Testing Darwin's transoceanic dispersal hypothesis for the inland nettle family (Urticaceae)

Dispersal is a fundamental ecological process, yet demonstrating the occurrence and importance of long‐distance dispersal (LDD) remains difficult, having rarely been examined for widespread, non‐coastal plants. To address this issue, we integrated phylogenetic, molecular dating, biogeographical, ecological, seed biology and oceanographic data for the inland Urticaceae. We found that Urticaceae originated in Eurasia c. 69 Ma, followed by ≥ 92 LDD events between landmasses. Under experimental conditions, seeds of many Urticaceae floated for > 220 days, and remained viable after 10 months in seawater, long enough for most detected LDD events, according to oceanographic current modelling. Ecological traits analyses indicated that preferences for disturbed habitats might facilitate LDD. Nearly half of all LDD events involved dioecious taxa, so population establishment in dioecious Urticaceae requires multiple seeds, or occasional selfing. Our work shows that seawater LDD played an important role in shaping the geographical distributions of Urticaceae, providing empirical evidence for Darwin's transoceanic dispersal hypothesis.     

Testing the enemy release hypothesis: a review and meta-analysis

One of the most cited hypotheses explaining the inordinate success of a small proportion of introduced plants that become pests is the ‘natural enemies hypothesis’. This states that invasive introduced plants spread rapidly because they are liberated from their co-evolved natural enemies. This hypothesis had not been properly tested until recently. Previous reviews on this topic have been narrative and vote counting in nature. In this review, we carried out quantitative synthesis and meta-analysis using existing literature on plants and their herbivores to test the different components of the enemy release hypothesis. We found supporting evidence in that (1) insect herbivore fauna richness is significantly greater in the native than introduced ranges, and the reduction is skewed disproportionally towards specialists and insects feeding on reproductive parts; and (2) herbivore damage levels are greater on native plants than on introduced invasive congeners. However, herbivore damage levels are only marginally greater for plants in native than in introduced ranges, probably due to the small numbers of this type of study. Studies quantifying herbivore impacts on plant population dynamics are too scarce to make conclusions for either comparison of plants in native vs introduced ranges or of co-occurring native and introduced congeners. For future research, we advocate that more than two-way comparisons between plants in native and introduced ranges, or native and introduced congeners are needed. In addition, the use of herbivore exclusions to quantify the impacts of herbivory on complete sets of population vital rates of native vs introduced species are highly desirable. Furthermore, three-way comparisons among congeners of native plants, introduced invasive, and introduced non-invasive plants can also shed light on the importance of enemy release. Finally, simultaneously testing the enemy release hypothesis and other competing hypotheses will provide significant insights into the mechanisms governing the undesirable success of invasive species.     

Why close relatives make bad neighbours: phylogenetic conservatism in niche preferences and dispersal disproves Darwin's naturalization hypothesis in the thistle tribe

The number of exotic plant species that have been introduced into the United States far exceeds that of other groups of organisms, and many of these have become invasive. As in many regions of the globe, invasive members of the thistle tribe, Cardueae, are highly problematic in the California Floristic Province, an established biodiversity hotspot. While Darwin's naturalization hypothesis posits that plant invaders closely related to native species would be at a disadvantage, evidence has been found that introduced thistles more closely related to native species are more likely to become invasive. To elucidate the mechanisms behind this pattern, we modelled the ecological niches of thistle species present in the province and compared niche similarity between taxa and their evolutionary relatedness, using fossil‐calibrated molecular phylogenies of the tribe. The predicted niches of invasive species were found to have higher degrees of overlap with native species than noninvasive introduced species do, and pairwise niche distance was significantly correlated with phylogenetic distance, suggesting phylogenetic niche conservatism. Invasive thistles also displayed superior dispersal capabilities compared to noninvasive introduced species, and these capabilities exhibited a phylogenetic signal. By analysing the modelled ecological niches and dispersal capabilities of over a hundred thistle species, we demonstrate that exapted preferences to the invaded environment may explain why close exotic relatives may make bad neighbours in the thistle tribe.     

Changes in the life history of Abbottina rivularis in Lake Fuxian

After invading Lake Fuxian, China, Abbottina rivularis showed marked changes in its life history traits, including alterations in annuli formation, extension to the length of the breeding season, and an increase in fecundity and growth rate. The results are explained in the context of the enemy release hypothesis.     

