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.     

Evidence for asymmetrical intraguild predation between native and introduced Anolis lizards

Since its introduction, Anolis sagrei (Sauria: Polychrotidae) has been replacing native A. carolinensis in Florida and native A. conspersus in Grand Cayman Island as the common anole of urban environments and other open habitats. To assess the likelihood that predation of juvenile native anoles by A. sagrei adults is an important interaction in this process, the propensities for intraguild predation and cannibalism were assessed for A. sagrei and A. carolinensis in Florida and for A. sagrei and A. conspersus in Grand Cayman. Predation experiments were conducted in cages, using freshly captured lizards, in which adult males of each species were presented with conspecific and heterospecific juveniles. Adult A. sagrei were (1) significantly more likely to eat juveniles than were adult A. carolinensis or A. conspersus, and (2) significantly more likely to eat heterospecific than conspecific juveniles, whereas adult A. carolinensis and A. conspersus were not. Thus, the propensity for intraguild predation is asymmetrical in favor of introduced A. sagrei in Florida and Grand Cayman. Further study is needed, however, to determine the importance of intraguild predation under field conditions. Received: 14 July 1999 / Accepted: 6 March 2000     

Character displacement and coexistence in mud snails (Hydrobiidae)

Summary Populations of coexisting and of allopatrically occurring species of hydrobiid snails (Hydrobia ulvae, H. neglecta, H. ventrosa and Potamopyrgus jenkinsi) have been studied in 90 localities within three different areas. When H. ventrosa coexists with H. ulvae they show character displacement, i.e., the average body size of the former is smaller and that of the latter is larger. When these species live alone they are of approximately the same size. It is shown here that the size ratio between the coexisting species usually found (1.3–1.5) allows stable coexistence based on food praticles size selectivity alone. Variation in the degree of character displacement from locality to locality is explained by different degrees of genetical isolation of the populations. Coexisting H. ventrosa and H. ulvae have shorter, more well-defined periods of reproduction than they do when they occur alone. H. neglecta is larger than coexisting H. ventrosa and smaller than coexisting H. ulvae. Due to the patchy distribution and the fluctuating populations of this species, and due to the fact that pure H. neglecta populations are rare, data on this species are difficult to interpret. Potamopyrgus jenkinsi shows a different food particle size selection than the Hydrobia spp. of identical sizes. Competitive interactions between P. jenkinsi and coexisting Hydrobia spp. are therefore probably weak. In accordance with this, P. jenkinsi does not show character displacement when coexisting with, e.g., H. ventrosa.The fact that the major study area, the Limfjord, is only 150 years old as a marine habitat as well as various more recent man-made changes of the coast line allow estimates of the time scale of the microevolutionary changes which lead to character displacement.     

Competitive abilities of introduced and native grasses

Differencesin competitive ability may explain the maintenance of existing plantpopulationsand the invasion of new areas by plant species. We used field experiments toexamine the competitive responses of Agropyron cristatum(L.) Gaertn., an introduced C₃ grass, and Boutelouagracilis (HBK.) Lag., a native C₄ grass, and thecompetitive effects of Agropyron-dominated vegetation andsuccessional prairie. We also tested whether the outcome of competitiveinteractions varied with water availability. In each vegetation type,transplants of each species were grown under two levels of competition(presenceor absence of neighboring vegetation) and three levels of water availability(high, medium, or low). Transplant survival, growth, and biomass allocationpatterns were measured. Water availability had no effect on the measuredvariables, suggesting that both species were limited by another resource.Growthrates were affected more by competition, while survival and root: shoot ratiowere affected more by transplant species identity. In the successional prairie,neighboring vegetation suppressed the growth of Agropyrontransplants less than that of Bouteloua transplants,suggesting that Agropyron has a stronger ability to resistcompetitive suppression in that vegetation type. The spread ofAgropyron into surrounding vegetation may relate to itsability to resist competitive suppression. In theAgropyron-dominated vegetation, neighboring vegetationsuppressed the growth of both species by the same extent. However, competitionaccounted for more variation in transplant growth inAgropyron-dominated vegetation than in successionalprairie, suggesting that Agropyron has strong competitiveeffects which hinder plant growth and prevent other species from establishinginAgropyron fields.     

