Enemy Release Research Articles

Invasiveness, ecological impacts and control of acacias in southwestern Europe – a review

Abstract. The most prolific acacias in southern Europe (Acacia dealbata, A. melanoxylon and A. longifolia) are rapidly spreading in its westernmost area: Portugal and NW Spain, where congeners with invasion potential are already established. We performed a bibliographic search of acacia invasions in southern Europe and used spatial data on acacia distribution and abiotic parameters in Iberia to check the influence of abiotic factors on acacia invasion. According to our results, in Iberia A. dealbata and A. melanoxylon seem limited by high soil pH (pHCaCl2>5.5), frequent frosts (>21 to 40 d yr−1) and low annual precipitation (<500 to 1000 mm); data were inconclusive for A. longifolia, while A. saligna prefers neutral soils in the driest and warmest areas. The percentage of area occupied by A. dealbata and A. melanoxylon increases significantly with the percentage of burnt surface. In the literature, acacias' invasiveness is usually attributed to their high resprouting and seeding capacity and to native exclusion through their allelopathic potential; symbiotic promiscuity with rhizobia; high environmental plasticity; and adaptation to burnt, cleared and resource-poor land. However, it is unknown how acacias became so invasive in western Iberia, where native Fabaceae shrubs with similar ecological traits (and invaders outside their natural range) are abundant. Invasive acacias can modify fire and water regimes, aboveground biodiversity, and topsoil characteristics (microbial communities, pH, organic matter and macronutrients levels); nevertheless, sound comparisons with mature stands of Iberian legumes for these and other soil properties (N fluxes, micronutrients) are lacking. As several acacias outcompete Iberian Fabaceae shrubs partly thanks to enemy release, the introduction of biocontrol agents (as for A. longifolia in Portugal) can be useful for invasion control.

Association between temporal patterns in helminth assemblages and successful range expansion of exotic Mus musculus domesticus in Senegal

Relationships between parasitism and invasion success are increasingly evidenced in the literature. However, the dynamic nature of the major parasite-related processes has been rarely taken into account until now, while the residence time of invaders in colonized regions was shown to be associated to crucial changes in parasite communities. Here, we strive to bridge this gap using a temporal survey of rodent populations along one invasion route of the exotic house mouse Mus musculus domesticus that currently invades North Senegal. In this study, we investigated whether gastrointestinal helminth (GIH) assemblages changed over time in native (Mastomys erythroleucus) and/or invasive (M. m. domesticus) rodent populations sampled at an invasion front, and whether these potential changes may be associated to the invasion success of the exotic mouse. Four years separated two rodent sampling campaigns (2013 and 2016/17) in six localities. Despite being relatively short, the timeframe considered here allowed to evidence significant patterns in rodent communities and their GIH assemblages. At the host community level, we showed that the exotic mouse was now established at all sites, becoming the dominant species in sites where it was not recorded before. At the GIH community level, increased infection of the single shared cestode (Mathevotaenia symmetrica) in both rodent species brought support to the “spill-back” hypothesis. Infection levels of GIH that remained low at the invasion front in invading mice over time also supported the “enemy release” hypothesis. Both hypotheses should deserve further experimental work to demonstrate their role in the invasion success of the house mouse in Senegal.

Lineage and latitudinal variation in Phragmites australis tolerance to herbivory: implications for invasion success

