Emergent impacts of cannibalism and size refuges in prey on intraguild predation systems

Many organisms undergo ontogenetic niche shifts due to considerable changes in size during their development. These ontogenetic shifts can alter the trophic position of individuals, the type and strength of ecological interactions across species, and allow for cannibalism within species. In this study we ask if and how the interaction of a size refuge and cannibalism in the prey alters the dynamics of intraguild predation (IGP) systems. By manipulating the composition of large cannibalistic (Aeshna umbrosa) and predatory (Anax junius) dragonfly larvae in mesocosms we show that the interaction of cannibals and predators was non-linear and increased the survival of prey. The structure of the final resource community shared by prey and predator differed between small and large dragonfly treatments but not within size classes across species. In general, the small prey stage showed similar shifts in microhabitat use and refuge use when exposed to either conspecific cannibals or predators, while large cannibals showed no clear anti-predator response. However, further behavioral experiments revealed that specific behavioral components, such as distances between individuals or number of movements, differed when individuals were exposed to either cannibals or predators. This indicates that individuals discriminated between conspecific or heterospecific predators. Furthermore, in similar experiments large cannibals and predators showed different behaviors when exposed to conspecifics rather than to each other. These changes in behavior are consistent with the observed increase in prey survival. In general, the results indicate that cannibalism and ontogenetic niche shifts can result in behavior-mediated indirect interactions that reduce the impact of the predator on the mortality of its prey and alter the interactions of IGP systems. However, they also indicate that size is not the sole determinant and that we also need to account for the species identity when predicting the dynamics of communities.     

Niche segregation among sympatric Amazonian teiid lizards

We examined standard niche axes (time, place, and food) for three sympatric teiid lizards in the Amazon rain forest. Activity times during the day were similar among species. Ameiva ameiva were in more open microhabitats and had higher body temperatures compared with the two species of Kentropyx. Microhabitat overlaps were low and not significantly different from simulations based on Monte Carlo analysis. Grasshoppers, crickets, and spiders were important in the diets of all three species and many relatively abundant prey were infrequently eaten (e.g., ants). Dietary overlaps were most similar between the two species of Kentropyx even though microhabitat overlaps were relatively low. A Monte Carlo analysis on prey types revealed that dietary overlaps were higher at all ranks than simulated overlaps indicating that use of prey is not random. Although prey size was correlated with lizard body size, there were no species differences in adjusted prey size. A. ameiva ate more prey items at a given body size than either species of Kentropyx. Body size varies among species, with A. ameiva being the largest and K. altamazonica the smallest. The two species of Kentropyx are most distant morphologically, with A. ameiva intermediate. The most distant species morphologically are the most similar in terms of prey types. A morphological analysis including 15 species from four genera revealed patterns of covariation that reflected phylogenetic affinities (i.e., taxonomic patterns are evident). A cluster analysis revealed that A. ameiva, K. pelviceps, and K. altamazonica were in the same morphological group and that within that group, A. ameiva differed from the rest of the species. In addition, K. pelviceps and K. altamazonica were distinguishable from other species of Kentropyx based on morphology. Received: 26 December 1998 / Accepted: 15 September 1999     

Edge effects and intraguild predation in native and introduced centipedes: evidence from the field and from laboratory microcosms

Human alteration of habitat has increased the proportion of forest edge in areas of previously continuous forest. This edge habitat facilitates invasion of exotic species into remaining fragments. The ability of native species to resist invasion varies and may depend on intrinsic variables such as dispersal and reproductive rates as well as external factors such as rate of habitat change and the density of populations of introduced species in edge habitat. We examined the distributional and competitive relationships of two members of the class Chilopoda, Scolopocryptops sexspinosus, a centipede native to the eastern US, and Lithobius forficatus, an exotic centipede introduced from Europe. We found that L. forficatus was most abundant in edge habitat and S. sexspinosus was most abundant in the interior habitat at our field sites. Although L. forficatus was present in habitat interiors at 11 of 12 sites, there was no correlation between fragment size and numbers of L. forficatus in interior habitat. The native centipede was rarely found occupying fragment edges. We used laboratory microcosms to examine potential competitive interactions and to indirectly assess prey preferences of the two species. In microcosms both species consumed similar prey, but the native centipede, S. sexspinosus, acted as an intraguild predator on the introduced centipede. Native centipedes were competitively superior in both intraspecific and interspecific pairings. Our results suggest that intraguild predation may aid native centipedes in resisting invasion of introduced centipedes from edge habitat.     

