Differences in herbivore damage and performance among <Emphasis Type="Italic">Arctium minus</Emphasis> (burdock) genotypes sampled from a geographic gradient: a common garden experiment

Performance of plant species does not necessarily decline as they approach their geographic range limits. One reason for this may be a loss of natural enemies in marginal populations. Such patterns have been found in native species, but also may occur for exotics if they have not already escaped their herbivores in invaded regions. For instance, the Eurasian biennial Arctium minus (common burdock) is attacked by a variety of native and introduced insects in its new North American range. Previously, research has shown that damage by these herbivores strongly decreases towards the northern range limit of this species. This gradient might reflect a genetic cline in resistance to herbivores, or geographic variation in herbivore abundance. To distinguish between these possibilities, herbivore damage to leaves and seeds of A. minus was measured in a common garden experiment with genotypes sampled from 11 populations along a 550 km transect extending from southern Ontario towards burdock’s northern range limit. As well, a freezing tolerance experiment was performed with the important lepidopteran seed predator Metzneria lappella, and palatability experiments were performed with two generalists, the snail Cepaea nemoralis and the moth Trichoplusia ni. Although there were some differences in damage among populations, results indicated that latitudinal differences in herbivore damage are not explained by genotypic differences among populations, but instead are likely to result from the absence of herbivores from colder sites. Escape of A. minus from its usual herbivores may increase performance of marginal populations, and contribute to future spread.     

Use of life tables to predict the impact of introducing exotic parasitoids,against the cabbage seedpod weevil in North America

ABSTRACT

Cabbage seedpod weevil, Ceutorhynchus obstrictus (Coleoptera: Curculionidae), is an invasive alien species that impacts seed production of canola, Brassica napus and B. rapa (Brassicaceae), one of Canada’s major agricultural crops. To determine the need for and potential impact of introduction of one or more natural enemies for biological control it is essential to understand the mortality factors associated with C. obstrictus both in the native range and in the introduced range. Life tables were constructed for Ontario and British Columbia populations to determine the mortality factors affecting C. obstrictus. These were compared with life tables generated in Switzerland, the area of origin of the weevil. Results indicated that introduction of an exotic specialist parasitoid into canola growing regions of Canada would substantially reduce C. obstrictus populations. This study is the first to compare life tables of an invasive pest in the areas of origin and introduction.     

Increased resistance to generalist herbivores in invasive populations of the California poppy (Eschscholzia californica)

Escape from enemies in the native range is often assumed to contribute to the successful invasion of exotic species. Following optimal defence theory, which assumes a trade‐off between herbivore resistance and plant growth, some have predicted that the success of invasive species could be the result of the evolution of lower resistance to herbivores and increased allocation of resources to growth and reproduction. Lack of evidence for ubiquitous costs of producing plant toxins, and the recognition that invasive species may escape specialist, but not generalist enemies, has led to a new prediction: invasive species may escape ecological trade‐offs associated with specialist herbivores, and evolve increased, rather than decreased, production of defensive compounds that are effective at deterring generalist herbivores in the introduced range. We tested the performance of two generalist lepidopteran herbivores, Trichoplusia ni and Orgyia vetusta, when raised on diets of native and invasive populations of the California poppy, Eschscholzia californica. Pupae of T. ni were significantly larger when reared on native populations. Similarly, caterpillars of O. vetusta performed significantly better when raised on native populations, indicating that invasive populations of the California poppy are more resistant to herbivores than native populations. The chance of successful establishment of some non‐indigenous plant species may be increased by retaining resistance to generalist herbivores, and in some cases, invasive species may be able to escape ecological trade‐offs in their new range and evolve, as we observed, even greater resistance to generalist herbivores than native plants.     