Testing the facilitation-competition paradigm under the stress-gradient hypothesis: decoupling multiple stress factors

While the facilitation-competition paradigm under the stress-gradient hypothesis has received recent attention, its rigorous testing is yet to be explored. Most of the studies have considered a switch in the net interactions from competition to facilitation with increasing environmental stress as primary evidence supporting the hypothesis, though few studies examined changes in interaction along a full range of a stress gradient. Here, we have conceptualized possible variations in the patterns of change in interaction strength along such gradient. Based on this, we empirically evaluated the temporal shift in the interaction between two marine sessile animals, goose barnacles (Capitulum mitella) and mussels (Septifer virgatus), under multiple stress factors. The net effect of goose barnacles on mussel survivorship was positively related to the total stress gradient encompassing two stress factors, physical disturbance and thermal stress, while no negative value occurred even under mild conditions. When the two stress factors were treated separately, however, the net effect demonstrated apparently different patterns: monotonic increase with physical disturbance versus a quasi-asymptotic pattern (no change over a wide range) with thermal stress. These variable situations have not previously been recognized in this discipline, and the present study emphasizes the importance of an integrative and mechanistic approach to testing and deciphering the facilitation-competition paradigm.     

Realized vs apparent reduction in enemies of the European starling

Release from parasites, pathogens or predators (i.e. enemies) is a widely cited ‘rule of thumb’ to explain the proliferation of nonindigenous species in their introduced regions (i.e. the ‘enemy release hypothesis’, or ERH). Indeed, profound effects of some parasites and predators on host populations are well documented. However, some support for the ERH comes from studies that find a reduction in the species richness of enemies in the introduced range, relative to the native range, of particular hosts. For example, data on helminth parasites of the European starling in both its native Eurasia and in North America support a reduction of parasites in the latter. However, North American ‘founder’ starlings were likely not chosen randomly from across Eurasia. This could result in an overestimation of enemy release since enemies affect their hosts on a population level. We control for the effects of subsampling colonists and find, contrary to previous reports, no evidence that introduced populations of starlings experienced a reduction in the species richness of helminth parasites after colonization of North America. These results highlight the importance of choosing appropriate contrast groups in biogeographical analyses of biological invasions to minimize the confounding effects of ‘propagule biases’.     

Testing multiple pathways for impacts of the non‐native Black‐headed Weaver Ploceus melanocephalus on native birds in Iberia in the early phase of invasion

Not all non‐native species have strong negative impacts on native species. It is desirable to assess whether a non‐native species will have a negative impact at an early stage in the invasion process, when management options such as eradication are still available. Although it may be difficult to detect early impacts of non‐native species, it is necessary to ensure that management decisions can be based on case‐specific scientific evidence. We assess the impacts of a non‐native bird, the Black‐headed Weaver Ploceus melanocephalus, at an early stage in its invasion of the Iberian Peninsula. To do this we identify potential pathways by which competition for shared resources by Black‐headed Weavers could lead to population declines in two ecologically similar native species, and generate hypotheses to test for evidence of competition along these pathways. Black‐headed Weavers could potentially impact native species by displacing them from nesting habitat, or by reducing habitat quality. We found no evidence for either potential competition pathway, suggesting that Black‐headed Weavers do not currently compete with the two native species. However, it is possible that mechanisms that currently allow coexistence may not operate once Black‐headed Weavers reach higher population densities or different habitats.     

Diets of sympatric native and introduced carnivores in the Barrington Tops,eastern Australia

Invasive predators have severe impacts on global biodiversity, and their effects in Australia have been more extreme than on any other continent. The spotted‐tailed quoll (Dasyurus maculatus), an endangered marsupial carnivore, coexists with three eutherian carnivores, the red fox (Vulpes vulpes), feral cat (Felis catus) and wild dog (Canis lupus ssp.) with which it did not coevolve. No previous study has investigated dietary overlap between quolls and the suite of three eutherian carnivores. By analysing scats, we aimed to quantify dietary overlap within this carnivore assemblage in eastern Australia, and to detect any differences that may facilitate coexistence. We also sought evidence of intraguild predation. Dietary overlap between predators was extensive, with the greatest similarity occurring between foxes and cats. However, some differences were apparent. For example, cats mainly consumed smaller prey, and wild dogs larger prey. Quolls showed greater dietary overlap with foxes and cats than with dogs. Intraguild predation was evident, with fox remains occurring in 3% of wild dog scats. Our results suggest wild dogs competitively dominate invasive foxes, which in turn are likely to compete with the endangered quoll.     

