Modeling biological invasions into periodically fragmented environments

Range expansion of a single species in a regularly striped environment is studied by using an extended Fisher model, in which the rates of diffusion and reproduction periodically fluctuate between favorable and unfavorable habitats. The model is analyzed for two initial conditions: the initial population density is concentrated on a straight line or at the origin. For each case, we derive a mathematical formula which characterizes the spatio-temporal pattern of range expansion. When initial distribution starts from a straight line, it evolves to a traveling periodic wave (TPW), whose frontal speed is analytically determinable. When the range starts from the origin, it tends to expand radially at a constant average speed in each direction (ray speed) keeping its frontal envelope in a similar shape. By examining the relation between the ray speed and the TPW speed, we derive the ray speed in a parametric form, from which the envelope of the expanding range can be predicted. Thus we analyze how the pattern and speed of the range expansion are affected by the pattern and scale of fragmentation, and the qualities of favorable and unfavorable habitats. The major results include: (1). The envelope of the expanding range show a variety of patterns, nearly circular, oval-like, spindle-like, depending on parameter values; (2). All these patterns are elongated in the direction of stripes; (3). When the scale of fragmentation is enlarged without changing the relative spatial pattern, the ray speed in any direction increases, i.e., the rate of range expansion increases.     

Deterministic genetic models in varying environments

Summary J. B. S. Haldane and S. D. Jayakar [J. Genet. 58, 237–242 (1963)] argue that, when genotype fitnesses fluctuate from generation to generation, if the geometric and arithmetic means of the fitnesses satisfy certain inequalities, there will be a protected polymorphism. Their assertions are biologically interesting, but their mathematical analysis is not sufficient to support their conclusions. We present a firm mathematical analysis and several examples that demonstrate the need for stronger hypotheses and, in some cases, weaker conclusions.Journal Paper No. J-10136 of the Iowa Agriculture and Home Economics Experiment Station, Ames, Iowa. Project 1669. Partial support by National Institutes of Health, Grant GM 13827     

Establishment versus population growth in spatio-temporally varying environments

We consider situations where repeated invasion attempts occur from a source population into a receptor population over extended periods of time. The receptor population contains two locations that provide different expected offspring numbers to invaders. There is demographic stochasticity in offspring numbers. In addition, temporal variation causes local invader fitnesses to vary. We show that effects of environmental autocorrelation on establishment success depend on spatial covariance of the receptor subpopulations. In situations with a low spatial covariance this effect is positive, whereas high spatial covariance and/or high migration probabilities between the subpopulations causes the effect to be negative. This result reconciles seemingly contradictory results from the literature concerning effects of temporal variation on population dynamics with demographic stochasticity. We study an example in the context of genetic introgression, where invasions of cultivar plant genes occur through pollen flow from a source population into wild-type receptor populations, but our results have implications in a wider range of contexts, such as the spread of exotic species, metapopulation dynamics and epidemics.     

Little evidence of invasion by alien conifers in Europe

Aim Conifers are invasive species in many parts of the world, especially in the Southern Hemisphere. There are many introduced conifers in Europe, but their status as alien species is poorly documented. We conducted a comprehensive literature review to ascertain the extent to which alien conifers can be considered invasive. Location Europe. Methods We reviewed the historical record of alien conifer invasion in Europe (i.e. species with a native range outside the continental boundaries of Europe) by screening the DAISIE database and the ISI Web of Science. Results According to DAISIE, there are 54 alien conifer species in Europe. Pseudotsuga menziesii is the species recorded as naturalized in the most countries (12) and the UK is the country with the most naturalized species (18). Thirty‐seven of these conifers have been studied, to some extent, in a total of 131 papers (212 records). Nevertheless, only a few papers have investigated aspects related to biological invasions. In fact, the species are not referred to as alien by the authors in more than half of the papers (66%). Twenty‐five per cent of the papers have investigated plant traits, 46% are about biotic and abiotic factors influencing tree performance and 29% deal with ecological and economic impacts. Most papers are related to entomology, dealing with natural enemies affecting the alien conifers. Main conclusions Scientists have not yet perceived alien conifers in Europe as problematic species. Moreover, the low introduction effort, long lag‐time since plantation and phylogenetic closeness between alien and native conifers are possible reasons for their low expansion in Europe to date. From a management point of view, careful observations of sites with alien conifers is necessary to watch for new invasions. From a scientific perspective, thorough analyses of the extent that introduction, rates of naturalization and biogeographical differences influence invasive spread between the two hemispheres will prove timely.     

