Feral pigs (Sus scrofa) are considered to be among the world’s worst invasive species due to their successful invasion and ecological and economic impact to native and agricultural plants and animals around the world. Feral pigs are significant disturbance agents that destroy plant communities, change soil characteristics, alter nutrient cycling, and create open sites for colonization of both native and non-native plant species through their foraging behavior called rooting. In contrast to native animal disturbances, rooting is a striking feature in the landscape that varies in space, seasonal timing, frequency (number of times rooted), and intensity (depth of rooting). During this study, feral pigs rooted 7.7% of the search area, which increased to 12% when abandoned patches (baseline patches that were not rooted during this study) were included. Overall, feral pigs rooted and re-rooted habitats along roads and trails significantly more than wetlands. Rooting also varied temporally with the most rooting occurring during July-November, which also corresponds to the peak in rooting intensity. Implications to land managers include avoiding the installation of roads and trails near wet to mesic habitats or other habitats that contain species of concern in order to conserve habitat quality and recreational value. Despite less rooting activity, feral pigs still pose a significant threat to wetlands as evidenced by the large amount of abandoned patches documented. In order to conserve natural areas, effective management and development of efficient control methods is needed to keep feral pig populations in check. As a large opportunistic omnivore, feral pigs have the potential to be important vectors for endozoochorus seed dispersal of a variety of plant species. Feral pigs can travel long iv distances and have a gut retention time up to 49 hours, therefore seeds can be deposited throughout the landscape far from the parent plant. Over the course of this study, feral pigs dispersed 50 plant species from a wide range of ecological and morphological characteristics, though the majority were native, small seeded, wetland species. For most plant species, location of deposition matched their habitat preference and suggests a high probability of survival. Feral pigs disperse mainly wetland plant species, which has important implications for wetland conservation. However, feral pigs also deposited unwanted species into wetlands and predated the seeds of important wetland canopy tree species.

Predicting the locations of new biological invasions has become a high priority for biologists as well as trying to predict if newly introduced species will become damaging to native ecosystems. Reefs of the eastern oyster Crassostrea virginica in Mosquito Lagoon, Florida have been highly disturbed in recent years resulting in dead reefs (piles of dead, disarticulated shells) some of which have been restored. I conducted oyster reef surveys for non-native invertebrates to determine if disturbance on these oyster reefs might assist invasion by two species, Mytella charruana and Perna viridis, recently introduced to the southeastern coast of the United States. Next, I investigated if M. charruana's temperature and aerial exposure tolerance limits may allow for it to establish permanently on intertidal oyster reefs. Temperature and aerial exposure tolerance experiments were conducted and oyster reef temperatures were collected. Oyster reef surveys could not predict if reef disturbance is assisting in the invasion process because only two non-native individuals (P.viridis) were found, one on a restored reef and one on a natural (reference) reef. Tolerance experiments showed that some Mytella charruana survived even after 7 days of 8??C temperatures if the mussels are exposed to air for 4 hours or less per day. Mytella charruana had near 0% survival after 4 hours of 44??C. However, only disturbed reefs reached this temperature in the field. It is likely that M. charruana could survive in the low intertidal zone on restored or reference reefs. This information is important for understanding the introduction of M. charruana in Mosquito Lagoon and also provides a data set of temperature tolerances for better understanding of whether the species might be able to invade other areas.