Testing the intermediate disturbance hypothesis on species diversity in herbaceous plant communities along a human trampling gradient using a 4-year experiment in an old-field

The intermediate disturbance hypothesis predicts unimodal relationships between species diversity and disturbance frequency/intensity. To test this hypothesis, species diversity in herbaceous plant communities along a human trampling gradient was investigated by conducting a 4-year experiment in an old-field. In general, species richness (S), the Shannon–Weiner index (H) from plant cover data and species evenness (J) showed negative linear relationships with trampling frequency, in contrast to the prediction of the intermediate disturbance hypothesis. However, the significant relationships between trampling frequency and species diversity were not observed in the fourth year without J, which showed a unimodal relationship. In all experimental years, the number of new species that colonized the plots after 1year was small under frequent trampling, and the number of species lost from the plots was large under infrequent trampling. The relative number and the relative cover of perennial species increased as trampling frequency increased in the first and second years, but this pattern was not observed in the following years because the dominance of perennials further increased at decreasing frequencies of trampling. The similarity in the species composition and the yearly changes in species dominance indicated that trampling at higher frequencies eliminated more trampling-intolerant species only in the early years of the experiment. These results suggest that trampling mediated early changes in species diversity patterns, but competitive interactions were more important in the later experimental years. The time lag in the effects of trampling and competition appears to be attributable to the infrequent occurrence of unimodal patterns of species diversity.     

Testing the hypothesis of greater eurythermality in invasive than in native ladybird species: from physiological performance to life‐history strategies

1. Global warming and biological invasions are important threats to biodiversity. Nonetheless, there is little information on how these factors influence performance or life‐history traits of invasive and native species. 2. The effects of temperature on physiological and fitness traits of two invasive alien species (Harmonia axyridis and Hippodamia variegata) and one native species (Eriopis chilensis) of coccinellid were evaluated, testing a model of eurythermality. Eggs of all species were exposed to four temperature treatments (20, 24, 30 and 33 °C). In adult F₂ we measured fecundity, locomotor performance, development time (total and per life stage), survival, and preferred body temperature in a thermal gradient. 3. It was found that H. axyridis had comparatively better performance at low temperatures (i.e. 20 °C), while the performance of H. variegata and E. chilensis did not change with temperature or was better at higher temperatures (30 °C). The standardised Levins index showed that all species are eurythermic. E. chilensis had a high niche overlap with the invasive alien ladybird species, rejecting the hypothesis of greater eurythermality of invasive species than native species. 4. Although there were differences in the temperature preferences and in the response of some physiological and life‐history traits of ladybirds to temperature, both the native and invasive alien species are eurythermic, contrary to the prediction. The better performance of H. axyridis at lower temperatures may result in displacement of its current distribution, and thus not all invasive species will respond favourably to global warming.     

Testing the resource‐matching hypothesis in the mast seeding tree Nothofagus truncata (Fagaceae)

Abstract: The genus Nothofagus in New Zealand and Australia exhibits strong mast seeding (i.e. highly variable seed crops between years). Seed crop variation is synchronized within and between species over large spatial scales, and results in greatly increased wind pollination efficiency which could provide a selective benefit favouring the maintenance of mast seeding. However, the null hypothesis (that plants simply match their reproductive effort to the variable resources available each year) has not been tested in Nothofagus. Here we use a 33‐year dataset on seedfall and wood ring increments for 19 individual Nothofagus truncata trees at Orongorongo, New Zealand, to test for the presence of switching (exaggeration of seedfall variability by diverting resources into, then out of, reproduction). A generalized least squares model explained 40.7% of the variance in standardized ring widths, using six weather variables (absolute minimum temperatures in March (lag 0) April (lag 0 and lag 1), May (lag 0) and rainfall in November and February (lag 0) ) and seedfall. Seedfall had a negative relationship with the current year's ring widths even after controlling for all significant weather variables. This shows that switching is occurring in N. truncata within individuals among years, and therefore that masting in this species is the result of selective forces such as increased wind pollination efficiency. As this result has been demonstrated for very few masting species, we call for this test to be applied more widely.     