Patterns of woody plant invasion in an Argentinean coastal grassland

Coastal dune grasslands are fragile ecosystems that have historically been subjected to various types of uses and human activities. In Buenos Aires Province (Argentina), these areas are frequently afforested for urban and touristic development. The introduction and subsequent spread of exotic tree species is one of the main threats to conservation of natural grasslands as invasive trees strongly transform their structure and composition. The aim of this study was to identify patterns of woody plant invasion comparing plant communities and environmental variables between invaded and non-invaded areas surrounding the coastal village of Mar Azul, Argentina. Coastal grasslands in this area are being invaded by Populus alba (white poplar) and Acacia longifolia (coast wattle). The height of the saplings and the richness of the accompanying vegetation were evaluated in relation to the distance from the edge of the mature tree patches. Also, the cover, richness and diversity of all species in the invaded and non-invaded areas were measured, as well as soil pH, temperature and particle size. Negative correlations were found between the height of the saplings and distance to mature tree patches in all areas. The richness of the accompanying vegetation was negatively and positively correlated with the distance from the poplar and acacia area, respectively. The most abundant native species was Cortaderia selloana. Less cover, richness and diversity of native plant species and greater soil particle size were found in invaded areas, where the proportion of bare soil was higher. Also, a higher proportion of leaf litter in the invaded areas was registered. The results emphasize the invasive capacity of P. alba and A. longifolia advancing on the native communities and reducing their richness. Knowledge of the impact of invasive woody plants in coastal grasslands is important to design active management strategies for conservation purposes.     

Landscape-scale impacts of transportation infrastructure on spatial dynamics of two vulnerable ungulate species in Ghamishloo Wildlife Refuge,Iran

Development of roads through protected areas can have deleterious effects on natural habitats containing species of conservation concern. During the past decades road construction has affected many former remote areas and led to fragmentation and isolation of wildlife populations. The present study focuses on the ecological impacts of Isfahan's West Freeway, which passes through Ghamishloo Wildlife Refuge; an IUCN category IV protected area, in Isfahan Province. The two key affected species, both classified as vulnerable by IUCN, the goitered gazelle (Gazella subgutturosa subgutturosa) and the wild sheep (Ovis orientalis isphahanica) were subject to impact analyses. We used habitat evaluation procedure (HEP) as a habitat-based impact assessment methodology which considers habitat quality and quantity. Habitat quality was measured as habitat suitability index (HSI) for each species. By literature review and field observations, five variables defining habitat suitability were identified and suitability maps for both species generated. Habitat units (HUs) were derived from multiplying the HSI for each species by the habitat area before and after freeway construction. The results showed that due to the construction of the freeway, about 14% of the HUs for goitered gazelle and about 9% of the HUs for wild sheep were lost. In addition, for quantifying landscape pattern change due to freeway construction, various landscape metrics were calculated for the species distribution polygons for two times before and after freeway construction. Results obtained through quantifying landscape metrics showed that mean nearest neighbor (MNN) and number of patches (NP) metrics increased. On the other hand, CONTAG metric decreased in both goitered gazelle and wild sheep distribution polygons, demonstrating the negative effect of freeway on these species distribution polygons integrity. According to the results of this study, mitigation and compensation activities should be considered in Ghamishloo Wildlife Refuge. Our study demonstrated that HEP method combined with quantifying landscape metrics might provide a powerful tool for assessing ecological impact of technical infrastructures on populations of far-ranging species of conservation concern.     

Predicting invasion dynamics of four alien Pinus species in a highly fragmented semi-arid shrubland in South Africa

