large populations, they may actually have higher levels of reproductive success than plants in large populations. To resolve this uncertainty, we investigated how the per capita fecundity of plants was affected by population size in three invasive milkweed species. Field surveys of seed production in natural populations of different sizes but similar densities were conducted for three pollinator-dependent invasive species, namely Asclepias curassavica, Gomphocarpus fruticosus and G. physocarpus. Additionally, supplemental hand-pollinations were performed in small and large populations in order to determine whether reproductive output was limited by pollinator activity in these populations. Reproductive Allee effects were not detected in any of the study species. Instead, plants in small populations exhibited remarkably high levels of reproductive output compared to those in large populations. Increased fruit production following supplemental hand-pollinations suggested that the lower reproductive output of naturally pollinated plants in large populations is a consequence of pollen limitation rather than limitation due to abiotic resources. This is consistent with increased intraspecific competition for pollination amongst plants in large populations. It is likely that the invasion of these milkweed species in Australia has been enhanced because plants in small founding populations experience less intraspecific competition for pollinators than those in large populations, and thus have the ability to produce copious amounts of seeds.     

Evidence of a component Allee effect driven by predispersal seed predation in a plant (Pedicularis rex,Orobanchaceae)

A small or sparse population may suffer a reduction in fitness owing to Allee effects. Here, we explored effects of plant density on pollination, reproduction and predation in the alpine herb Pedicularis rex over two years. We did not detect a significant difference in the pollination rate or fecundity (fruit set and the initial seed set) before predation between sparse and dense patches in either year, indicating no pollination-driven Allee effect. However, dense patches experienced significantly fewer attacks by predispersal seed predators in both years, resulting in a significantly decreased realized fecundity (final seed set), suggesting a component Allee effect driven by predispersal seed predation. Predation-driven Allee effects have been predicted by many models and demonstrated for a range of animals, but there is scant evidence for such effects in plants. Our study provides strong evidence of a component Allee effect driven by predation in a plant species.     

Dynamics of a small re-introduced population of wild dogs over 25 years: Allee effects and the implications of sociality for endangered species’ recovery

We analysed 25 years (1980–2004) of demographic data on a small re-introduced population of endangered African wild dogs (Lycaon pictus) in Hluhluwe-iMfolozi Park (HiP), South Africa, to describe population and pack dynamics. As small populations of cooperative breeders may be particularly prone to Allee effects, this extensive data set was used to test the prediction that, if Allee effects occur, aspects of reproductive success, individual survival and population growth should increase with pack and population size. The results suggest that behavioural aspects of wild dogs rather than ecological factors (i.e. competitors, prey and rainfall) primarily have been limiting the HiP wild dog population, particularly a low probability of finding suitable mates upon dispersal at low pack number (i.e. a mate-finding Allee effect). Wild dogs in HiP were not subject to component Allee effects at the pack level, most likely due to low interspecific competition and high prey availability. This suggests that aspects of the environment can mediate the strength of Allee effects. There was also no demographic Allee effect in the HiP wild dog population, as the population growth rate was significantly negatively related to population size, despite no apparent ecological resource limitation. Such negative density dependence at low numbers indicates that behavioural studies of the causal mechanisms potentially generating Allee effects in small populations can provide a key to understanding their dynamics. This study demonstrates how aspects of a species’ social behaviour can influence the vulnerability of small populations to extinction and illustrates the profound implications of sociality for endangered species’ recovery. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Pollinator Abundance and Pollen Limitation of a Solanaceous Shrub at Premontane and Lower Montane Sites

Populations at geographic range limits may frequently encounter changing biotic interactions as well as abiotic constraints. A conundrum in plant population biology, the frequent apparent overproduction of flowers, is thought to be a response by plant populations to unpredictable pollinator environments, such as those that may be expected at the range limit. We studied pollinator visitation rates and pollen limitation of fruit set in populations of Witheringia solanacea , a widespread bee-pollinated species. We compared two populations in the south and two in the north of Costa Rica, with each region encompassing a lower montane site at the upper elevational limit of the species range with small plant populations, and a premontane site where population densities were high. Populations in the south, where the climate was favorable, had higher pollinator visitation and higher natural fruit set, and showed only minor pollen limitation as compared with northern populations at windy sites near the Continental Divide. Lower montane populations had lower pollinator visitation rates than premontane populations but did not show more pollen limitation of fruit set. In general, pollen limitation was not inversely proportional to pollinator abundance but was higher where the climate is unpredictable.     