A Bayesian approach to conservation genetics of Blanding’s turtle (Emys blandingii) in Ontario,Canada

Blanding’s turtle, Emys blandingii, is a globally endangered species with a range centred on the Great Lakes of North America. Several disjunct populations also occur along the East Coast of North America. Previous studies suggest that the Great Lakes portion of the species’ range exhibits panmixia. However, E. blandingii is restricted to relatively small populations in many areas around the Great Lakes. Therefore, panmixia across large geographic distances in this area is unlikely. Here, we apply Bayesian analyses of population structure to samples collected across southern Ontario (N = 97) to test a null hypothesis of panmixia and assess possible management units (MUs), and to estimate rates of gene flow across the study area. Sampled sites in Ontario represent a minimum of four distinct genetic clusters of E. blandingii, which we recommend should be considered as independent MUs. Preliminary evidence suggests that further structure may be present in less robustly sampled areas, which deserve further consideration. Genetic diversity at sampled sites is comparable to that reported for other freshwater turtles. Our comparison between this study and previous work confirms that genetic diversity in E. blandingii is reduced in disjunct eastern populations compared to populations centred on the Great Lakes. Genetic diversity in E. blandingii is not correlated with latitude, and instead may reflect post-glacial dispersal of this species from multiple Pleistocene glacial refugia.     

Conservation in Hine’s sight: the conservation genetics of the federally endangered Hine’s emerald dragonfly,Somatochlora hineana

Hine’s emerald dragonfly (Somatochlora hineana) is distributed in discrete fen and wet meadow habitats over its range from Ontario, Canada, to Missouri, USA. Habitat destruction in the vicinity of Chicago, IL, and other areas lead to its designation as an US federal endangered species in 1995. Our main goal was to delineate the population genetic structure of the species within the northern recovery unit centered on the Door Peninsula in Wisconsin and the southern recovery unit in the Des Plaines River Valley near Chicago, IL. Sites on the Door Peninsula, WI, are in a matrix of agricultural development and second-growth forest and were used as a best available approximation of a pristine system for the dragonfly. We nondestructively sampled 557 adults and larvae from 16 sites in Illinois, Michigan, and Wisconsin from 2008 through 2011 and used ten microsatellite markers to estimate levels of genetic variability, and genetic structure. Mean allelic richness across all sites and years was 5.03 (±0.64) and expected heterozygosity was 0.52 (±0.032). Northern and southern recovery units as designated in the original recovery plan were genetically distinct. We delineated two genetic populations in the northern unit and three within the southern including two disjunct sites.     

Lack of sex-biased dispersal promotes fine-scale genetic structure in alpine ungulates

Identifying patterns of fine-scale genetic structure in natural populations can advance understanding of critical ecological processes such as dispersal and gene flow across heterogeneous landscapes. Alpine ungulates generally exhibit high levels of genetic structure due to female philopatry and patchy configuration of mountain habitats. We assessed the spatial scale of genetic structure and the amount of gene flow in 301 Dall’s sheep (Ovis dalli dalli) at the landscape level using 15 nuclear microsatellites and 473 base pairs of the mitochondrial (mtDNA) control region. Dall’s sheep exhibited significant genetic structure within contiguous mountain ranges, but mtDNA structure occurred at a broader geographic scale than nuclear DNA within the study area, and mtDNA structure for other North American mountain sheep populations. No evidence of male-mediated gene flow or greater philopatry of females was observed; there was little difference between markers with different modes of inheritance (pairwise nuclear DNA F _ST = 0.004–0.325; mtDNA F _ST = 0.009–0.544), and males were no more likely than females to be recent immigrants. Historical patterns based on mtDNA indicate separate northern and southern lineages and a pattern of expansion following regional glacial retreat. Boundaries of genetic clusters aligned geographically with prominent mountain ranges, icefields, and major river valleys based on Bayesian and hierarchical modeling of microsatellite and mtDNA data. Our results suggest that fine-scale genetic structure in Dall’s sheep is influenced by limited dispersal, and structure may be weaker in populations occurring near ancestral levels of density and distribution in continuous habitats compared to other alpine ungulates that have experienced declines and marked habitat fragmentation.     