Intraguild predation and successful invasion by introduced ladybird beetles

Introductions of two ladybird beetle (Coleoptera: Coccinellidae) species, Coccinella septempunctata and Harmonia axyridis, into North America for aphid biocontrol have been followed by declines in native species. We examined intraguild predation (IGP) between larvae of these two exotic species and larvae of the two most abundant native coccinellids in eastern Washington State, C. transversoguttata and Hippodamia convergens. In pairings between the two native species in laboratory microcosms containing pea (Pisum sativum) plants, neither native had a clear advantage over the other in IGP. When the natives were paired with either Harmonia axyridis or C. septempunctata, the natives were more frequently the victims than perpetrators of IGP. In contrast, in pairings between the exotic species, neither had an IGP advantage, although overall rates of IGP between these two species were very high. Adding alternative prey (aphids) to microcosms did not alter the frequency and patterns of relative IGP among the coccinellid species. In observations of encounters between larvae, the introduced H. axyridis frequently survived multiple encounters with the native C. transversoguttata, whereas the native rarely survived a single encounter with H. axyridis. Our results suggest that larvae of the native species face increased IGP following invasion by C. septempunctata and H. axyridis, which may be contributing to the speed with which these exotic ladybird beetles displace the natives following invasion.     

The role of antipredator behavior in an experimental community of jumping spiders with intraguild predation

 The role of direct and indirect interactions in intraguild predation (IGP) was investigated in a laboratory study. The study system contained two spider species, Phidippus audax and Phidippus octopunctatus, and the fruit fly, Drosophila melanogaster. P. audax and P. octopunctatus eat D. melanogaster. P. audax (top predators) also eat P. octopunctatus (intermediate predators). Thus, P. audax and P. octopunctatus compete for the shared resource and also interact as predator and prey. Experiments consisted of two treatments: risk-IGP and full-IGP. In the risk-IGP treatments, I examined the effects of trait-mediated indirect effects generated by antipredator behavior of P. octopunctatus on the survival of fruit flies. P. audax chelicerae were waxed so that P. audax could not attack a prey. The result indicated a significant positive indirect effect of P. audax on the survival of D. melanogaster due to the antipredator behavior of P. octopunctatus (a trait-mediated indirect effect). In the full-IGP treatments, P. audax chelicerae were not restricted, so that it could attack prey; this resulted in decreased survival of D. melanogaster. Because of predation of P. audax on P. octopunctatus, even stronger positive interactions occurred between P. audax and D. melanogaster in full-IGP than in risk-IGP. Received: May 7, 2002 / Accepted: June 27, 2002     

The usefulness of amplified fragment length polymorphism markers for taxon discrimination across graduated fine evolutionary levels in Caribbean Anolis lizards

Fine-level taxon discrimination is important in biodiversity assessment and ecogeographical research. Genomic markers are often required for studies on closely related taxa, however, most existing mitochondrial and nuclear markers require prior knowledge of the genome and are impractical for use in small conservation projects. This study describes the application of amplified fragment length polymorphism (AFLP) to discriminate at four progressively finer evolutionary levels of Caribbean Anolis lizards from the central Lesser Antilles. AFLP is shown to be a rapid and effective method for discriminating between species. Separation increases with primer pair number and choice of primer combination appears to be noncritical. Initial population-level results show markedly less discriminatory power. A screening technique for the identification of population informative markers combining principal component and principal coordinate analyses is presented and assessed. Subsequent results show selected conspecific AFLP data to be remarkably congruent with those of mitochondrial DNA, microsatellite and morphological markers. The use of AFLP as a low-cost nuclear marker in species-level taxon discrimination is supported, whereas population level application demands further consideration.     