Pollination of invasive Eichhornia crassipes (Pontederiaceae) by the introduced honeybee (Apis mellifera L.) in South China

Heterostyly functions as an outcrossing mechanism facilitating accurate pollen transfer from anthers to stigmas of particular heights as a result of the behavior of specialist pollinators. However, heterostylous plants are also visited by generalist pollinators, which may affect the plant–pollinator mutualism. Eichhornia crassipes is a tristylous invasive species, with only the mid- and long-styled morphs (M and L) found in China. We recorded flower-visiting insects in Zhuhai, Zhongshan and Nanning in South China. We hand-pollinated the two morphs to determine their compatibility. In addition, by allowing controlled insect pollination in artificial isoplethically monomorphic and bimorphic populations, we undertook a detailed analysis of pollen deposition between the floral morphs, and fruit and seed set. Ranked by relative abundance, the flower-visiting insects were: Apis mellifera, A. cerana, Lasioglossum sp. and Eristalis arvorum. Hand pollination showed that both the M and L morphs were self-compatible, but the former was probably more so than the latter. Intra-morph pollen transfer by A. mellifera within a population was significantly greater than legitimate pollen transfer between populations, suggesting that the pollen exchange between populations was limited. Seed set of the L morph was significantly greater than that of the M morph in monomorphic populations, indicating intra-morph pollen deposition in the former was higher than in the latter. The results showed that A. mellifera was the major pollinator in South China and able to pollinate E. crassipes legitimately and to promote its fruit and seed set, even though high levels of intra-morph pollination occurred.     

Number of conspecifics and reproduction in the invasive plant Eschscholzia californica (Papaveraceae): is there a pollinator‐mediated Allee effect?

The component Allee effect has been defined as ‘a positive relationship between any measure of individual fitness and the number or density of conspecifics’. Larger plant populations or large patches have shown a higher pollinator visitation rate, which may give rise to an Allee effect in reproduction of the plants. We experimentally tested the effect of number of conspecifics on reproduction and pollinator visitation in Eschscholzia californica Cham., an invasive plant in Chile. We then built patches with two, eight and 16 flowering individuals of E. californica (11 replicates per treatment) in an area characterised by dominance of the study species. We found that E. californica exhibits a component Allee effect, as the number of individuals of this species has a positive effect on individual seed set. However, individual fruit production was not affected by the number of plants examined. Pollinator visitation rate was also independent of the number of plants, so this factor would not explain the Allee effect. This rate was positively correlated with the total number of flowers in the patches. We also found that the number of plants did not affect the seed mass or proportion of germinated seeds in the patches. Higher pollen availability in patches with 16 plants and pollination by wind could explain the Allee effect. The component Allee effect identified could lead to a weak demographic Allee effect that might reduce the rate of spread of E. californica. Knowledge of this would be useful for management of this invasive plant in Chile.     

Neutral hybridization can overcome a strong Allee effect by improving pollination quality

Small populations of plant species can be susceptible to demographic Allee effects mainly due to pollen limitation. Although sympatry with a common, co-flowering species may somewhat alleviate the problem of pollinator visitation (pollination quantity), the interspecific pollen transfer, IPT, (pollination quality) may remain a barrier to reproduction in small populations such as new introductions. However, if the two species are crosscompatible, our hypothesis is that neutral hybridization can help the small founding population overcome the Allee effect by improving the quality of pollination. We tested this hypothesis by using a novel modelling approach based on the theory of kinetic reactions wherein pollinators act as enzymes to catalyse the reaction between the two substrates: pollen and unselfed ovule. Using a single locus, two-allele genetic model, we developed a generic model that allows for hybridization between the invading and the native genotypes. Analysing the stability properties of the trivial equilibria in hybridization model as compared with the single genotype invasion model, we found that hybridization can either remove or reduce the Allee effect by making an otherwise stable trivial equilibrium unstable. Our study suggests that hybridization can be neutral but still be the key driver of a successful invasion by mediating pollen limitation. Conservation programmes should therefore account for this cryptic role that hybridization could play in plant invasions.     