Geographic patterns of genetic variation in nuclear and chloroplast genomes of two related oaks (<Emphasis Type="Italic">Quercus aliena</Emphasis> and <Emphasis Type="Italic">Q. serrata</Emphasis>) in Japan: implications for seed and seedling transfer

In this study, we assessed geographic patterns of genetic variations in nuclear and chloroplast genomes of two related native oaks in Japan, Quercus aliena and Q. serrata, in order to facilitate development of genetic guidelines for transfer of planting stocks for each species. A total of 12 populations of Q. aliena and 44 populations of Q. serrata were analyzed in this study. Genotyping of nuclear microsatellites in Q. aliena was done with only nine populations (n = 212) due to limited numbers of individuals in two populations, while all 12 populations (n = 89) were used in sequencing chloroplast DNA (cpDNA). In Q. serrata, 43 populations (n = 1032) were genotyped by nuclear microsatellite markers, while cpDNA of 44 populations (n = 350) was sequenced. As anticipated, geographic patterns detected in the variations of Q. aliena’s nuclear genome and its chloroplast haplotype distribution clearly distinguished northern and southern groups of populations. However, those of Q. serrata were inconsistent. The geographic distribution of its chloroplast haplotypes tends to show the predicted differentiation between northern and southern lineages, but geographic signals in the genetic structure of its nuclear microsatellites are weak. Therefore, treating northern and southern regions of Japan as genetically distinct transferrable zones for planting stocks is highly warranted for Q. aliena. For Q. serrata, the strong NE-SW geographic structure of cpDNA should be considered.     

Herbivory Differentially Affects Plant Fitness in Three Populations of the Perennial Herb Lythrum salicaria along a Latitudinal Gradient

Herbivory can negatively and selectively affect plant fitness by reducing growth, survival and reproductive output, thereby influencing plant population dynamics and evolution. Latitudinal variation in intensity of herbivory is common, but the extent to which it translates into corresponding variation in effects on plant performance is still poorly known. We tested the hypothesis that variation in the fitness-consequences of herbivory mirror differences in intensity of herbivory among three natural populations of the perennial herb Lythrum salicaria along a latitudinal gradient from southern to northernmost Sweden. We documented intensity of herbivory and examined its effect on survival, growth and reproductive output over two years by experimentally removing herbivores with insecticide. The intensity of herbivory and the effects of herbivory on plant fitness were strongest in the southern population, intermediate in the central population and weakest in the northern population. The mean proportion of the leaf area removed ranged from 11% in the southern to 3% in the northern population. Herbivore removal increased plant height 1.5-fold in the southern and 1.2-fold in the central population, the proportion plants flowering 4-fold in the southern and 2-fold in the central population, and seed production per flower 1.6-fold in the southern and 1.2-fold in the central population, but did not affect plant fitness in the northern population. Herbivore removal thus affected the relative fecundity of plants in the three populations: In the control, seed output per plant was 8.6 times higher in the northern population compared to the southern population, whereas after herbivore removal it was 2.5 times higher in the southern population. The results demonstrate that native herbivores may strongly affect the demographic structure of L. salicaria populations and thereby shape geographic patterns of seed production. They further suggest that the strength of herbivore-mediated selection varies among populations and decreases towards the north.     

Parasitoids follow herbivorous insects to a novel host plant,generalist predators less so