Herbivores play a critical role in plant invasions either by facilitating or inhibiting species establishment and spread. However, relatively few studies with invasive plant species have focused on the role of plant tolerance and how it varies geographically to influence invasion success. We conducted a common garden study using two lineages (native and invasive) of the grass Phragmites australis that are prevalent in North American wetlands. Using 31 populations collected across a broad geographic range, we tested five predictions: 1) the invasive lineage is more tolerant to simulated folivory than the native lineage, 2) tolerance to herbivory decreases with increasing latitude of origin of the populations, 3) estimates of tolerance are correlated with putative tolerance traits and plasticity in those traits, 4) a tradeoff exists between tolerance and resistance to herbivory and 5) tolerance has a fitness cost. Response to folivory varied substantially among populations of P. australis, ranging from intolerance to overcompensation. Our model selection procedure deemed lineage to be an important predictor of tolerance but, contrary to our prediction, the native lineage was 19% more tolerant to folivory than the invasive lineage. Tolerance for both lineages exhibited a u‐shaped relationship with latitude. A tolerance–resistance tradeoff was evident within the invasive but not the native lineage. Also, tolerance was positively correlated with belowground biomass allocation, leaf silica concentrations, specific leaf area and plasticity in stem density, and negatively correlated with the relative growth rate of the population and plasticity in putative resistance traits. Lastly, although we did not detect costs of tolerance, our results highlight that fast growth rates can maintain high fitness in the presence of herbivory. Herbivory and plant defense strategies for P. australis lineages in North America exhibit complex biogeographic patterns that cause substantial heterogeneity in enemy release and biotic resistance and, consequently, invasion success.

Plant–soil feedback contributes to predicting plant invasiveness of 68 alien plant species differing in invasive status

Understanding what species characteristics allow some alien plants to become invasive while others fail is critical to our understanding of community assembly processes. While many characteristics have been shown to predict plant invasiveness, the importance of plant–soil feedback (PSF) in invasions has been difficult to assess since individual studies include only a few species and use disparate methodology. We studied PSF of 68 invasive and non‐invasive alien species in a single two‐phase common garden experiment, and compared the relative importance of PSF, residence time, phylogenetic novelty and plant traits for plant invasiveness. Additionally, we explored relationships between PSF, residence time and phylogenetic novelty. PSF for seedling establishment, but not for biomass, was a significant predictor of invasive status, with invasive species having more positive PSF than non‐invasive species. Its explanatory power was, however, much lower than that of specific leaf area, height and residence time. Phylogenetically novel species experienced less negative PSF than species with native congeners, suggesting they benefit more from enemy release. PSF of non‐invasive species, contrary to that of invasive species, was becoming more negative with increasing residence time. We demonstrated that PSF for seedling establishment plays a role in predicting plant invasiveness and is a better predictor than more commonly studied PSF for plant biomass. Other species traits, such as specific leaf area, however, predict plant invasiveness much better than the PSF.

Parasites in sympatric populations of native and invasive freshwater bivalves

An increasing threat to local, native freshwater mussels (Unionida)—an ecologically important but globally alarmingly declining group—is the invasion by exotic bivalves. The Enemy Release Hypothesis predicts that introduced species should benefit from enemy-mediated competition because they are less likely to be harmed by natural enemies, such as parasites, than their native competitors. We investigated within-site differences in parasitism between sympatric native (tot. five spp.) and invasive (tot. three spp.) bivalves in eight northern European waterbodies, which harboured totally 15 parasite taxa. In paired comparisons using within-site averages, the mean number of parasite species in the native bivalves was 2.3 times higher, and the sum of parasite prevalences 2.4 times higher, than in the invasive bivalves. This may lead to enemy-mediated competitive release of invaders and contribute to the success of invasive freshwater bivalves, in general. However, while the invasive clam Corbicula fluminea was completely free from parasites, parasite parameters of the other invader, Sinanodonta woodiana, were relatively high, indicating that the role of parasites can be invader-specific and urges further research. Understanding the factors affecting success of freshwater bivalve invasions, such as parasitism, can aid invasion control and conservation of local, native (endangered) bivalves.

Elucidating mechanisms of invasion success: effects of parasite removal on growth and survival rates of invasive and native frogs.