For this dissertation, I measured how plant communities in Appalachian forests responded to disturbances caused by forest management activities. I had two primary objectives including (1) testing theories of biological diversity and invasions by nonnative species; and (2) providing empirical data that will help guide the sustainable use of forest resources. This work is part of the Southern Appalachian Silviculture and Biodiversity (SASAB) experiment that was established in the early 1990s to investigate ecosystem responses to a gradient of timber harvesting disturbances. Ranging from undisturbed controls to silvicultural clearcuts, the disturbance gradient is replicated at sites located throughout the Appalachian mountains of Virginia and West Virginia. The plant community was sampled across a wide range of spatial scales (2 hectares to 1 m2) using a nested sampling design and was also sampled at three times including pre-disturbance, one year post-disturbance, and ten year post-disturbance. For one element of the study I tested modern theories of biological invasions and investigated how the forest disturbance gradient interacted with species diversity to influence invasion by nonnative species (Chapter 2). Contrary to popular ecological theories of biotic resistance, the most diverse sites tended to be more easily invaded following intense canopy disturbance. Interestingly, none of the sites in this study were dominated by invasive plants, which led me to ask whether dominant tree species in forests provided resistance to nonnative plant establishment and growth through the quality of the litter they produce. I also asked how might animals that are known to alter litter layers interact with dominant tree species to influence plant invasions. Therefore, I conducted an experiment investigating how changes in litter from dominant tree species and invasions by nonnative earthworms might influence invasibility of forests using forest floor mesocosms (Chapter 3). I found that plant invasion was inhibited by native oak litter even when earthworms were present, suggesting that oak forests may resist plant invasions via oak tree litter. In contrast, plant invasion was greater under invasive tree litter and earthworm activity tended to facilitate invasive plant success only under invasive tree litter. I was also interested in understanding how disturbance might alter relationships between local and regional diversity. The long-term data of the forest disturbance experiment allowed me to investigate how local species richness is mediated by regional species richness after disturbance and during forest community development (Chapter 4). Local richness depended strongly on regional richness only after disturbance via colonization of species, but this relationship changed during forest aggradation. These results suggest that regional species pools are important to maintain local diversity following disturbance, but that local interactions (through canopy closure of dominant trees) exert control over species diversity during community reorganization. Lastly, I tested current theories on how diversity influences compositional stability after disturbance (Chapter 5). Disturbance consistently resulted in decreased compositional stability, but diversity was associated with stability in complex ways, which depended on how stability was measured and at what scale. Species-rich areas were in some instances less stable; in other instances areas with intermediate levels of diversity were more stable. These results suggest that disturbance causes shifts in species composition via colonization, but the ways in which diversity of sites influences compositional stability is complex and depends on methods used and the scales of observation. Taken together, these results suggest that disturbance influences invasibility, species saturation, and compositional stability of ecological communities. These properties change immediately following disturbance, and during forest development and canopy closure. Data from this project were useful in testing existing theories of community ecology, and may ultimately prove useful for forest managers as they decide how to protect biodiversity while planning for other uses of forest resources. Overall, these results suggest that colonization of species is the primary process driving plant community patterns in Appalachian forests following disturbance. / Ph. D.

Nonnative fishes are a major threat to biodiversity and new species continue to be introduced. In this dissertation, I described patterns and assessed determinants of fish invasions in the Mid-Atlantic region of the United States. Data on nonnative fish distributions were obtained from the United States Geological Survey's Nonindigenous Aquatic Species Database (NASD). Nonnative fishes are introduced by a variety of pathways, and prevention efforts can be optimized by focusing on pathways posing the greatest risk of new invasions. To assess the importance of existing pathways, I described the species associated with each pathway, analyzed the number of species introduced by decade for certain pathways, and estimated the detectability and probability of establishment of species introduced by each pathway. Additionally, I reviewed the efficacy of existing regulations for preventing introductions via each pathway. Historically, the intentional introduction of centrarchids and salmonids for sport was the dominant pathway. Pathways currently posing the greatest risks included bait release, illegal introductions, stocking of private ponds, and several pathways associated with economic activities. These pathways involved cyprinids, catostomids, and species exotic to North America. Regulations varied considerably among states, and I suggest that region-wide prohibitions on the release of nonnative species into the wild would help prevent additional introductions. Mid-Atlantic region watersheds differ considerably in nonnative species richness (NNSR), suggesting they are not equally invasible. I analyzed relationships between