Partner limitation and restoration of sexual reproduction in the clonal dwarf shrubLinnaea borealis L. (Caprifoliaceae)

Summary Reports suggest that there is widespread reproductive failure inLinnaea borealis in Britain, Scandinavia, and North America. Our investigations of Scottish populations of this clonal dwarf shrub indicate that, although visited by a number of different insects, pollen transfer in this species is highly effective and principally occurs by small flies (Muscidae). However, natural levels of fruit set varied between populations (from 0% to 25.1%) and reproductive failure was most severe in populations which were composed of single clones. Microscopic examination of stigmas showed that there is no barrier to pollen flow since at least 85% of stigmas have sufficient germinating pollen to effect seed set. Fluorescence microscopy of germinating pollen grains indicates high levels of pollen rejection in the style and only a small proportion of the pollen tubes were able to reach the ovary. It was concluded that lack of xenogamous pollination limits fruit formation in populations ofL. borealis. Reproductive success in an isolated population with extreme reproductive failure was restored by experimental field pollination with viable pollen imported from plants from another population. In Scotland,L. borealis occurs in small, isolated populations and restoration of reproduction can be achieved by the reintroduction of compatible mating partners. This is an important consideration for the conservation management of isolated populations since their long-term recovery may only be possible by translocation of different genotypes from elsewhere into the population. Habitat fragmentation in any part of the species range may impose a potential reproductive bottleneck by causing loss of population diversity and this could explain the low levels of seed set recorded for this species in other parts of its range.     

Reproductive assurance through self-fertilization does not vary with population size in the alien invasive plant Datura stramonium

Autonomous self-fertilization is suggested to be associated with invasiveness in plants because it offers reproductive assurance when there is a shortage of suitable mates or pollinators. Given that shortages of mates and pollinators are a common cause of Allee effects in small plant populations, we predict that the benefits of self-fertilization in terms of reproductive assurance should be greatest in small populations. We tested this idea for the invasive herb Datura stramonium , a self-fertilizing species which is also cross-pollinated to some extent by insects (mainly hawkmoths and honeybees). During two consecutive years, we studied 20 and 55 populations, respectively, of different sizes. Untreated flowers of D. stramonium showed high levels of fruit and seed set in all populations studied. Although, fruit and seed set were generally reduced by about 90% in flowers in which self-fertilization was prevented through emasculation, this effect did not vary according to population size. By using a natural color (anthocyanin) dimorphism in 12 populations, we showed that the average outcrossing rate was low (1.3%) and that there was no relationship between outcrossing rate and population size. Pollen removal from flowers also did not vary according to population size, suggesting that the pollinator visitation rate is not lower in small populations. However, decreasing deviations of observed from expected fruit set with population size imply that small populations may have an increased chance of extinction due to demographic stochasticity. Overall, our results suggest that reproductive assurance through self-fertilization in invasive plants may be important for all stages of population establishment, and not just in the founder population.     

Invasion of rocky shores by a mytilid mussel reveals an abundant-centre distribution coupled with moderate increases in densities at its absolute range limits

Semimytilus patagonicus is an invasive mussel on the coast of southern Africa and has extended its range in recent years. We asked whether its distribution and abundance are consistent with the abundant-centre hypothesis (ACH). Marginal populations were located by monitoring 33 rocky shore sites in South Africa and southern Namibia in 2021. This revealed no changes to its distributional limits since 2020. At nine of these sites, population demography was measured to allow a comparison of their densities and size structure. Four were central populations on the west coast of South Africa (including the site where the species was first detected in 2009). Four were marginal populations in South Africa: two towards the cold range edge in the north and two towards the warm range edge to the south. The ninth population was in southern Namibia, representing a recent invasion event first detected in 2014. Across the species' South African range, the distribution of its abundance was generally consistent with the ACH, with the greatest abundance at its range centre and a gradual decrease towards the range edges. However, the ultimate marginal population at both its cold and warm range edges showed moderate upticks in abundance compared to the penultimate marginal populations. Additionally, marginal populations in South Africa typically included a greater proportion of large individuals. Recruitment intensity was greater in warm range edge populations than cold range edge populations. The size structure of the population in Namibia resembled those of central populations in South Africa. Moderate increases in densities at the absolute range limits suggest that the species is currently undergoing spread into regions associated with moderately optimal environmental conditions (ultimate range edge sites) after encountering regions associated with suboptimal environmental conditions (penultimate range edge sites).     