Identifying the mechanisms underlying biological invasions can inform the management of invasive species. The enemy release hypothesis (ERH) suggests that invasive species have a competitive advantage in their introduced range because they leave behind many of their predators and parasites from their native range, allowing them to shift resources from defenses to growth, reproduction, and dispersal. Many studies have demonstrated that invasive species have fewer parasites than their native counterparts, but few studies have tested whether the loss of these natural enemies appears to be a primary driver of the invasion process.To test the ERH, we conducted a mark-recapture study in which we used an anthelmintic drug to successfully reduce parasitic worms in invasive Cuban treefrogs (Osteopilus septentrionalis) and native treefrogs (Hyla spp.) at half of 12 wetlands, marking nearly 4,200 frogs. If the ERH is supported, we would expect that treating for parasitic worms would have a greater benefit to native than invasive hosts.Growth and survival rates of invasive and native treefrogs responded similarly to the anthelmintic treatment, suggesting that the Cuban treefrog's release from parasitic worms does not appear to significantly contribute to its invasiveness in established areas. Instead, it appears that the overall faster rates of growth and maturation, higher survival rates, and larger body sizes of Cuban treefrogs that we observed may contribute to their expansion and proliferation.Synthesis and applications. Although Cuban treefrogs have a lower diversity of parasitic worms in their invasive than native range, this does not appear to significantly contribute to their invasion success in areas where they have been established for more than 20 years. This suggests that any manipulation of parasites in invasive or native hosts would not be an effective method of controlling Cuban treefrogs or reducing their impacts. Further research into other hypotheses is needed to explain the Cuban treefrog's success and help guide management actions to reduce their spread and negative impacts. Our study demonstrates that enemy release may not be a primary driver of invasiveness, highlighting the need for more experimental tests of the enemy release hypothesis to examine its generality.

Evolution of defense and herbivory in introduced plants-Testing enemy release using a known source population, herbivore trials, and time since introduction.

The enemy release hypothesis is often cited as a potential explanation for the success of introduced plants; yet, empirical evidence for enemy release is mixed. We aimed to quantify changes in herbivory and defense in introduced plants while controlling for three factors that might have confounded past studies: using a wide native range for comparison with the introduced range, measuring defense traits without determining whether they affect herbivore preferences, and not considering the effect of time since introduction. The first hypothesis we tested was that introduced plants will have evolved lower levels of plant defense compared to their source population. We grew South African (source) and Australian (introduced) beach daisies (Arctotheca populifolia) in a common‐environment glasshouse experiment and measured seven defense traits. Introduced plants had more ash, alkaloids, and leaf hairs than source plants, but were also less tough, with a lower C:N ratio and less phenolics. Overall, we found no difference in defense between source and introduced plants. To determine whether the feeding habits of herbivores align with changes in defense traits, we conducted preference feeding trials using five different herbivore species. Herbivores showed no overall preference for leaves from either group. The second hypothesis we tested was that herbivory on introduced plant species will increase through time after introduction to a new range. We recorded leaf damage on herbarium specimens of seven species introduced to eastern Australia and three native control species. We found no change in the overall level of herbivory experienced by introduced plants since arriving in Australia.ConclusionIn the field of invasion ecology, we need to rethink the paradigm that species introduced to a new range undergo simple decreases in defenses against herbivores. Instead, plants are likely to employ a range of defense traits that evolve in both coordinated and opposing ways in response to a plethora of different biotic and abiotic selective pressures.

Release from below- and aboveground natural enemies contributes to invasion success of a temperate invader

Efforts to concurrently test for enemy release of both above- and belowground enemies for invasive plants in non-native range are limited. Moreover, direct evidence for the causal link between enemy release and performance is rare for invasive plants. Thus, we sought to investigate if above- and/or belowground enemies are involved in the enemy release mechanisms at various life stages of a temperate invader and their consequences on plant community properties. We conducted field surveys of enemy damage and plant performance, and plant-soil feedback experiments in the glasshouses using Ambrosia trifida in both its non-native and native ranges across the seed, seedling, and adult life stages. Field surveys showed that seeds of A. trifida were more severely damaged by aboveground enemies in the native relative to the non-native range, while the difference in leaf damage between ranges was small. Plant-soil feedback experiments showed that release from belowground enemies was also important, as seed germination and plant growth were significantly reduced in soils from the native range compared with soils from the non-native range. Consistent with the above results, A. trifida was larger, produced more seeds, had higher density, and exerted stronger impacts on co-occurring native plants in the non-native relative to the native range. Our results demonstrated that release from both above- and belowground enemies at various life stages contributed to the invasion success of A. trifida in its non-native range and highlighted the importance of considering both above- and belowground enemy release when studying plant invasions.

