Aim Invasion ecology includes many hypotheses. Empirical evidence suggests that most of these can explain the success of some invaders to some degree in some circumstances. If they all are correct, what does this tell us about invasion? We illustrate the major themes in invasion ecology, and provide an overarching framework that helps organize research and foster links among subfields of invasion ecology and ecology more generally. Location Global. Methods We review and synthesize 29 leading hypotheses in plant invasion ecology. Structured around propagule pressure (P), abiotic characteristics (A) and biotic characteristics (B), with the additional influence of humans (H) on P, A and B (hereon PAB), we show how these hypotheses fit into one paradigm. P is based on the size and frequency of introductions, A incorporates ecosystem invasibility based on physical conditions, and B includes the characteristics of invading species (invasiveness), the recipient community and their interactions. Having justified the PAB framework, we propose a way in which invasion research could progress. Results By highlighting the common ground among hypotheses, we show that invasion ecology is encumbered by theoretical redundancy that can be removed through integration. Using both holistic and incremental approaches, we show how the PAB framework can guide research and quantify the relative importance of different invasion mechanisms. Main conclusions If the prime aim is to identify the main cause of invasion success, we contend that a top‐down approach that focuses on PAB maximizes research efficiency. This approach identifies the most influential factors first, and subsequently narrows the number of potential causal mechanisms. By viewing invasion as a multifaceted process that can be partitioned into major drivers and broken down into a series of sequential steps, invasion theory can be rigorously tested, understanding improved and effective weed management techniques identified.
We suggest Milankovitch climate oscillations as a common cause for geographical patterns in species diversity, species' range sizes, polyploidy, and the degree of specialization and dispersability of organisms. Periodical changes in the orbit of the Earth cause climatic changes termed Milankovitch oscillations, leading to large changes in the size and location of species' geographical distributions. We name these recurrent changes ''orbitally forced species' range dynamics'' (ORD). The magnitude of ORD varies in space and time. ORD decreases gradual speciation (attained by gradual changes over many generations), increases range sizes and the proportions of species formed by polyploidy and other ''abrupt'' mechanisms, selects against specialization, and favor dispersability. Large ORD produces species prone neither to extinction nor gradual speciation. ORD increases with latitude. This produces latitudinal patterns, among them the gradient in species diversity and species' range sizes (Rapoport's rule). Differential ORD and its evolutionary consequences call for new conservation strategies on the regional to global scale.
Hydrochory, or the passive dispersal of organisms by water, is an important means of propagule transport, especially for plants. During recent years, knowledge about hydrochory and its ecological consequences has increased considerably and a substantial body of literature has been produced. Here, we review this literature and define the state of the art of the discipline. A substantial proportion of species growing in or near water have propagules (fruits, seeds or vegetative units) able to disperse by water, either floating, submerged in flowing water, or with the help of floating vessels. Hydrochory can enable plants to colonize sites out of reach with other dispersal vectors, but the timing of dispersal and mechanisms of establishment are important for successful establishment. At the population level, hydrochory may increase the effective size and longevity of populations, and control their spatial configuration. Hydrochory is also an important source of species colonizing recruitment-limited riparian and wetland communities, contributing to maintenance of community species richness. Dispersal by water may even influence community composition in different landscape elements, resulting in landscape-level patterns. Genetically, hydrochory may reduce spatial aggregation of genetically related individuals, lead to high gene flow among populations, and increase genetic diversity in populations receiving many propagules. Humans have impacted hydrochory in many ways. For example, dams affect hydrochory by reducing peak flows and hence dispersal capacity, altering the timing of dispersal, and by presenting physical barriers to dispersal, with consequences for riverine plant communities. Hydrochory has been inferred to be an important vector for the spread of many invasive species, but there is also the potential for enhancing ecosystem restoration by improving or restoring water dispersal pathways. Climate change may alter the role of hydrochory by modifying the hydrology of water-bodies as well as conditions for propagule release and plant colonization.
I propose that global patterns in numbers of range-restricted endemic species are caused by variation in the amplitude of climatic change occurring on time-scales of 10-100 thousand years (Milankovitch oscillations). The smaller the climatic shifts, the more probable it is that palaeoendemics survive and that diverging gene pools persist without going extinct or merging, favouring the evolution of neoendemics. Using the change in mean annual temperature since the last glacial maximum, estimated from global circulation models, I show that the higher the temperature change in an area, the fewer endemic species of mammals, birds, reptiles, amphibians and vascular plants it harbours. This relationship was robust to variation in area (for areas greater than 10(4) km2), latitudinal position, extent of former glaciation and whether or not areas are oceanic islands. Past climatic change was a better predictor of endemism than annual temperature range in all phylads except amphibians, suggesting that Rapoport's rule (i.e. species range sizes increase with latitude) is best explained by the increase in the amplitude of climatic oscillations towards the poles. Globally, endemic-rich areas are predicted to warm less in response to greenhouse-gas emissions, but the predicted warming would cause many habitats to disappear regionally, leading to species extinctions.
▪ Abstract Variations in Earth's orbit with periods of 10–100 thousand years (kyr) (Milankovitch oscillations) have led to recurrent and rapid climatic shifts throughout Earth's history. These cause changes in the geographical distributions of clades, which we term orbitally forced range dynamics (ORD). The magnitude of ORD varies geographically, e.g., with latitude. Climatic shifts cause extinction, splitting, and merging of gene pools and clades. They select among individuals and clades for traits enhancing the ability to survive in situ and to establish new populations. There is also nonadaptive sorting caused by the large geographical variation in ORD, as only gene pools that are in the right place when climate shifts survive. ORD lead to sorting at many levels of genealogic inclusiveness. Clades that have survived climatic shifts during at least one entire period of the longest significant Milankovitch oscillations (100 kyr), we name β-clades. The products of more recent cladogenesis are α-clades, which are always nested within a β-clade. We conclude that ORD may promote α-clade formation but curb rates of β-clade formation. In areas with little ORD, where gene pools persist without going extinct or merging, clade splits and divergence may accumulate leading to high rates of β-clade formation and β-anagenesis (evolutionary change persisting >100 kyr). High ORD should lead to low numbers of β-clades, β-clades with low levels of spatial genetic divergence, little geographical subdivision and large ranges, organisms with high vagility and low specialization, high proportions of β-clades formed by polyploidization, and little β-anagenesis. We predict global and interregional geographic patterns in these variables caused by differential ORD. Thus, ORD potentially explains a wide array of patterns, suggesting ORD as a fundamental factor in evolution. The vulnerability of biotas to many human activities should vary with the magnitude of ORD.
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