On periodic cohort solutions of a size-structured population model

 We consider a size-structured population model with discontinuous reproduction and feedback through the environmental variable ‘substrate’. The model admits solutions with finitely many cohorts and in that case the problem is described by a system of ODEs involving a bifurcation parameter β. Existence of nontrivial periodic n-cohort solutions is investigated. Moreover, we discuss the question whether n cohorts (n≧2) with small size differences will tend to a periodic one-cohort solution as t→∞. Received 16 March 1995; received in revised form 7 January 1997     

Habitat use and ecological interactions of an introduced and a native species of Anolis lizard on Grand Cayman,with a review of the outcomes of anole introductions

Since its introduction ten years ago, Anolis sagrei has spread over much of Grand Cayman and is now more common in some habitats than the native anole, A. conspersus. Interspecific differences in body size, perch height, and microclimatic preference may have facilitated the colonization. Nonetheless, competition may be occurring between the species; comparisons with studies of habitat use prior to the arrival of A. sagrei indicate that in open habitats, where A. sagrei is now abundant, A. conspersus perches higher, but in closed habitats, where A. sagrei is absent, no change in perch height is evident. Review of data concerning 23 Anolis introductions indicates that the presence or absence of an ecologically similar native species may be an important determinant of colonization success or failure.     

Mutual predation between and cannibalism within several freshwater gammarids: Dikerogammarus villosus versus one native and three invasives

Dikerogammarus villosus, a Ponto-Caspian species, is known to be a particularly successful invader, and is currently the prevailing invasive gammarid in Southern German large waters. Field observations suggest that D. villosus has replaced the native G. pulex and the invasive D. haemobaphes, also a Ponto-Caspian species, in some reaches of the German Danube. Dikerogammarus villosus is also believed to be the reason why Echinogammarus berilloni, a Mediterranean species, and Dikerogammarus bispinosus, a further Ponto-Caspian gammaridean species, could not build significant populations in Southern German rivers. Because intraguild predation (IGP) is regarded as a major force leading to species displacements, we hypothesized that superiority in predation by D. villosus is responsible for the disappearance or limited occurrence of several native and invasive gammarideans in many reaches of Southern German waters. To test this hypothesis, we conducted a series of laboratory IGP experiments with 1:1 combinations in which D. villosus was allowed to prey on other gammarids and vice versa. We also studied the extent of cannibalism within species. Dikerogammarus villosus was a stronger predator than G. pulex and E. berilloni. However, D. villosus was not stronger in predation than its relatives D. haemobaphes and D. bispinosus, although results with the combinations of D. villosus and D. bispinosus were less conclusive, especially in the female–female combination. Cannibalism rate was high in D. haemobaphes compared with other species. We conclude that superiority in IGP experiments could explain the disappearance of G. pulex and the missing or limited occurrence of E. berilloni since the arrival of D. villosus in Southern German rivers. However, the results of IGP experiments do not explain why D. haemobaphes was displaced by D. villosus in many places and why D. bispinosus could not build significant populations in Southern German waters. Possibly specific combinations between mutual predation and cannibalism or factors other than predation could have affected displacement and occurrence patterns of Dikerogammarus species in Southern German waters.     

Ability of native ectomycorrhizal fungi from northern Spain to colonize Douglas-fir and other introduced conifers

 Thirty-six isolates from 27 species of native ectomycorrhizal fungi collected in northern Spain were tested for ectomycorrhiza formation with Pseudotsuga menziesii seedlings in pure culture syntheses. Thirteen of those species were also tested for ectomycorrhiza formation with six other species of conifers (two native and four introduced) to compare their colonization potential. Twenty-three fungal isolates from 18 species formed ectomycorrhizas with Pseudotsuga menziesii. The colonization level of the root system varied markedly among the different fungal species. Eight fungi colonized over 50% of the short roots. Nine fungi did not form ectomycorrhizas even though some of them were collected in pure stands of Pseudotsuga menziesii. Laccaria laccata, Lyophyllum decastes, Pisolithus tinctorius, and Scleroderma citrinum formed abundant ectomycorrhizas on all the conifers tested. Lactarius deliciosus, Rhizopogon spp., and Suillus luteus showed the greatest host specificity. The success in the introduction of some exotic conifers for reforestation in northern Spain is discussed in relation to their compatibility with native ectomycorrhizal fungi. Accepted: 28 August 1995     

Genetic evidence for hybridization between the native Spartina maritima and the introduced Spartina alterniflora (Poaceae) in South-West France: Spartina × neyrautii re-examined