Checklist of parasites for Ponto‐Caspian gobies (Actinopterygii: Gobiidae) in their native and non‐native ranges

This study provides a comprehensive checklist of parasites reported for native and non-native populations of seven Ponto-Caspian goby species, i.e. Babka gymnotrachelus, Neogobius fluviatilis, N. melanostomus, Ponticola gorlap, P. kessleri, Proterorhinus semilunaris and P. semipellucidus, all of which have recently expanded their natural range, either by transoceanic translocation or along the three main European intracontinental invasion corridors. In some cases, invasion success is facilitated by loss of natural enemies, such as predators and parasites; thus, knowledge of a species natural parasite fauna is essential for assessments of enemy release in the host's non-native range. The study is based on published literature and personal unpublished data related to parasites of Ponto-Caspian gobiids. A total of 249 parasite taxa have been reported during 1931—2019, with higher numbers reported from native (214) than non-native (151) ranges. This checklist indicates apparent reduction in native parasites in non-native ranges in all goby species, although parasite colonisation in non-native ranges differs between fish species and invasion corridors. Interestingly, the cumulative number of species reported is still gradually increasing in both ranges. Acquisition of high number of local parasite species also indicates competence of Ponto-Caspian gobies to a wide range of generalist parasites.

Soil biotic and abiotic conditions negate invasive species performance in native habitat

BackgroundMost studies on plant invasion consider the enemy release hypothesis when analyzing native habitats. However, the lower performance of invasive species in the native habitats can be the result of unfavorable soil conditions in the native habitats. While soil biotic and abiotic factors have a potential to restrict the growth of invasive species in their native habitats, our understanding of belowground environment of invasive species in their native habitats is very limited. In this study, we analyzed soil characteristics associated with an exotic invasive plant, Old World Climbing Fern (Lygodium microphyllum), in its native habitat in Australia and the recipient habitat in South Florida. Rhizosphere soil samples from both habitats were analyzed for soil physical, chemical and biological characteristics.ResultsSoil characteristics in the recipient habitats were significantly different compared to those in the native habitats. Soil samples from the native habitat had low soil pH, and high concentrations of elements such as aluminum and zinc which are phytotoxic in acidic soil environments. Additionally, mycorrhizal fungi spores were more diverse in the recipient habitat in Florida compared to the native habitat in Australia.ConclusionOverall, our results indicate that growth of an invasive plant in its native habitats could be restricted by the toxic effects associated with strong soil acidity. Results from this study indicate that invasive plants not only escape from their natural herbivores but also from toxic soil environment in their native habitats.

Review: classical biological control of invasive stink bugs with egg parasitoids - what does success look like?

Although the enemy release hypothesis forms the theoretical basis for classical (=importation) biological control of invasive pests, its core assumptions are not always examined. This could contribute to unrealistic expectations for some biological control programs. In this paper we examine the assumptions that: (i) enemy release has contributed to the invasive nature of four exotic pentatomids in North America; and (ii) classical biological control with egg parasitoids has been or will be successful in reducing populations of these pests below economically significant levels. First, we review the history of biological control programs against invasive stink bugs to highlight the variable and controversial levels of success of introducing egg parasitoids against stink bugs. Then, we use simple stage-structured matrix models to demonstrate that it may be easy to overestimate the contribution of egg parasitism alone to a reduction in stink bug population growth. Finally, we discuss what realistic expectations might be for success of biological control against invasive stink bugs using egg parasitoids in the context of integrated pest management programs. © 2020 Her Majesty the Queen in Right of Canada Pest Management Science © 2020 Society of Chemical Industry.

Top-down and bottom-up controls on an herbivore on a native and introduced plant in a tropical agricultural landscape.