The ‘enemy‐free space’ hypothesis predicts that herbivorous insects can escape their natural enemies by switching to a novel host plant, with consequences for the evolution of host plant specialisation. However, if natural enemies follow herbivores to their novel host plants, enemy‐free space may only be temporary. We tested this by studying the colonisation of the introduced tree Eucalyptus grandis (Hill) Maiden (Myrtaceae) by insects in Brazil, where various species of herbivores have added eucalyptus to their host plant range, which consists of native myrtaceous species such as guava. Some herbivores, for example, Thyrinteina leucoceraea Ringe (Lepidoptera: Geometridae), cause outbreaks in eucalyptus plantations but not on guava, possibly because eucalyptus offers enemy‐free space. We sampled herbivores (mainly Lepidoptera species) and natural enemies on eucalyptus and guava and assessed parasitism of Lepidoptera larvae on both host plant species during ca. 2 years. Overall, predators were encountered more frequently on guava than on eucalyptus. In contrast, parasitoids were encountered equally and parasitism rates of Lepidoptera larvae were similar on both host plants. This indicates that herbivores may escape some enemies by moving to a novel host plant. However, this escape may be temporary and may vary with time. We argue that studying temporal and spatial patterns of enemy‐free space and the response of natural enemies to host use changes of their herbivorous prey is essential for understanding the role of natural enemies in the evolution of host plant use by herbivorous arthropods.     

Spatio-temporal variation in seed predation by a native weevil in the invasive Prunus serotina

Invasive species may escape the enemies from their native range (‘enemy release’), but they can also acquire new enemies in their introduced range, which will affect the invasion process. For the invasive tree species Prunus serotina, seed predation by the native weevil Furcipus rectirostris has been reported in forests in its introduced range. In this study, we quantified how common the infestation of P. serotina seeds by F. rectirostris is in a 4000 km² area in northern Belgium. Seeds were sampled on P. serotina trees in different habitats and in two years, i.e., with low and high P. serotina fruit production. Infestation was found throughout the study region, in 43 and 62% of the sampled trees in the two years of the study; the maximum infestation levels of infested seed samples were 50 and 69%. Overall, predation occurred in 4.4 and 10.8% of the sampled seeds. The level of infestation differed between habitats and years, and the number of fruits per raceme was inversely related to the infestation level. Notwithstanding the rather high incidence of F. rectirostris infestation in our study, the impact on P. serotina's invasiveness might remain low seeing the overall high seed production and dispersal capacity of the species.     

Herbivores differentially limit the seedling growth and sapling recruitment of two dominant rain forest trees

Resource heterogeneity may influence how plants are attacked and respond to consumers in multiple ways. Perhaps a better understanding of how this interaction might limit sapling recruitment in tree populations may be achieved by examining species’ functional responses to herbivores on a continuum of resource availability. Here, we experimentally reduced herbivore pressure on newly established seedlings of two dominant masting trees in 40 canopy gaps, across c. 80 ha of tropical rain forest in central Africa (Korup, Cameroon). Mesh cages were built to protect individual seedlings, and their leaf production and changes in height were followed for 22 months. With more light, herbivores increasingly prevented the less shade-tolerant Microberlinia bisulcata from growing as tall as it could and producing more leaves, indicating an undercompensation. The more shade-tolerant Tetraberlinia bifoliolata was much less affected by herbivores, showing instead near to full compensation for leaf numbers, and a negligible to weak impact of herbivores on its height growth. A stage-matrix model that compared control and caged populations lent evidence for a stronger impact of herbivores on the long-term population dynamics of M. bisulcata than T. bifoliolata. Our results suggest that insect herbivores can contribute to the local coexistence of two abundant tree species at Korup by disproportionately suppressing sapling recruitment of the faster-growing dominant via undercompensation across the light gradient created by canopy disturbances. The functional patterns we have documented here are consistent with current theory, and, because gap formations are integral to forest regeneration, they may be more widely applicable in other tropical forest communities. If so, the interaction between life-history and herbivore impact across light gradients may play a substantial role in tree species coexistence.     