 Spartina alterniflora, a perennial grass native to the North American Atlantic coast, was introduced during the 19th century in western Europe (Southern England and western France) where it hybridized with the native Spartina maritima. In England, the sterile hybrid S. × townsendii gave rise by chromosome doubling to the highly fertile allopolyploid Spartina anglica, which has now invaded many salt marshes and estuaries in western Europe, and has been introduced in several continents. In South-West France, another sterile hybrid was discovered in 1892 in the Bidassoa Estuary, and named Spartina × neyrautii. According to their morphology, some authors suggested that S. × neyrautii and S. × townsendii result from reciprocal crosses. During the 20th century, the hybridization site was severely disturbed, and surviving of S. × neyrautii was questioned. In this paper, various Spartina populations are investigated in the Basque region (France and Spain), and compared to the hybrid taxa formed in England (S. × townsendii and S. anglica). The samples were analyzed using molecular fingerprinting (RAPD and ISSR) and Chloroplast DNA sequence (trnL-trnT spacer, trnL intron and trnL-trnF spacer). In the Bidassoa estuary, a hybrid isolated clone has been found, that displays additive species-specific nuclear markers of S. maritima and S. alterniflora, and that is subsequently considered as a surviving clone of S. × neyrautii. The molecular analyses indicate that S. × neyrautii and S. × townsendii share the same maternal (S. alterniflora), and paternal (S. maritima) parental species, but also that the two independent hybridization events have involved different parental (nuclear) genotypes in England and in South-West France. Received July 12, 2002; accepted October 4, 2002 Published online: March 20, 2003     

Asymmetric larval interactions between introduced and indigenous ladybirds in North America

Understanding the mechanisms that result in the success of introduced species will contribute to predicting future invasions and managing invaded systems. We examined interactions between larvae of two predatory ladybird species recently introduced to North America, Coccinella septempunctata (CS) and Harmonia axyridis (HA), and two indigenous ladybirds, Coccinella transversoguttata (CT) and Hippodamia convergens (HC). By pairing young and old larvae in the laboratory at low and high levels of aphid availability, we assessed the degree of asymmetry in intraguild predation (IGP), the strength of competitive effects on growth and development of larvae escaping predation, and the nature of attack and escape behavior among the species. Interactions were generally asymmetric, with larvae of introduced species acting most frequently as intraguild predators and larvae of indigenous species serving most frequently as intraguild prey (the two Coccinella spp., however, preyed on each other at similar rates). Because they were especially aggressive and because other larvae were least successful in escaping their attacks, larvae of HA had stronger negative effects on larvae of the two indigenous species than did larvae of CS. Such negative effects, expressed most strongly when aphid availability was low, were especially adverse for the smaller of the two indigenous species, HC. In general, older larvae interacted with each other more strongly than young larvae did, and older larvae had especially strong negative effects on young larvae when interactions occurred between age classes. Our results suggest that HA more than CS may represent a threat to indigenous ladybirds as an intraguild predator, and that IGP in turn may play a stronger role for HA than for C. septempunctata in promoting the successful invasion of North America.     

An acanthocephalan parasite mediates intraguild predation between invasive and native freshwater amphipods (Crustacea)

1. The balance of predation between closely related invasive and native species can be an important determinant of the success or failure of biological invasions. In Irish freshwaters, the introduced amphipod Gammarus pulex has replaced the native G. duebeni celticus, possibly through differential mutual intraguild predation (IGP). Theoretically, parasitism could mediate such predation and hence the invasion outcome. However, this idea remains poorly studied. 2. In a field survey, we show that the acanthocephalan parasite Echinorynchus truttae is present in more G. pulex populations than G. d. celticus populations. In addition, within parasitised populations, E. truttae is more prevalent in the invader than in the native. 3. We show for the first time that an acanthocephalan parasite mediates predation between its intermediate macroinvertebrate hosts. In a field experiment, E. truttae parasitism of the invader lowered IGP upon the unparasitised native. In laboratory experiments, parasitism of G. pulex significantly reduced their predatory impact on recently moulted female G. d. celticus. Parasitism also appeared to cause reduction in predatory behaviour, such as attacks per contact on precopula guarded female natives. 4. We conclude that higher parasite prevalence in invaders as compared with natives, by mediation of interspecific interactions, could promote species coexistence, or at least slow species replacements, in this particular biological invasion.     