The recent introduction in a tropical agricultural environment of a weedy open-habitat plant (Solanum myriacanthum) and subsequent host range expansion of a common forest-edge butterfly (Mechanitis menapis) onto that plant provides an opportunity to examine reconfiguration of tritrophic networks in human-impacted landscapes. The objectives of this study were (1) determine if the caterpillars on the exotic host are more or less limited by plant defenses (bottom-up forces) and if they experience enemy release (decrease of top-down pressure) and (2) define how anthropic open pasture habitat influences the herbivore’s tritrophic niche. Field and laboratory monitoring of larval survival and performance on a native (Solanum acerifolium) host plant and the exotic (S. myriacanthum) host plant were conducted in the Mindo Valley, Ecuador. Plant physical defenses were also measured. Results showed that larval mortality was mostly top-down on S. acerifolium, linked to parasitism, but mostly bottom-up on S. myriacanthum, possibly linked to observed increased plant defenses. Thus, in the absence of co-evolved relationships, herbivores on the exotic host experienced little top-down regulation, but stronger bottom-up pressures from plant defenses. These findings provide a rare empirical example of enemy-free space as a mechanism underlying host-range expansion. S. myriacanthum was less colonized in open pastures than in semi-shaded habitats (forest edges, thickets): fewer eggs were found, suggesting limited dispersal of adult butterflies into the harsh open environments, and the survival rate of first instar larvae was lower than on semi-shaded plants, likely linked to the stronger defenses of sun-grown leaves. These findings show how environmental conditions modulate the rewiring of trophic networks in heavily impacted landscapes, and limit a biocontrol by a native herbivore on an invasive plant in open habitats.

Lesser leaf herbivore damage and structural defense and greater nutrient concentrations for invasive alien plants: Evidence from 47 pairs of invasive and non-invasive plants

Empirical evidence of enemy release is still inconsistent for invasive alien plant species, although enemy release is the key assumption for both the enemy release hypothesis (ERH) and the evolution of increased competitive ability hypothesis (EICA). In addition, little effort has been made to test this assumption in terms of defense investment using a multi-species comparative approach. Using a phylogenetically controlled within-study meta-analytical approach, we compared leaf herbivore damage, structural defenses and nutrients between 47 pairs of invasive versus native and/or non-invasive alien plants in China. The invasive relative to the co-occurring native or non-invasive (native and non-invasive alien) plants incurred lesser leaf herbivore damage, had lesser leaf concentrations of cellulose, hemicellulose, lignin and carbon, lesser leaf density and carbon or lignin to nitrogen ratio but greater nutrients, which may facilitate success of the invasive plants. The lesser structural investment did not result in lesser leaf construction costs for the invaders, which may be associated with their greater leaf nitrogen concentration. However, the invasive plants were not significantly different from the non-invasive alien plants in any trait. Our results provide strong evidence for ERH, also are consistent with EICA, and indicate that enemy release may be an important factor in alien plant invasions.

Severe Insect Pest Impacts on New Zealand Pasture: The Plight of an Ecological Outlier.

New Zealand’s intensive pastures, comprised almost entirely introduced Lolium L. and Trifolium L. species, are arguably the most productive grazing-lands in the world. However, these areas are vulnerable to destructive invasive pest species. Of these, three of the most damaging pests are weevils (Coleoptera: Curculionidae) that have relatively recently been controlled by three different introduced parasitoids, all belonging to the genus Microctonus Wesmael (Hymenoptera: Braconidae). Arguably that these introduced parasitoids have been highly effective is probably because they, like many of the exotic pest species, have benefited from enemy release. Parasitism has been so intense that, very unusually, one of the weevils has now evolved resistance to its parthenogenetic parasitoid. This review argues that New Zealand’s high exotic pasture pest burden is attributable to a lack of pasture plant and natural enemy diversity that presents little biotic resistance to invasive species. There is a native natural enemy fauna in New Zealand that has evolved over millions of years of geographical isolation. However, these species remain in their indigenous ecosystems and, therefore, play a minimal role in creating biotic resistance in the country’s exotic ecosystems. For clear ecological reasons relating to the nature of New Zealand pastures, importation biological control can work extremely well. Conversely, conservation biological control is less likely to be effective than elsewhere.