Competitive interaction between introduced rainbow trout and native silverside in a Chilean stream

The field experiment presented here elucidates the competitive effect of exotic trout on Chilean silverside. This is the first local experimental work that demonstrates the negative effect of trout on native fish from Chile. Interspecific competition between the Chilean Silverside Basilichthys australis and the rainbow trout Oncorhynchus mykiss was evaluated by an experimental design that used enclosures in a stream of central Chile. Nine enclosures were equally assigned to three treatments, low density, with two silversides; interspecific treatment, with two silversides and four trout; and high density treatment, with six silversides. The results indicate that silversides were weaker competitors, negatively affected by the presence of trout; they showed significant weight loss and reduction in stomach content in the presence of trout.     

Selective crab predation on native and introduced bivalves in British Columbia

Experiments were conducted to determine whether locally abundant crab species prefer co-occurring littleneck clams, Protothaca staminea (Conrad, 1837) and Tapes philippinarum (A. Adams and Reeve, 1850), relative to a recently introduced species, the varnish clam, Nuttallia obscurata, (Reeve, 1857). Prey preference, handling time, pick-up success, profitability and consumption rates were investigated for two crab species, Dungeness crab, Cancer magister (Dana, 1852) and red rock crab, Cancer productus (Randall, 1839) crabs. Both crab species preferred varnish clams over the native species. This may be attributable to the lower handling time, higher pick-up success and increased profitability of consuming varnish clams. Handling time appeared to be a factor not only in species preference, but also in the degree of preference, with shorter handling times corresponding to stronger preference values. Both native and introduced bivalves burrow into the substratum, with the varnish clam burrowing deepest. When feeding on clams in limited substratum both crab species preferred the varnish clam. In the unlimited substratum trials Dungeness crabs preferred varnish clams (although to a lesser degree) while red rock crabs preferred littleneck clams. This was likely due to the significantly deeper burial of the varnish clam, making it less accessible. Although the morphology (i.e. thin shell, compressed shape) of the invader increases its vulnerability to predation, burial depth provides a predation refuge. These results demonstrate how interactions between native predators and the physical characteristics and behaviour of the invader can be instrumental in influencing the success of an invasive species.     

Parasite-mediated predation between native and invasive amphipods

Parasites can structure biological communities directly through population regulation and indirectly by processes such as apparent competition. However, the role of parasites in the process of biological invasion is less well understood and mechanisms of parasite mediation of predation among hosts are unclear. Mutual predation between native and invading species is an important factor in determining the outcome of invasions in freshwater amphipod communities. Here, we show that parasites mediate mutual intraguild predation among native and invading species and may thereby facilitate the invasion process. We find that the native amphipod Gammarus duebeni celticus is host to a microsporidian parasite, Pleistophora sp. (new species), with a frequency of infection of 0-90%. However, the parasite does not infect three invading species, G. tigrinus, G. pulex and Crangonyx pseudogracilis. In field and laboratory manipulations, we show that the parasite exhibits cryptic virulence: the parasite does not affect host fitness in single-species populations, but virulence becomes apparent when the native and invading species interact. That is, infection has no direct effect on G. d. celticus survivorship, size or fecundity; however, in mixed-species experiments, parasitized natives show a reduced capacity to prey on the smaller invading species and are more likely to be preyed upon by the largest invading species. Thus, by altering dominance relationships and hierarchies of mutual predation, parasitism strongly influences, and has the potential to change, the outcome of biological invasions.     