Interactions between native and invasive gecko lizards in tropical Australia

Invasive geckos of the genus Hemidactylus (Gekkonidae) are spreading rapidly through urban environments in many tropical and subtropical parts of the world. The invaders have caused rapid declines in native gecko abundance in some areas, but their interactions with Australian native species remain unknown. In a small rural village near Darwin, we found that Hemidactylus frenatus is abundant around both lighted and unlighted buildings, but rarely found in surrounding bushland. It is sympatric with the larger Gehyra australis (Gekkonidae) in this disturbed site, and often forages on the same buildings, but is active mostly during the dry‐season (vs. wet‐season for G. australis) and is competitively subordinate to the larger native species. In laboratory encounters, H. frenatus fled from G. australis, and modified its refuge‐site use in the presence of the native lizard. In those same trials, the native gecko often attacked and rarely fled; and did not shift its refuge‐site selection. In field surveys, the two taxa frequently co‐occurred. However, substrate use of the invasive H. frenatus was modified by the presence of the native G. australis, consistent with competitive displacement. Our counts of H. frenatus were highest during the dry‐season, when G. australis (like most other small native reptiles) is relatively inactive. The invasive gecko thus appears to be exploiting a ‘vacant niche’ around buildings, rather than displacing the native gecko taxon. This outcome may reflect the size disparity between the native species and the invader; Hemidactylus frenatus may well have significant ecological impacts on smaller native lizards.     

Inferring relationships between native plant diversity and Lonicera japonica in upland forests in north Mississippi,USA

Question: Do anthropogenic disturbances interact with local environmental factors to increase the abundance and frequency of invasive species, which in turn exerts a negative effect on native biodiversity? Location: Mature Quercus‐Carya and Quercus‐Carya‐Pinus (oak‐hickory‐pine) forests in north Mississippi, USA. Methods: We used partial correlation and factor analysis to investigate relationships between native ground cover plant species richness and composition, percent cover of Lonicera japonica, and local and landscape‐level environmental variables and disturbance patterns in mature upland forests. We directly measured vegetation and environmental variables within 34 sampling subplots and quantified the amount of tree cover surrounding our plots using digital color aerial photography. Results: Simple bivariate correlations revealed that high species richness and a high proportion of herbs were associated with low Lonicera japonica cover, moist and sandy uncompacted soils, low disturbance in the surrounding landscape, and periodic prescribed burning. Partial correlations and factor analysis showed that once we accounted for the environmental factors, L japonica cover was the least important predictor of composition and among the least important predictors of species richness. Hence, much of the negative correlation between native species diversity and this invasive species was explained by soil texture and local and landscape‐level land‐use practices. Conclusions: We conclude that negative correlations between the abundance of invasive species and native plant diversity can occur in landscapes with a gradient of human disturbance, regardless of whether there is any negative effect of invasive species on native species.     

Reciprocal interactions between native and introduced populations of common milkweed,Asclepias syriaca,and the specialist aphid,Aphis nerii

Following its introduction into Europe (EU), the common milkweed (Asclepias syriaca) has been free of most specialist herbivores that are present in its native North American (NA) range, except for the oleander aphid Aphis nerii. We compared EU and NA populations of A. nerii on EU and NA milkweed populations to test the hypothesis that plant–insect interactions differ on the two continents. First, we tested if herbivore performance is higher on EU plants than on NA plants, because the former have escaped most of their herbivores and have perhaps been selected for lower defence levels following introduction. Second, we compared two A. nerii lines (one from each continent) to test whether genotypic differences in the herbivore may influence species interactions in plant–herbivore communities in the context of species introductions. The NA population of A. nerii developed faster, had higher fecundity and attained higher population growth rates than the EU population. There was no overall significant continental difference in aphid resistance between the plants. However, milkweed plants from EU supported higher population growth rates and faster development of the NA line of A. nerii than plants from NA. In contrast, EU aphids showed similar (low) performance across plant populations from both continents. In a second experiment, we examined how chewing herbivores indirectly mediate interactions between milkweeds and aphids, and induced A. syriaca plants from each continent by monarch caterpillars (Danaus plexippus) to compare the resulting changes in plant quality on EU aphid performance. As specialist chewing herbivores of A. syriaca are only present in NA, we expected that plants from the two continents may affect aphid growth in different ways when they are challenged by a specialist chewing herbivore. Caterpillar induction decreased aphid developmental times on NA plants, but not on EU plants, whereas fecundity and population growth rates were unaffected by induction on both plant populations. The results show that genetic variation in the plants as well as in the herbivores can determine the outcome of plant–herbivore interactions.     