Female Fecundity Is Dependent on Substrate, Rather Than Male Abundance, in the Wind-Pollinated, Dioecious Understory Palm Chamaedorea radicalis

We examined the impact of substrate, population density, and sexual composition on female reproductive success in harvested populations of the tropical understory palm Chamaedorea radicalis . Leaf harvest for the international floral greenery trade does not kill palms, but has been shown to reduce flowering frequency and has been projected to reduce population growth. Because C. radicalis is dioecious and wind-pollinated, a reduction in flowering density from leaf harvest or other anthropogenic factors may lower pollen movement between flowering conspecifics and lower fruit production. Such reduced fecundity in harvested populations can contribute to an Allee effect, where the per capita rate of population growth declines at low density. We tested for these effects by sampling C. radicalis along transects in ten populations, spanning a range of densities, during May (flowering peak) and August (fruiting peak) 2002. We applied path analysis using structural equation modeling (SEM) to test a set of hypothesized relationships between substrate, female size, several population parameters, and female fecundity. SEM models revealed that female fecundity was not dependent on any measure of population density or sexual composition, providing no evidence for an Allee effect. The strongest model ( R ²= 92.9%) related C. radicalis reproductive output only to substrate, female size, flower number, and proportional fruit set. This model revealed that palms on rock outcrops were larger, produced more flowers, and had higher proportional fruit set than palms on the forest floor. This finding suggests that protection of females on outcrops should enhance population growth and viability.     

Allee effects and population regulation: a test for biotic resistance against an invasive leafroller by resident parasitoids

Resident natural enemies can impact invasive species by causing Allee effects, leading to a reduction in establishment success of small founder populations, or by regulating or merely suppressing the abundance of established populations. Epiphyas postvittana, the Light Brown Apple Moth, an invasive leafroller in California, has been found to be attacked by a large assemblage of resident parasitoids that cause relatively high rates of parasitism. Over a 4-year period, we measured the abundance and per capita growth rates of four E. postvittana populations in California and determined parasitism rates. We found that at two of the sites, parasitism caused a component Allee effect, a reduction in individual survivorship at lower E. postvittana population densities, although it did not translate into a demographic Allee effect, an impact on per capita population growth rates at low densities. Instead, E. postvittana populations at all four sites exhibited strong compensatory density feedback throughout the entire range of densities observed at each site. As we found no evidence for a negative relationship between per capita population growth rates and parasitism rates, we concluded that resident parasitoids were unable to regulate E. postvittana populations in California. Despite a lack of evidence for regulation or a demographic Allee effect, the impact of resident parasitoids on E. postvittana populations is substantial and demonstrates significant biotic resistance against this new invader.     

Allee effects, aggregation, and invasion success

Understanding the factors that influence successful colonization can help inform ecological theory and aid in the management of invasive species. When founder populations are small, individual fitness may be negatively impacted by component Allee effects through positive density dependence (e.g., mate limitation). Reproductive and survival mechanisms that suffer due to a shortage of conspecifics may scale up to be manifest in a decreased per-capita population growth rate (i.e., a demographic Allee effect). Mean-field population level models are limited in representing how component Allee effects scale up to demographic Allee effects when heterogeneous spatial structure influences conspecific availability. Thus, such models may not adequately characterize the probability of establishment. In order to better assess how individual level processes influence population establishment and spread, we developed a spatially explicit individual-based stochastic simulation of a small founder population. We found that increased aggregation can affect individual fitness and subsequently impact population growth; however, relatively slow dispersal—in addition to initial spatial structure—is required for establishment, ultimately creating a tradeoff between probability of initial establishment and rate of subsequent spread. Since this result is sensitive to the scaling up of component Allee effects, details of individual dispersal and interaction kernels are key factors influencing population level processes. Overall, we demonstrate the importance of considering both spatial structure and individual level traits in assessing the consequences of Allee effects in biological invasions.     