Decreased resistance and increased tolerance to native herbivores of the invasive plant Sapium sebiferum

Release from natural enemies may favor invasive plants evolving traits associated with reduced herbivore‐resistance and faster‐growth in introduced ranges. Given a genetic trade‐off between resistance and tolerance, invasive plants could also become more tolerant to herbivory than conspecifics in the native range. We conducted a field common garden study in the native range of Sapium sebiferum using seeds from native Chinese populations and invasive North American populations to compare their growth and herbivory resistance. We also performed a cage‐pot experiment to compare their resistance and tolerance to Bikasha collaris beetles that are specialist feeders on S. sebiferum trees in China. Results of the common garden study showed that Sapium seedlings of invasive populations relative to native populations were more frequently attacked by native herbivores. Growth and leaf damage were significantly higher for invasive populations than for native populations. Growth of invasive populations was not significantly affected by insecticide spray, but insecticide spray benefited that of native populations. In the bioassay trial, beetles preferentially consumed leaf tissue of invasive populations compared to native populations when beetles had a choice between them. Regression of percent leaf damage on biomass showed that invasive populations tolerated herbivory more effectively than native populations. Our results suggest that S. sebiferum from the introduced range had lower resistance but higher tolerance to specialist herbivores. Both defense strategies could have evolved as a response to the escape from natural enemies in the introduced range.     

Release from herbivory does not confer invasion success for Eugenia uniflora in Florida

One of the most commonly cited hypotheses explaining invasion success is the enemy release hypothesis (ERH), which maintains that populations are regulated by coevolved natural enemies where they are native but are relieved of this pressure in the new range. However, the role of resident enemies in plant invasion remains unresolved. We conducted a field experiment to test predictions of the ERH empirically using a system of native, introduced invasive, and introduced non-invasive Eugenia congeners in south Florida. Such experiments are rarely undertaken but are particularly informative in tests of the ERH, as they simultaneously identify factors allowing invasive species to replace natives and traits determining why most introduced species are unsuccessful invaders. We excluded insect herbivores from seedlings of Eugenia congeners where the native and invasive Eugenia co-occur, and compared how herbivore exclusion affected foliar damage, growth, and survival. We found no evidence to support the ERH in this system, instead finding that the invasive E. uniflora sustained significantly more damage than the native and introduced species. Interestingly, E. uniflora performed better than, or as well as, its congeners in terms of growth and survival, in spite of higher damage incidence. Further, although herbivore exclusion positively influenced Eugenia seedling survival, there were few differences among species and no patterns in regard to invasion status or origin. We conclude that the ability of E. uniflora to outperform its native and introduced non-invasive congeners, and not release from insect herbivores, contributes to its success as an invader in Florida.     

The impact of transportation and translocation on dispersal behaviour in the invasive cane toad

Biological invasions transport organisms to novel environments; but how does the translocation process influence movement patterns of the invader? Plausibly, the stress of encountering a novel environment, or of the transport process, might induce rapid dispersal from the release site—potentially enhancing (or reducing) invader success and spread. We investigated the effect of transportation and release to novel environments on dispersal-relevant traits of one of the world’s most notorious invaders, the cane toad (Rhinella marina). We collected toads in northern New South Wales from heath and woodland habitats, manipulated the level of transport stress and either returned toads to their exact collection point (residents) or reciprocally translocated them to a novel site. Both translocation and the level of transport stress drastically altered toad dispersal rates for at least 5 days post-release. Translocated toads (depending on their level of transport stress and release habitat) moved on average two to five times further per day (mean range 67–148 m) than did residents (mean range 22–34 m). Translocated toads also moved on more days, and moved further from their release point than did resident toads, but did not move in straighter lines. A higher level of transport stress (simulating long-distance translocation) had no significant effect on movements of resident toads but amplified the dispersal of translocated toads only when released into woodland habitat. These behavioural shifts induced by translocation and transportation may affect an invader’s ability to colonise novel sites, and need to be incorporated into plans for invader control.     