Ocean acidification increases the vulnerability of native oysters to predation by invasive snails

There is growing concern that global environmental change might exacerbate the ecological impacts of invasive species by increasing their per capita effects on native species. However, the mechanisms underlying such shifts in interaction strength are poorly understood. Here, we test whether ocean acidification, driven by elevated seawater pCO₂, increases the susceptibility of native Olympia oysters to predation by invasive snails. Oysters raised under elevated pCO₂ experienced a 20% increase in drilling predation. When presented alongside control oysters in a choice experiment, 48% more high-CO₂ oysters were consumed. The invasive snails were tolerant of elevated CO₂ with no change in feeding behaviour. Oysters raised under acidified conditions did not have thinner shells, but were 29–40% smaller than control oysters, and these smaller individuals were consumed at disproportionately greater rates. Reduction in prey size is a common response to environmental stress that may drive increasing per capita effects of stress-tolerant invasive predators.     

Genetic diversity and molecular markers in introduced and Thai native apple snails (Pomacea and Pila)

The genetic diversity and species-diagnostic markers in the introduced apple snail, Pomacea canaliculata and in the native Thai apple snails; Pila ampullacea, P. angelica, P. pesmei, and P. polita, were investigated by restriction analysis of COI and are reported for the first time. Twenty-one composite haplotypes showing non-overlapping distributions among species were found. Genetic heterogeneity analysis indicated significant differences between species (P < 0.0001) and within P. pesmei (P < 0.0001) and P. angelica (P < 0.0004). No such heterogeneity was observed in Pomacea canaliculata (P > 0.0036 as modified by the Bonferroni procedure), P. ampullacea (P = 0.0824-1.000) and P. polita (P = 1.0000). A neighbor-joining tree based on genetic distance between pairs of composite haplotypes differentiated all species and indicated that P. angelica and P. pesmei are closely related phylogenetically. In addition, the 16S rDNA of these species was cloned and sequenced. A species-specific PCR for P. canaliculata was successfully developed with a sensitivity of detection of approximately 50 pg of the target DNA template. The amplification of genomic DNA (50 pg and 25 ng) isolated from the fertilized eggs, and juveniles (1, 7, and 15 d after hatching) of Pomacea canaliculata was also successful, and suggested that Pomacea canaliculata and Pila species can be discriminated from the early stages of development.     

Phenotypic and genetic differentiation between native and introduced plant populations

Plant invasions often involve rapid evolutionary change. Founder effects, hybridization, and adaptation to novel environments cause genetic differentiation between native and introduced populations and may contribute to the success of invaders. An influential idea in this context has been the Evolution of Increased Competitive Ability (EICA) hypothesis. It proposes that after enemy release plants rapidly evolve to be less defended but more competitive, thereby increasing plant vigour in introduced populations. To detect evolutionary change in invaders, comparative studies of native versus introduced populations are needed. Here, we review the current empirical evidence from: (1) comparisons of phenotypic variation in natural populations; (2) comparisons of molecular variation with neutral genetic markers; (3) comparisons of quantitative genetic variation in a common environment; and (4) comparisons of phenotypic plasticity across different environments. Field data suggest that increased vigour and reduced herbivory are common in introduced plant populations. In molecular studies, the genetic diversity of introduced populations was not consistently different from that of native populations. Multiple introductions of invasive plants appear to be the rule rather than the exception. In tests of the EICA hypothesis in a common environment, several found increased growth or decreased resistance in introduced populations. However, few provided a full test of the EICA hypothesis by addressing growth and defence in the same species. Overall, there is reasonable empirical evidence to suggest that genetic differentiation through rapid evolutionary change is important in plant invasions. We discuss conceptual and methodological issues associated with cross-continental comparisons and make recommendations for future research. When testing for EICA, greater emphasis should be put on competitive ability and plant tolerance. Moreover, it is important to address evolutionary change in characteristics other than defence and growth that could play a role in plant invasions.     

Competition between a native and introduced pollinator in unmanaged urban meadows

Biological Invasions - There is little understanding of the role of exotic bees in the observed pollinator decline. Recent models of species invasions predict that aggressive species have more...     