The potential for biological control on cryptic plant invasions

Cryptic invasions can be defined as ‘the occurrence of an invasive species or genotype that was not previously recognised as alien in origin or not distinguished from other aliens’. Such invasions can result in negative impacts on the recipient ecosystems and disturb the evolutionary history of native plant populations. Many cryptic invasions have become so problematic that there is a need to implement control measures. This paper explores the potential for biological control to be implemented as a means of managing cryptic invasions. Firstly, the paper defines the different forms of cryptic invasion, differentiating between interspecific and intraspecific invasions; this hierarchy influences how to detect, study and ultimately implement biological control when cryptic invasions occur. Secondly, unique challenges associated with biological control programmes for cryptic invasions are addressed, including: the need for intraspecific level host specificity in agents, the occurrence of hybridisation between native species/lineages and the target weed, the role of enemy release in cryptic invasions in the presence of closely related native plant species/lineages, and a review of potential stakeholder conflicts of interest and legislation. Biological control of cryptic invasions has been shown to be possible, however the process will be more difficult and complex than controlling traditional targets and will likely take up more time and resources. If these challenges are overcome, then biological control programmes against cryptic invasions should be able to proceed and maintain the same standards as traditional biological control programmes.

Insights from excrement: invasive gastropods shift diet to consume the coffee leaf rust and its mycoparasite

Agroecosystems are almost always by definition composed of novel assemblages of organisms from various parts of the world (Perfecto and Vandermeer 2015). As ecologists, we have little ability to predict a priori how interactions within these novel assemblages will organize themselves and what their impacts will be within and adjacent to agricultural production. While it may be possible to make coarse predictions about well-studied organisms, as with natural enemy release in non-native ranges, it is less often the case that we are able to predict the development of novel interactions which result from host shifts in new ecological contexts (Agosta 2006; Nylin et al. 2018).

Urbanization and translocation disrupt the relationship between host density and parasite abundance

The species interactions that structure natural communities are increasingly disrupted by radical habitat change resulting from the widespread processes of urbanization and species translocations. Although many species are disadvantaged by these changes, others thrive in these new environments, achieving densities that exceed those in natural habitats. Often the same species that benefit from urbanization are successful invaders in introduced habitats, suggesting that similar processes promote these species in both environments. Both processes may especially benefit certain species by modifying their interactions with harmful parasites ('enemy release'). To detect such modifications, we first need to identify the mechanisms underlying host-parasite associations in natural populations, then test whether they are disrupted in cities and introduced habitats. We studied the interaction between the cane toad Rhinella marina, a globally invasive species native to South America, and its Amblyomma ticks. Our field study of 642 cane toads across 46 sites within their native range in French Guiana revealed that 56% of toads carried ticks, and that toads with ticks were in poor body condition relative to uninfected conspecifics. Across natural and disturbed habitats tick prevalence and abundance increased with toad density, but this association was disrupted in the urban environment, where tick abundance remained low even where toad densities were high, and prevalence decreased with density. Reductions in the abundance of ticks in urban habitats may be attributable to pesticides (which are sprayed for mosquito control but are also lethal to ticks), and our literature review shows that tick abundance is generally lower in cane toads from urban habitats across South America. In the invasive range, ticks were either absent (in 1,960 toads from Puerto Rico, Hawai'i, Japan and Australia) or less abundant (in Florida and the Caribbean; literature review). The positive relationship between host density and parasite abundance is thought to be a key mechanism through which parasites regulate host populations; anthropogenic processes that disrupt this relationship may allow populations in urban and introduced habitats to persist at densities that would otherwise lead to severe impacts from parasites.