Differences in growth and herbivory damage of native and invasive kudzu (Peuraria montana var. lobata) populations grown in the native range

The invasion success of exotic plants is often attributed to escape from natural enemies in their introduced ranges and subsequent evolutionary change in resource allocation from defense to growth and reproduction. We tested this idea by comparing resistance, tolerance, and growth between native (China) and invasive (US) populations of kudzu (Peuraria montana var. lobata) exposed to natural herbivores in the native range. The percentage of foliar damage was much higher in invasive populations than in native populations, indicating that plants from invasive populations had lower resistance to herbivory. Regression of total mass on percentage of foliar damage showed no significant differences in tolerance to herbivory between native and invasive populations. However, stem diameter and mass were significantly greater in invasive populations than in native populations. Our results may suggest geographic variation in herbivory damage and plant growth among kudzu native and invasive populations, but the role of herbivores influencing kudzu invasion requires further investigation.     

Continent-wide tracking to determine migratory connectivity and tropical habitat associations of a declining aerial insectivore

North American birds that feed on flying insects are experiencing steep population declines, particularly long-distance migratory populations in the northern breeding range. We determine, for the first time, the level of migratory connectivity across the range of a songbird using direct tracking of individuals, and test whether declining northern populations have higher exposure to agricultural landscapes at their non-breeding grounds in South America. We used light-level geolocators to track purple martins, Progne subis, originating from North American breeding populations, coast-to-coast (n = 95 individuals). We show that breeding populations of the eastern subspecies, P. s. subis, that are separated by ca. 2000 km, nevertheless have almost completely overlapping non-breeding ranges in Brazil. Most (76%) P. s. subis overwintered in northern Brazil near the Amazon River, not in the agricultural landscape of southern Brazil. Individual non-breeding sites had an average of 91 per cent forest and only 4 per cent agricultural ground cover within a 50 km radius, and birds originating from declining northern breeding populations were not more exposed to agricultural landscapes than stable southern breeding populations. Our results show that differences in wintering location and habitat do not explain recent trends in breeding population declines in this species, and instead northern populations may be constrained in their ability to respond to climate change.     

Reduced seed predation after invasion supports enemy release in a broad biogeographical survey

The Enemy Release (ER) hypothesis predicts an increase in the plant invasive capacity after being released from their associated herbivores or pathogens in their area of origin. Despite the large number of studies on biological invasions addressing this hypothesis, tests evaluating changes in herbivory on native and introduced populations and their effects on plant reproductive potential at a biogeographical level are relatively rare. Here, we tested the ER hypothesis on the South African species Senecio pterophorus (Asteraceae), which is native to the Eastern Cape, has expanded into the Western Cape, and was introduced into Australia (>70–100 years ago) and Europe (>30 years ago). Insect seed predation was evaluated to determine whether plants in the introduced areas were released from herbivores compared to plants from the native range. In South Africa, 25 % of the seedheads of sampled plants were damaged. Plants from the introduced populations suffered lower seed predation compared to those from the native populations, as expected under the ER hypothesis, and this release was more pronounced in the region with the most recent introduction (Europe 0.2 % vs. Australia 15 %). The insect communities feeding on S. pterophorus in Australia and Europe differed from those found in South Africa, suggesting that the plants were released from their associated fauna after invasion and later established new associations with local herbivore communities in the novel habitats. Our study is the first to provide strong evidence of enemy release in a biogeographical survey across the entire known distribution of a species.     

SEED MORTALITY IN DAUCUS CAROTA POPULATIONS: LATITUDINAL EFFECTS

Daucus carota, a common herbaceous weed, grows over a wide latitudinal range in eastern North America. Viability and germination tests of mature seeds collected from 36° to 45°N were conducted to measure predispersal seed mortality. Viability and germination declined as latitude of the seed source decreased. Only 30–50% of the seeds from southern populations germinated owing to high embryo inviability and absence of embryos. Sixty to ninety percent of the seeds from northern populations germinated. Reciprocal planting of seeds in outdoor experimental plots at three latitudes and testing of seeds over two generations together showed that the environment in which seeds mature, rather than environmental preconditioning over generations or genetically-based differences among populations, explain this variation in germination ability. Within-latitude germination declined in experimental plots as population age of the seed source within latitudes increased. The data indicate that predispersal seed mortality can influence local population persistence and that seed mortality is an increasingly important factor in population regulation at the southern limit of the species’ range.     