This study explored the determinants of spread of four alien Pinus species and the ability of models to predict invasion dynamics in a complex fragmented landscape. The role of environmental factors, natural and anthropogenic disturbance in relation to invasion history was assessed for different stages in the invasion process using a Geographic Information System. Pines escaped from plantations over the past 30 years and spread into the natural semi-arid shrubland (renosterveld). The pattern of spread was compared with a simulated random distribution using two different techniques, a standard logistic regression, and a new recursive modelling approach (Formal Inference-based Recursive Modelling; FIRM). FIRM analysis improved the accuracy of predictions and revealed interactive effects of variables hidden by the logistic regression analysis. More than 80% of isolated pine individuals were found in 20% of the habitat classified as suitable by the models. Soil pH was the most important predictor for the distribution of isolated trees, whereas the establishment of dense pine stands was largely determined by fire history. Differences in invasive behaviour could be explained by species attributes such as limited dispersal for P. canariensis, and better drought-tolerance for P. halepensis. Sixty-five percent of the current pine distribution was accurately predicted by the spatial distribution of the first trees to have invaded. Such models could be used to predict potential spread of invasive plants and gain a better understanding of the main factors driving the invasion process. However, the spread of invasive species in fragmented landscapes, strongly modified by human activities, is very complicated, and the spread remains difficult to predict in the long term. The dynamics of invasion are discussed in relation to changes in land use and disturbance regime.     

Spatial patterns in insect community composition in coldwater springs

The insects living in 27 coldwater springs in southern Ontario, Canada were analyzed using several permutation and ordination methods, in order to detect patterns in the composition of their communities. These patterns were compared against simulations using artificial landscapes. A Moving Window Mantel Correlogram indicated the presence of both a strong local spatial aggregation and a global trend toward higher dissimilarity of sites with increasing geographical distance, especially beyond 50 km. Ordination plots showed a clear global gradient from a southern group of springs to a northern group (likely related to temperature and pH), which dominated the global, total, and local plots in both a SCA (Spatial CA) and a SPCA (Spatial PCA) analysis. However, the results of the local analysis differed markedly between the two methods: while the local plot for the SCA analysis indicated considerable variation among sites belonging to the same spatial group (thought to be related to discharge and random colonization episodes) and showed a pattern that was different from both the global and total analysis plots, the local SPCA plot showed little within-group variation and resembled the global and total plots.     

Marine invasion history and vector analysis of California: a hotspot for western North America

Aim We examine the regional dominance of California as a beachhead for marine biological invasions in western North America and assess the relative contribution of different transfer mechanisms to invasions over time. Location Western North America (California to Alaska, excluding Mexico). Methods We undertook extensive analysis of literature and collections records to characterize the invasion history of non‐native species (invertebrates, microalgae and microorganisms) with established populations in coastal marine (tidal) waters of western North America through 2006. Using these data, we estimated (1) the proportion of first regional records of non‐native species that occurred in California and (2) the relative contribution of transfer mechanisms to California invasions (or vector strength) over time. Results Excluding vascular plants and vertebrates, we identified 290 non‐native marine species with established populations in western North America, and 79% had first regional records from California. Many (40–64%) of the non‐native species in adjacent states and provinces were first reported in California, suggesting northward spread. California also drives the increasing regional rate of detected invasions. Of 257 non‐native species established in California, 59% had first regional records in San Francisco Bay; 57% are known from multiple estuaries, suggesting secondary spread; and a majority were attributed to vessels (ballast water or hull fouling) or oysters, in some combination, but their relative contributions are not clear. For California, more than one vector was possible for 56% of species, and the potential contribution of ballast water, hull fouling and live trade increased over time, unlike other vectors. Main conclusions California, especially San Francisco Bay, plays a pivotal role for marine invasion dynamics for western North America, providing an entry point from which many species spread. This pattern is associated historically with high propagule supply and salinity. Any effective strategies to minimize new invasions throughout this region must (1) focus attention on California and (2) address current uncertainty and future shifts in vector strength.     

The more you introduce the more you get: the role of colonization pressure and propagule pressure in invasion ecology

Aim  We argue that 'propagule pressure', a key term in invasion biology, has been attributed at least three distinct definitions (with usage of a related term causing additional confusion). All of the definitions refer to fundamental concepts within the invasion process, with the result that the distinct importance of these different concepts has been at best diluted, and at worst lost.
Location  Global.
Methods  We reviewed pertinent literature on propagule pressure to resolve confusion about different uses of the term 'propagule pressure' and we introduced a new term for one variant, colonization pressure. We conducted a computer simulation whereby the introduction of species is represented as a simple sampling process to elucidate the relationship between propagule and colonization pressure.
Results  We defined colonization pressure as the number of species introduced or released to a single location, some of which will go on to establish a self-sustaining population and some of which will not. We subsequently argued that colonization pressure should serve as a null hypothesis for understanding temporal or spatial differences in exotic species richness, as the more species that are introduced, the more we should expect to establish. Finally, using a simple simulation, we showed that propagule pressure is related to colonization pressure, but in a non-linear manner.
Main conclusion  We suggest that the nature of the relationship between propagule pressure and colonization pressure, as well as the efficacy of various proxy measures of each, require more detailed exploration if invasion ecology is to continue to develop into a more predictive science.     