ecosystem characteristics and invasibility by compiling data on watershed characteristics and correlating these with NNSR. I included measures of colonization pressure (i.e., the number of species introduced) and research effort, which can bias patterns of NNSR. After controlling for these factors, the range in elevation in a watershed explained the greatest variation in NNSR. Highland watersheds had greater NNSR, probably because of greater habitat heterogeneity due in part to human activity. I suggest that NNSR can be reduced by restoration activities that reduce the diversity of artificial habitats available in highland watersheds. Ecosystems with similar NNSR may be invaded by different species, because differences in ecosystem characteristics may regulate the types of species that are able to establish. To explain differences in nonnative species among ecosystem types, I grouped Mid-Atlantic region watersheds by nonnative community and tested for differences in ecosystem characteristics among groups. Four groups were identified. A large, speciose group in the north-west portion region was characterized by smallmouth bass (Micropterus dolomieu). A large, speciose group in the south-west portion of the region was characterized by largemouth bass (Micropterus salmoides). Two smaller groups with few species were found on the coastal plain; one to the north, characterized by black crappie (Pomoxis nigromaculatus) and a second to the south characterized by white crappie (Pomoxis annularis). Nonnative community type was correctly predicted 80% of the time by models based on temperature and range in elevation. Relatively uninvaded watersheds in the south-east part of the region were predicted to host the most diverse nonnative community, suggesting that risks of invasion are high there. These results demonstrate the importance of species identity in determining ecosystem invasibility. There is no consensus on how to estimate the relative impacts of nonnative species. I developed and compared several approaches for doing so. I estimated impact by surveying fish biologists regarding the abundance and socioeconomic and ecological impacts of each species. I obtained fish collection records as an additional estimate of abundance and consulted reports of impacts in the NASD. I consulted reports of impacts in global invasive species databases as a basis for comparison. I compared top-ranked species among approaches, and game and non-game biologists' ratings of game and non-game species for each survey question. Top-ranked species differed considerably among approaches. Non-game biologists gave higher ecological impact ratings to both game and non-game species. Approaches assessing socioeconomic impacts are most appropriate for informing social decisions, such as restricting the possession or trade of a species. A combination of data from approaches assessing ecological impacts and abundance is most appropriate for studies of ecological patterns, such as testing for differences in traits between high- and low-impact species. These approaches are transferrable to other regions and taxa, and can inform management decisions and improve efforts to identify factors correlated with high-impact invaders. Collectively, my results can aid in reducing the effects of nonnative fish invasions by enabling managers to focus prevention efforts on high-impact species likely to invade particular ecosystems via known pathways. For example, bait releases, illegal introductions, private stocking, and several pathways associated with economic activities present the highest risks of future invasions, and warrant attention aimed at preventing invasions. Prevention could also be focused on several watersheds in the south-east part of the region, which currently have few established species but were predicted to be invasible by bluegill (Lepomis gibbosus), bluntnose minnow (Pimephales notatus) channel catfish (Ictalurus punctatus), and warmouth (Lepomis gulosus). This work represents major advancements in invasion biology, including new links between species identity and ecosystem invasibility and the development of methods for quantifying impact. / Ph. D.

Le dialecte syrien de Tartous ne dispose pas d’adverbes dérivés d’un adjectif, comme c’est le cas en français, pouvant fonctionner comme adverbes de manière. Par contre,Les deux langues utilisent deux moyens similaires pour exprimer la manière. Le premier concerne la construction introduite par la préposition avec en français et parla préposition bə (‘associatif’) en syrien de Tartous, et le second, l’objet interne en français et l’objet dit absolu en syrien de Tartous. Compte tenu de la différence entre les systèmes linguistiques des deux langues, nous pourrons faire l’hypothèse que même dans les cas des deux moyens comparables dont disposent les deux langues pour exprimer la manière, ils ne sont pas soumis aux mêmes types de contraintes syntaxiques et sémantiques en français et en syrien de Tartous. Cette étude comparative nous permettra de comprendre le fonctionnement de chacun des moyens utilisés dans les deux langues pour exprimer la manière, et par là de faire ressortir les différences syntaxique et sémantique qui séparent les compléments de manière dans les deux langues. / The Syrian dialect of Tartous does not have adverbs derived from adjectives that work as adverbs of manner, like in French. On the other hand, both languages use two similar ways of expressing condition ; the first one is construct introduced by thepreposition ‘with’ (‘avec’ in French, ‘bi’ in Syrian). The second way is the cognate object, or what is known in Syrian as ‘absolute effect’.With the differences between the two language systems in mind, we can assume that even if we have similar ways of expressing manner, yet they are controlled by different grammatical and semantic rules.Comparing the two languages allows us to understand the method of expressing manner in each language, and then identify the semantic and grammatical distinctions between the adverbs of manner in the two languages.