The hidden faces of a biological invasion: parasite dynamics of invaders and natives

One of the primary drivers of emerging infectious diseases (EIDs) is human intervention via host or parasite translocations. A unique opportunity to study host and parasite dispersal during a bio-invasion currently exists in Ireland due to the introduction of the bank vole (Myodes glareolus) in the 1920s. The continuing range expansion of M. glareolus within Ireland presents a natural large-scale perturbation experiment. This study used the Irish M. glareolus model to conduct a spatiotemporal study analysing the parasite dynamics of native and invasive species throughout their range. Myodes glareolus and native Apodemus sylvaticus were trapped in woodlands across Ireland and surveyed for their helminth parasites. Myodes glareolus in Ireland were found to have lower parasite diversity in comparison to records of M. glareolus from across Europe and A. sylvaticus in Ireland. Increased density of M. glareolus resulted in a dilution effect, with significantly lower levels of parasitism overall in native hosts, where M. glareolus has been established longest. However, three helminth parasite species of A. sylvaticus increased in abundance in the presence of M. glareolus. Furthermore, M. glareolus at the expansion front were less parasitised (lower abundance and prevalence of certain parasites and lower parasite diversity) than M. glareolus from the core population. This “enemy release” is believed to be mediating the continued successful spread of the invader across Ireland. Our results identify two important variables, seasonality and the stage of the invasion, which should not be overlooked when investigating or managing the changing distribution of hosts and their parasites. Studies of bio-invasions and parasite transmission have primarily focused on the invasive host species or the native host species in cases where virulent pathogen spillover is observed. Our results demonstrate how the concurrent study of invasive and native hosts, and the careful identification of their parasite communities, allows the dynamic processes influencing the parasite component and intracommunity to be identified.

Sea urchin grazing preferences on native and non-native macroalgae

Abstract Herbivory plays a major role in shaping community dynamics across freshwater, marine and terrestrial habitats, by controlling patterns of abundance and distribution of primary producers, including seaweeds. In the context of biological invasions, the proliferation of non-native seaweeds has been often attributed to limited grazing by native herbivores on introduced species (“Enemy Release Hypothesis”, ERH). In our study, we aimed to explore the potential of an abundant generalist herbivore (the sea urchin Paracentrotus lividus) to graze on non-native in comparison to native macroalgae species. For this purpose, we used manipulative experiments to assess sea urchin preference on native and non-native seaweed species present in the northwest coast of Portugal. Specifically, we determined the preferences of P. lividus on brown seaweeds i.e. Laminaria ochroleuca, Saccorhiza polyschides and the non-native Undaria pinnatifida, and red seaweeds i.e. Chondrus crispus, Mastocarpus stellatus and the non-native Grateloupia turuturu. For each group of seaweed species, sea urchin preference and biomass consumption were examined. The nutritional (organic carbon and nitrogen) and chemical (phenolic content) features were also analysed, in order to assess their effect on sea urchin feeding. According to the results, P. lividus did not show a specific preference for any of the different seaweeds of each phylum. These results suggest that P. lividus is a generalist herbivore, not exerting a differential grazing pressure on non-native seaweeds when compared to native ones.

Evidence for enemy release in invasive common dace Leuciscus leuciscus in Ireland: a helminth community survey and systematic review.

Invasive species lose parasites in the process of invasion and tend to be less parasitized than conspecifics in the native range and sympatric native species in the invasive range (enemy release). We evaluated enemy release in an invasive freshwater fish in Ireland, common dace Leuciscus leuciscus, using helminth parasite community surveys at the core and front of the invasive range of common dace. Furthermore, we undertook a systematic literature review of helminth infection in common dace across its native range in Great Britain and Europe and invasive range in Ireland. The helminth parasite community survey revealed that invasive common dace were infected with fewer helminth species at the invasion front than at the core. Four helminth taxa - Acanthocephala, Monogenea, Digenea and Nematoda - were present in dace at the invasion core compared to only a single helminth species (Pomphorhynchus tereticollis) at the front. The systematic review revealed that invasive common dace in Ireland hosted fewer species of helminths than common dace in the native range. We report a total of three helminth species in common dace in Ireland compared to 24 in Great Britain and 84 in Continental Europe. Our results support the hypotheses that invasive populations are less parasitized than native populations and that more recently established populations host fewer parasites. However, we demonstrate that invasive species may continue to experience release from parasites long after initial invasion.