Stress relief may promote the evolution of greater phenotypic plasticity in exotic invasive species: a hypothesis

Invasion ecologists have often found that exotic invaders evolve to be more plastic than conspecific populations from their native range. However, an open question is why some exotic invaders can even evolve to be more plastic given that there may be costs to being plastic. Investigation into the benefits and costs of plasticity suggests that stress may constrain the expression of plasticity (thereby reducing the benefits of plasticity) and exacerbate the costs of plasticity (although this possibility might not be generally applicable). Therefore, evolution of adaptive plasticity is more likely to be constrained in stressful environments. Upon introduction to a new range, exotic species may experience more favorable growth conditions (e.g., because of release from natural enemies). Therefore, we hypothesize that any factors mitigating stress in the introduced range may promote exotic invaders to evolve increased adaptive plasticity by reducing the costs and increasing the benefits of plasticity. Empirical evidence is largely consistent with this hypothesis. This hypothesis contributes to our understanding of why invasive species are often found to be more competitive in a subset of environments. Tests of this hypothesis may not only help us understand what caused increased plasticity in some exotic invaders, but could also tell us if costs (unless very small) are more likely to inhibit the evolution of adaptive plasticity in stressful environments in general.     

Exploring the potential of using priority effects during ecological restoration to resist biological invasions in the neotropics

Manipulating plant order of arrival, a process that creates priority effects, may be an unexplored powerful tool to hinder the establishment of invasive non‐native plants in sites under restoration. Knowledge and experimental studies on priority effects in the neotropics are scarce. Here, we propose a research agenda that investigates whether manipulating plant order of arrival can create priority effects in the neotropics, and if this strategy can be used to avoid the spread of undesired species in restoration projects. We also describe our view on expanding existing knowledge on priority effects to the neotropics and identifying key questions for future research.     

Biotic resistance in marine environments

Biological invasions depend in part on the resistance of native communities. Meta‐analyses of terrestrial experiments demonstrate that native primary producers and herbivores generally resist invasions of primary producers, and that resistance through competition strengthens with native producer diversity. To test the generality of these findings, we conducted a meta‐analysis of marine experiments. We found that native marine producers generally failed to resist producer invasions through competition unless the native community was diverse, and this diversity effect was weaker in marine than in terrestrial systems. In contrast, native consumers equally resisted invasive producers in both ecosystems. Most marine experiments, however, tested invasive consumers and these invasions were resisted more strongly than were producer invasions. Given these differences between ecosystems and between marine trophic levels, we used a model‐selection approach to assess if factors other than the resistance mechanism (i.e. competition vs. consumption) are more important for predicting marine biotic resistance. These results suggest that understanding marine biotic resistance depends on latitude, habitat and invader taxon, in addition to distinguishing between competition with and consumption by native species. By examining biotic resistance within and across ecosystems, our work provides a more complete understanding of the factors that underlie biological invasions.     

Testing a new version of the size-advantage hypothesis for sex change: sperm competition and size-skew effects in the bucktooth parrotfish, Sparisoma radians

A variety of field studies suggest that sex change in animalsmay be more complicated than originally depicted by the size-advantagehypothesis. A modification of the size-advantage hypothesis,the expected reproductive success threshold model, proposesthat sperm competition and size-fecundity skew can stronglyaffect reproductive pay-offs. Size-fecundity skew occurs ifa large female's fecundity is markedly higher than the aggregateof the other members of her social group and, together withpaternity dilution from sperm competition, can produce situationsin which large females benefit by deferring sex change to smallerfemales. Deferral by large females can create sex-size distributionscharacterized by the presence of large females and small sex-changedmales, and it is precisely these distributions that the traditionalsize-advantage model cannot explain. We tested the predictionsof the new model with the bucktooth parrotfish, Sparisoma radians,on coral reefs in St. Croix, U.S. Virgin Islands. Collectionsand spawning observations determined that the local environmentalregime of S. radians is characterized by pervasive sperm competition(accompanying 30% of spawns) and factors that can produce substantialsize-fecundity skew in social groups. Dominant male removalexperiments demonstrate that the largest females in social groupsoften do not change sex when provided an opportunity. Instead,smaller, lower-ranking females change sex when a harem vacancyarises. This pattern of sex change is in contrast to virtuallyall previous studies of social control of sex change in fishes,but provides strong support for the general predictions of theexpected reproductive success threshold model.