This thesis focuses on the guppy, Poecilia reticulata, as an invasive species. Its non-native distribution, the biology behind its success and the reasons for its introduction are examined. A worldwide email survey revealed that the guppy is established in at least 73 countries outside of its native range and that mosquito control schemes and the release of unwanted aquarium fish are the two primary routes of introduction. Knowledge gaps were identified; primarily the scarcity of scientific evidence for negative impacts of guppy introductions and similarly for mosquito control efficacy. Replicated mesocosm experiments demonstrated that female guppies are capable of routinely establishing populations, and that these retain behavioural viability over several generations. The first mesocosm study suggested that founders with very different evolutionary histories were equally good at establishing populations. The second mesocosm study suggested that monandrous females were extremely successful at establishing behaviourally viable populations, with no decline in behavioural variation. The effectiveness of guppies as mosquito control agents was examined in two related foraging experiments. The first study found little evidence for the presence of ‘prey switching’ in guppies, questioning the validity of previous work advocating their introduction to stabilise prey populations. The second study revealed a preference for non-vector mosquito larvae in a two-prey system. However, both mosquito species were consumed equally readily when habitat complexity increased. The presence of conspecifics affected female foraging behaviour. The presence of males reduced the strength of prey preference in the first study, and the presence of conspecifics of either sex removed prey preference in the second. Both demonstrate that multi-prey systems have important implications for the efficacy of poeciliids in biological control. Despite severe demographic bottlenecks, their adaptability and ability to rapidly increase in numbers enable guppies to establish and persist when introduced. Such bottlenecks are typical of introduction scenarios, warning that particular caution should be exerted when introducing this species, or other livebearing fish, to natural water bodies.

Predicting the impacts of non-native species remains one of the greatest challenges to invasion ecologists. Because of their insularity, freshwater systems are particularly vulnerable to invasions, especially from non-native predators. The research in this thesis explores the role of abiotic and biotic factors in mediating the distribution and impact of Bythotrephes longimanus, a predatory cladoceran that has been introduced to freshwater systems in North America. Although the general impacts of this invasion have been documented, little is known about the factors that modulate them. Using a combination of field surveys and experiments, I tested several hypotheses concerning the influence of interactions with native species, as well as the role of heterogeneity in the light environment, in mediating the impact of Bythotrephes. Results demonstrated that biotic resistance by native macroinvertebrate predators does not play a limiting role in the establishment success of Bythotrephes. However, the within-lake distribution of Bythotrephes was influenced by these macroinvertebrates, suggesting that the native predator context matters when trying to understand the impacts of non-native predators. This was demonstrated with a mesocosm experiment in which the impact of Bythotrephes was constrained by the native Chaoborus larvae. In terms of the abiotic environment, in situ feeding experiments demonstrated that refuges from impact could exist for some prey taxa, as the outcome of predation by Bythotrephes was dependent on light availability and some prey taxa were more successful at evading predation under low light conditions. Finally, results show that adaptive behaviour by prey is also an important determinant of impact, as migrating Daphnia can escape predation effects by Bythotrephes. The combination of light-limited predation and a shallow distribution by Bythotrephes selects for prey that occupy relatively deeper positions during the day. As a whole, this research highlights the importance of complex interactions in mediating the impact of Bythotrephes and may help to explain some of the variation that has been documented among invaded lakes. A better understanding of these complex interactions can improve our ability to anticipate impacts as Bythotrephes continues to spread, as well as provide insight on some of the long-term effects following invasion. / Thesis (Ph.D, Biology) -- Queen's University, 2013-06-17 09:26:35.221