Influence of prey availability and conspecifics on patch quality for a cannibalistic forager: laboratory experiments with the wolf spider Schizocosa

 Because cannibals are potentially both predator and prey, the presence of conspecifics and alternative prey may act together to influence the rate at which cannibals prey upon each other or emigrate from a habitat patch. Wolf spiders (Lycosidae) are cannibalistic-generalist predators that hunt for prey with a sit-and-wait strategy characterized by changes in foraging site. Little information is available on how both prey abundance and the presence of conspecifics influence patch quality for these cursorial, non-web-building spiders. To address this question, laboratory experiments were conducted with spiderlings and older juveniles of the lycosid genus Schizocosa. The presence of insect prey consistently reduced rates of spider emigration when spiders were housed either alone or in groups. Solitary juvenile Schizocosa that had been recently collected from the field exhibited a median giving-up time (GUT) of 10 h in the absence of prey (Collembola); providing Collembola increased the median GUT to 64 h. For solitary spiders, the absence of prey increased by about fourfold the rate of emigration during the first 24 h. In contrast, for spiders in patches with a high density of conspecifics, the absence of prey increased the 24-h emigration rate by only 1.6-fold. For successful cannibals in the no-prey patches, the presence of conspecifics improved patch quality by providing a source of food. Mortality by cannibalism was affected by both prey availability and openness of the patch to net emigration. In patches with no net emigration, the presence of prey reduced rates of cannibalism from 79% to 57%. Spiders in patches open to emigration but not immigration experienced a rate of cannibalism (16%) that was independent of prey availability. The results of these experiments indicate that for a cannibalistic forager such as the wolf spider Schizocosa, (1) the presence of conspecifics can improve average patch quality when prey are absent, and (2) cannibalism has the potential to be a significant mortality factor under natural field conditions because cannibalism persisted in prey patches that were open to emigration. Received: 12 April 1996 / Accepted: 14 August 1996     

Aggression and competition for shelter between a native and an introduced crayfish in Europe

1. The introduced North American crayfish, Pacifastacus leniusculus Dana, is expanding its range in Europe and locally often replaces two native crayfish species, Astacus astacus L. and Austropotamobius pallipes Lereboullet. Pacifastacus leniusculus is also expected to invade the habitat of a third native crayfish, the endangered Austropotamobius torrentium Schrank. Interspecific aggressive interactions and competition for shelter were experimentally studied in the laboratory to assess the potential impact of P. leniusculus on A. torrentium .
2. Neither species was inherently dominant in aggressive interactions, but dominance was strongly size-dependent, favouring the larger and faster growing species, P. leniusculus .
3. Access to limited shelter was generally determined by aggressive dominance, although species-specific preference also influenced the outcome of competition for shelter. Austropotamobius torrentium had a higher preference for experimental shelters and often defended these even against larger P. leniusculus .
4. In accordance with theoretical models of animal conflicts, agonistic interactions between equally sized contestants were more severe than between animals of different size.
5. The P. leniusculus used in the present experiment were infected with the crayfish plague, Aphanomyces astaci , to which the animals are resistant. The crayfish transmitted the disease to non-resistant A. torrentium which died ≈ 2 weeks after contact with P. leniusculus .     

Marine invasion processes: interactions between native and introduced marsh snails

We test two hypotheses to explain the success of a biological invader in marshes on the Pacific coast of North America. The first hypothesis focuses on the competitive superiority of an invading species over native species; the second hypothesis focuses on the absence of ecological equivalents in the aboriginal biota, such that no competitive interactions are necessary to facilitate successful invasion. As a model system we experimentally manipulated the introduced pulmonate snail Ovatella myosotis and the native prosobranch snails Assiminea californica and Littorina subrotundata. Manipulations included pair-wise enclosures to determine of Ovatella depressed the growth of the native;species; pair-wise associations to determine whether or not snails demonstrated negative or positive clustering; removal experiments to see if native snails invaded regions occupied by Ovatella, and reciprocal transplants on a vertical gradient to determine the physiological breath of the target species. In addition to monitoring population abundances of all three species seasonally, dietary overlap of these gastropods was also examined. No evidence of competitive superiority by Ovatella was found. Ovatella possesses the physiological capacities to exist in a semiterrestrial environment and is therefore able to take advantage of resources unavailable to native gastropods. We conclude that the successful establishment of this Atlantic snail in the Pacific Northwest did not arise at the expense of native species.