Spatial sorting drives morphological variation in the invasive bird, Acridotheris tristis

The speed of range expansion in many invasive species is often accelerating because individuals with stronger dispersal abilities are more likely to be found at the range front. This 'spatial sorting' of strong dispersers will drive the acceleration of range expansion. In this study, we test whether the process of spatial sorting is at work in an invasive bird population (Common myna, Acridotheris tristis) in South Africa. Specifically, we sampled individuals across its invasive range and compared morphometric measurements relevant and non-relevant to the dispersal ability. Besides testing for signals of spatial sorting, we further examined the effect of environmental factors on morphological variations. Our results showed that dispersal-relevant traits are significantly correlated with distance from the range core, with strong sexual dimorphism, indicative of sex-biased dispersal. Morphological variations were significant in wing and head traits of females, suggesting females as the primary dispersing sex. In contrast, traits not related to dispersal such as those associated with foraging showed no signs of spatial sorting but were significantly affected by environmental variables such as the vegetation and the intensity of urbanisation. When taken together, our results support the role of spatial sorting in facilitating the expansion of Common myna in South Africa despite its low propensity to disperse in the native range.     

Contrasting mitochondrial diversity of European starlings (Sturnus vulgaris) across three invasive continental distributions

European starlings (Sturnus vulgaris) represent one of the most widespread and problematic avian invasive species in the world. Understanding their unique population history and current population dynamics can contribute to conservation efforts and clarify evolutionary processes over short timescales. European starlings were introduced to Central Park, New York in 1890, and from a founding group of about 100 birds, they have expanded across North America with a current population of approximately 200 million. There were also multiple introductions in Australia in the mid‐19th century and at least one introduction in South Africa in the late 19th century. Independent introductions on these three continents provide a robust system to investigate invasion genetics. In this study, we compare mitochondrial diversity in European starlings from North America, Australia, and South Africa, and a portion of the native range in the United Kingdom. Of the three invasive ranges, the North American population shows the highest haplotype diversity and evidence of both sudden demographic and spatial expansion. Comparatively, the Australian population shows the lowest haplotype diversity, but also shows evidence for sudden demographic and spatial expansion. South Africa is intermediate to the other invasive populations in genetic diversity but does not show evidence of demographic expansion. In previous studies, population genetic structure was found in Australia, but not in South Africa. Here we find no evidence of population structure in North America. Although all invasive populations share haplotypes with the native range, only one haplotype is shared between invasive populations. This suggests these three invasive populations represent independent subsamples of the native range. The structure of the haplotype network implies that the native‐range sampling does not comprehensively characterize the genetic diversity there. This study represents the most geographically widespread analysis of European starling population genetics to date.     

Early warning signals of demographic regime shifts in invading populations

The Allee effect can cause alternative stable states in population abundance of invasive species. Sudden eruption of invading populations from low to high abundance may be viewed as a regime shift from one alternative state to another. Previous research proposed several types of early warning signals to predict regime shifts in ecological systems such as polluted lakes and semiarid grasslands. This paper explores theoretically the potential of such indicators in predicting demographic regime shifts of invading populations. I analyzed a stochastic differential equation model for the population dynamics of an invasive species subject to Allee effects and propagule pressure. Diffusion approximation to the stochastic model suggests that persistent propagule pressure makes demographic regime shifts inevitable, but Allee effects can lengthen the mean time until regime shifts virtually indefinitely. To compare the potential of indicators, I examined standard deviation, skewness, and estimated return rates of longitudinal population abundance. I found that standard deviation showed a distinct increase as regime shifts became more likely, but skewness and return rates showed no clear trends. This result suggests that standard deviation might be a useful warning signal for forecasting an impending demographic regime shift of invading populations during the period when their abundance is still low.