The Argentine black and white tegu, Salvator merianae, is a nonnative species that has invaded parts of Florida. The potential impacts of this species on native Florida wildlife are not yet known. This study looks at the stomach contents of 169 S. merianae captured between 2011 and 2013 in south Florida to infer potential impacts of S. merianae and spatial or seasonal shifts in diet. Analysis of 169 GI tracts showed that S. merianae is an omnivorous, terrestrial forager with a broad dietary range which includes insects, fruits, plants, snails, crayfish, carrion, birds, small mammals, turtles, snakes, lizards, frogs and eggs. S. merianae diet composition varied with capture habitat, the fattest tegus were collected from disturbed/agricultural areas and these samples contained, primarily, gastropods and insects. Tegu dietary habits threaten local endangered and state listed species such as the American crocodile, Crocodylus acutus, and the Cape Sable seaside sparrow, Ammodramus maritimus mirabilis. Dedicated funds and efforts need to be focused on this species to limit its further spread and future impact on native species. / Includes bibliography. / Thesis (M.S.)--Florida Atlantic University, 2015. / FAU Electronic Theses and Dissertations Collection

The decline of the Oregon Spotted Frog (Rana pretiosa), a Pacific Northwest endemic now federally listed as threatened, has been attributed to several aspects of ecosystem alteration, primarily habitat degradation and loss. The introduced American Bullfrog (Rana (Aquarana) catesbeiana) has been widely implicated in those declines, but the basis of that contention has been difficult to characterize. The bullfrog occurring at every site of recent Oregon Spotted Frog extirpation has focused concern about its impact. Here, I present a suite of interconnected studies that examine the behavioral ecology of both species to better understand the potential for bullfrog-mediated Oregon Spotted Frog extirpation. I quantified Oregon Spotted Frog anti-predator behavior from the only known population successfully co-occurring with bullfrogs (Conboy Lake) and a population devoid of bullfrog impact (Big Marsh), and compared these behaviors to the predatory traits of the bullfrog. The initial study revealed that captive-reared individuals from the Oregon Spotted Frog population that has successfully co-occurred with bullfrogs respond faster to a predatory stimulus (measured as latency to response) than Oregon Spotted Frogs from a population not to exposed to bullfrogs. Subsequent field investigations of the approach distance allowed by a predator stimulus before taking evasive action (termed the flight initiation distance: FID) conducted with the Oregon Spotted Frog population co-occurring with bullfrogs first demonstrated that FID of recently metamorphosed bullfrogs is consistently greater than that of recently metamorphosed Oregon Spotted Frogs. Further, examination of FID across all post-metamorphic age classes of Oregon Spotted Frogs revealed that older frogs do not allow as close approach as recently metamorphosed Oregon Spotted Frogs. This age class shift in FID did not occur in the Oregon Spotted Frog population not exposed to bullfrogs. In the latter population, FID did not differ among age classes. Since the bullfrog might be driving this age-based change in anti-predator behavior, I explored the variation in strike distance of bullfrogs from the site of co-occurrence in both the field and laboratory to determine the extent of overlap with Oregon Spotted Frog FID. I found that the bullfrog strike distance significantly overlaps the FID of all ages of Oregon Spotted Frogs from the bullfrog-free site but only that of youngest (recently metamorphosed) frogs at the site of co-occurrence. Older Oregon Spotted Frogs from the site of co-occurrence generally escaped at distances greater than the strike distance of bullfrogs. I also collected > 880 bullfrogs from the site of co-occurrence and analyzed the stomach contents to assess their dietary trends. I found that bullfrogs consume Oregon Spotted Frogs at the site, but do not eat the larger (older) frogs. Moreover, the body size ratio between Oregon Spotted Frogs as prey and bullfrogs as predators suggests that nearly all of the adult size distribution of bullfrogs at Conboy would be incapable of preying on adult Oregon Spotted Frogs. Collectively, these studies strongly suggest that bullfrogs have altered the escape behavior of Oregon Spotted Frogs at Conboy Lake and that most adult Oregon Spotted Frogs at Conboy may have a size-based release from predation by bullfrogs. Implicit in this finding is that bullfrogs may pose a real threat via predation to other Oregon Spotted Frog populations with which they might come into contact where the distribution of bullfrog body sizes differ substantially from that at Conboy Lake.