Competition between introduced and native spiders (Araneae: Linyphiidae)

The European sheet-web spider Linyphia triangularis (Araneae: Linyphiidae) has become established in Maine, where it often reaches very high densities. Two lines of evidence from previous work suggest that L. triangularis affects populations of the native linyphiid spider Frontinella communis. First, F. communis individuals are relatively scarce in both forest and coastal habitat where L. triangularis is common, but more common where L. triangularis is at low density. Second, in field experiments, F. communis species are less likely to settle in experimental plots when L. triangularis is present, and F. communis disappears from study plots when L. triangularis is introduced. Here we test two mechanisms that may underlie these patterns. First, we tested whether L. triangularis invades and usurps the webs of F. communis. When spiders were released onto webs of heterospecifics, L. triangularis was more likely to take over or share webs of F. communis than the reverse. We also observed natural takeovers of F. communis webs. Second, we explored the hypothesis that L. triangularis reduces prey availability for native species. We sampled flying prey in areas with L. triangularis and those where it had been removed, and found no effect of spider presence on measured prey density. We also found no effect of prey supplementation on web tenacity in F. communis, suggesting that F. communis movements are not highly dependent on prey availability. We conclude that web takeover is likely more important than prey reduction in driving negative effects of L. triangularis on F. communis.     

Hybridization dynamics between Colorado's native cutthroat trout and introduced rainbow trout

Newly formed hybrid populations provide an opportunity to examine the initial consequences of secondary contact between species and identify genetic patterns that may be important early in the evolution of hybrid inviability. Widespread introductions of rainbow trout (Oncorhynchus mykiss) into watersheds with native cutthroat trout (Oncorhynchus clarkii) have resulted in hybridization. These introductions have contributed to the decline of native cutthroat trout populations. Here, we examine the pattern of hybridization between introduced rainbow trout and 2 populations of cutthroat trout native to Colorado. For this study, we utilized 7 diagnostic, codominant nuclear markers and a diagnostic mitochondrial marker to investigate hybridization in a population of greenback cutthroat trout (Oncorhynchus clarkii stomias) and a population of Colorado River cutthroat trout (Oncorhynchus clarkii pleuriticus). We infer that cutthroat-rainbow trout hybrid swarms have formed in both populations. Although a mixture of hybrid genotypes was present, not all genotype combinations were detected at expected frequencies. We found evidence that mitochondrial DNA introgression in hybrids is asymmetric and more likely from rainbow trout than from cutthroat trout. A difference in spawning time of the 2 species or differences in the fitness between the reciprocal crosses may explain the asymmetry. Additionally, the presence of intraspecific cytonuclear associations found in both populations is concordant with current hypotheses regarding coevolution of mitochondrial and nuclear genomes.     

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 native lady beetle decline

Coccinellid communities across North America have experienced significant changes in recent decades, with declines in several native species reported. One potential mechanism for these declines is interference competition via intraguild predation; specifically, increased predation of native coccinellid eggs and larvae following the introduction of exotic coccinellids. Our previous studies have shown that agricultural fields in Michigan support a higher diversity and abundance of exotic coccinellids than similar fields in Iowa, and that the landscape surrounding agricultural fields across the north central U.S. influences the abundance and activity of coccinellid species. The goal of this study was to quantify the amount of egg predation experienced by a native coccinellid within Michigan and Iowa soybean fields and explore the influence of local and large-scale landscape structure. Using the native lady beetle Coleomegilla maculata as a model, we found that sentinel egg masses were subject to intense predation within both Michigan and Iowa soybean fields, with 60.7% of egg masses attacked and 43.0% of available eggs consumed within 48 h. In Michigan, the exotic coccinellids Coccinella septempunctata and Harmonia axyridis were the most abundant predators found in soybean fields whereas in Iowa, native species including C. maculata, Hippodamia parenthesis and the soft-winged flower beetle Collops nigriceps dominated the predator community. Predator abundance was greater in soybean fields within diverse landscapes, yet variation in predator numbers did not influence the intensity of egg predation observed. In contrast, the strongest predictor of native coccinellid egg predation was the composition of edge habitats bordering specific fields. Field sites surrounded by semi-natural habitats including forests, restored prairies, old fields, and pasturelands experienced greater egg predation than fields surrounded by other croplands. This study shows that intraguild predation by both native and exotic predators may contribute to native coccinellid decline, and that landscape structure interacts with local predator communities to shape the specific outcomes of predator-predator interactions.