When do herbivores affect plant invasion? Evidence for the natural enemies and biotic resistance hypotheses

Two venerable hypotheses, widely cited as explanations for either the success or failure of introduced species in recipient communities, are the natural enemies hypothesis and the biotic resistance hypothesis. The natural enemies hypothesis posits that introduced organisms spread rapidly because they are liberated from their co‐evolved predators, pathogens and herbivores. The biotic resistance hypothesis asserts that introduced species often fail to invade communities because strong biotic interactions with native species hinder their establishment and spread. We reviewed the evidence for both of these hypotheses as they relate to the importance of non‐domesticated herbivores in affecting the success or failure of plant invasion.
To evaluate the natural enemies hypothesis, one must determine how commonly native herbivores have population‐level impacts on native plants. If native herbivores seldom limit native plant abundance, then there is little reason to think that introduced plants benefit from escape from these enemies. Studies of native herbivore‐native plant interactions reveal that plant life‐history greatly mediates the strength with which specialist herbivores suppress plant abundance. Relatively short‐lived plants that rely on current seed production for regeneration are most vulnerable to herbivory that reduces seed production. As such, these plants may gain the greatest advantage from escaping their specialist enemies in recipient communities. In contrast, native plants that are long lived or that possess long‐lived seedbanks may not be kept “in check” by native herbivores. For these species, escape from native enemies may have little to do with their success as exotics; they are abundant both where they are native and introduced.
Evidence for native herbivores providing biotic resistance to invasion by exotics is conflicting. Our review reveals that: 1) introduced plants can attract a diverse assemblage of native herbivores and that 2) native herbivores can reduce introduced plant growth, seed set and survival. However, the generality of these impacts is unclear, and evidence that herbivory actually limits or reduces introduced plant spread is scarce. The degree to which native herbivores provide biotic resistance to either exotic plant establishment or spread may be greatly determined by their functional and numerical responses to exotic plants, which we know little about. Generalist herbivores, through their direct effects on seed dispersal and their indirect effects in altering the outcome of native–non‐native plant competitive interactions, may have more of a facilitative than negative effect on exotic plant abundance.     

Comparison of phototrophic shell-degrading endoliths in invasive and native populations of the intertidal mussel Mytilus galloprovincialis

The intertidal mussel Mytilus galloprovincialis is a successful invader worldwide. Since its accidental introduction onto the South African west coast in the late 1970s, it has become the most successful marine invasive species in South Africa. One possible explanation for this phenomenon is that M. galloprovincialis suffers less from phototrophic shell-degrading endoliths in its invasive than in its native range. We assessed photoautotrophic endolithic pressure on M. galloprovincialis in native (Portugal) and invasive (South Africa) ranges. Invasive populations were more heavily infested than native populations. In Portugal, only the biggest/oldest mussels displayed endolithic erosion of the shell and the incidence of infestation was greater at higher shore levels where more prolonged exposure to light enhances endolith photosynthesis. In South Africa, even the smallest size classes of mussels were heavily infested throughout the shore. In Portugal, endolithic-induced mortality was observed at only one location, while in South Africa it occurred at all locations and at significantly higher rates than in Portugal. Important sub-lethal effects were detected in infested native mussels, confirming previous studies of invasive populations and suggesting an energy trade-off between shell repair and other physiological constraints. We observed a positive relationship between infestation rates and barnacle colonization on mussel shells, suggesting possible facilitation of barnacle settlement/survival by shell-boring pathogens. Identification of endoliths revealed common species between regions. However, two species were unique in the invasive range while another was unique in the native region. Different levels of endolithic infestation in the invasive and the native range were not explained by the effect of major environmental determinants (Photosynthetically Available Radiation and wave height). The results reject our initial hypothesis, indicating that invasion success of M. galloprovincialis is not simply explained by escape from its natural enemies but results from complex interactions between characteristics of the invaded community and properties of the invader.