Lethal biotic interactions strongly influence the potential for aquatic non-native species to establish and endure in habitats to which they are introduced. Predators in the recipient area, including native and previously established non-native predators, can prevent establishment, limit habitat use, and reduce abundance of non-native species. Management efforts by humans using methods designed to cause mass mortality (e.g., trapping, biocide applications) can reduce or eradicate non-native populations. However, the impacts of predator and human induced mortality may be mitigated by the behavior or population-level responses of a given non-native species. My dissertation examined the responses of non-native aquatic species to the risk of predation by novel (i.e., no previous exposure) predators in the recipient community and indicators of potential compensatory responses by non-native populations to increased mortality resulting from removal efforts. My dissertation addresses four primary questions. 1) Can first generation, naïve invaders recognize and defend against predators found within the region of invasion through the expression of inducible defenses? 2) Can the overcompensatory potential of a population be predicted through examinations of intraspecific interactions of individuals from the population? 3) What is the relationship between removal effort outcome (i.e., successful or unsuccessful reduction of the target population) and compensatory population responses? 4) Is there a relationship between characteristics of removal efforts that are typically available to managers (e.g., target area size, target area connectivity, removal methodology) and compensatory population responses that could indicate the relative likelihood of compensation resulting from removal efforts? An invading species should be more likely to establish if it can successfully identify and defend against predators in the recipient range, such as through the expression of inducible defenses. Inducible defenses are behavioral or physiological changes that reduce an organism's susceptibility to predation. Through a series of laboratory experiments, I tested whether inducible defenses, in the form of increased burrowing depth, may have benefited the early stage of invasion of Nuttallia obscurata (purple varnish clam), an established Northeast Pacific invader. Specimens of N. obscurata were collected from introduced populations in the Northeast Pacific and from a native population in Japan. The clams were exposed to chemical and physical cues from Northeast Pacific crab predators, including the native Metacarcinus magister (Dungeness crab), an abundant and frequent predator of N. obscurata. While introduced N. obscurata increased their burrowing depth in the physical presence of M. magister, clams collected from their native range showed no such response. This lack of increased burrowing depth by naïve clams in response to a predator native to the newly invaded range, but a significant increase in depth for clams from populations established in the range suggests that while inducible defenses likely did not contribute to the initial establishment of N. obscurata in the Northeast Pacific, they may contribute to their continued persistence and expansion in their introduced range. Some efforts to reduce invasive populations have paradoxically led to population increases. This phenomenon, referred to as overcompensation, occurs when strong negative density-dependent interactions are reduced through increased mortality within a population, resulting in an increase in the population's recruitment rate sufficient to increase the population's overall abundance. Increases in a population's recruitment rate can result from reduced cannibalism of juveniles resulting in lower mortality of new recruits, from increased adult reproductive output, which increases the number of potential recruits, or from reductions in size and/or age at maturity of the unharvested population, which increases the number of reproductive individuals. I predicted the overcompensatory potential of a population of Carcinus maenas (European green crab) in Bodega Harbor, California, using a series of laboratory and field experiments examining intraspecific pressures of adults on juveniles in the population. This measure of intraspecific pressure was used to predict the overcompensatory potential of the population in response to increased mortality from ongoing removal efforts. This prediction was then assessed using pre- and post-removal surveys of juvenile recruitment in Bodega Harbor compared to nearby populations, testing for evidence of overcompensation. While adult C. maenas in Bodega Harbor had limited negative impacts on juveniles, I concluded it was unlikely to result in overcompensation. Relative juvenile abundance did not statistically increase in removal compared to reference populations, consistent with my conclusion from the experiments. Increases in recruitment rates can occur as a result of efforts to remove non-native species. This increase in recruitment can result in overcompensation, but more commonly results in compensation, where recruitment rates increase relative to pre-removal recruitment but does not result in in the population's abundance exceeding pre-removal levels. However, a detailed and accurate prediction of the response of a population to harvest is time consuming and data intensive. This is not feasible for most efforts to eradicate non-native species, which have the greatest chance of success when enacted rapidly after detection. For my final chapter, I performed a literature review and accompanying statistical analysis to determine if typically available information related to the removal effort (site size, site connectivity, and removal technique) could be used to determine increased risk of compensation for a given effort to remove aquatic invasive species. Compensation was closely linked to unsuccessful removal efforts and was observed only among efforts utilizing physical removal methods. However, the frequency with which compensation occurred varied with the exact technique employed, occurring most frequently in removal utilizing electrofishing. Additionally, evidence of compensation was more frequent among larger removal areas with variable connectivity. While other predictors (temperature, effort, etc) might add to the predicative power, the findings of the review provide criteria for managers to determine the relative risk of compensation prior to the start of removal. Further understanding of how invasive species respond to lethal biotic interactions, including anthropogenically mediated control measures, can aid in assessing the risk of invasion for a given species and inform managers of the risk of complications resulting from removal efforts. While inducible defenses may contribute to the long-term success of an introduced species in their recipient range, my findings did not support the idea that inducible defenses triggered by predator cues contributed to their initial introduction in this case. However, research on other non-native species and offspring of previously naïve prey would allow for a clearer picture of the role of inducible defenses in the invasion process. Compensation resulting from removal efforts does not guarantee failure, and certain characteristics of removal efforts seem to indicate increased risk of compensation. Together these components help identify how biotic interactions surrounding mortality risk of an invading species help shape the trajectory of invasion.