A simple model for the dynamics of a host-parasite-hyperparasite interaction

Hyperparasites can play a crucial role in the control of a host-parasite interaction if they are successfully established in the community. We investigated the specific traits of the hyperparasite and those of the release event which allow a successful regulation of primary parasite populations. This study has been motivated by the case study of chestnut-Cryphonectria parasitica-Cryphonectria Hypovirus interaction. We use a model of SIR/SIS type which assumes a limited diffusion of the parasite. Our model emphasizes the thresholds for invasion linked to the ecological specificities of both the pathogen and the hyperparasite (transmission rates and virulence) and to the initial conditions of the system (population sizes of the different categories). The predictions are consistent with data on the observed spread of the virus. "Mild" strains of the hyperparasite, characterized by a high vertical transmission rate and low virulence, are more prone to establish than "severe" strains. It also demonstrates that the horizontal transmission of the virus, which is controlled by a vegetative incompatibility system in the fungus, is not the unique constraint for the virus establishment. This study may contribute to theoretical and practical aspects of the biological control of plant diseases with a hyperparasite and to the ecology of biological invasions.     

Mathematical analysis of the optimal habitat configurations for species persistence

We study a reaction-diffusion model in a binary environment made of habitat and non-habitat regions. Environmental heterogeneity is expressed through the species intrinsic growth rate coefficient. It was known that, for a fixed habitat abundance, species survival depends on habitat arrangements. Our goal is to describe the spatial configurations of habitat that maximise the chances of survival. Through numerical computations, we find that they are of two main types - ball-shaped or stripe-shaped. We formally prove that these optimal shapes depend on the habitat abundance and on the amplitude of the growth rate coefficient. We deduce from these observations that the optimal shape of the habitat realises a compromise between reducing the detrimental habitat edge effects and taking advantage of the domain boundary effects. In the case of an infinite-periodic environment, we prove that the optimal habitat shapes can be deduced from those in the case of a bounded domain.     

Reducing redundancy in invasion ecology by integrating hypotheses into a single theoretical framework

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.     

Population and evolutionary dynamics in spatially structured seasonally varying environments

Increasingly imperative objectives in ecology are to understand and forecast population dynamic and evolutionary responses to seasonal environmental variation and change. Such population and evolutionary dynamics result from immediate and lagged responses of all key life‐history traits, and resulting demographic rates that affect population growth rate, to seasonal environmental conditions and population density. However, existing population dynamic and eco‐evolutionary theory and models have not yet fully encompassed within‐individual and among‐individual variation, covariation, structure and heterogeneity, and ongoing evolution, in a critical life‐history trait that allows individuals to respond to seasonal environmental conditions: seasonal migration. Meanwhile, empirical studies aided by new animal‐tracking technologies are increasingly demonstrating substantial within‐population variation in the occurrence and form of migration versus year‐round residence, generating diverse forms of ‘partial migration’ spanning diverse species, habitats and spatial scales. Such partially migratory systems form a continuum between the extreme scenarios of full migration and full year‐round residence, and are commonplace in nature. Here, we first review basic scenarios of partial migration and associated models designed to identify conditions that facilitate the maintenance of migratory polymorphism. We highlight that such models have been fundamental to the development of partial migration theory, but are spatially and demographically simplistic compared to the rich bodies of population dynamic theory and models that consider spatially structured populations with dispersal but no migration, or consider populations experiencing strong seasonality and full obligate migration. Second, to provide an overarching conceptual framework for spatio‐temporal population dynamics, we define a ‘partially migratory meta‐population’ system as a spatially structured set of locations that can be occupied by different sets of resident and migrant individuals in different seasons, and where locations that can support reproduction can also be linked by dispersal. We outline key forms of within‐individual and among‐individual variation and structure in migration that could arise within such systems and interact with variation in individual survival, reproduction and dispersal to create complex population dynamics and evolutionary responses across locations, seasons, years and generations. Third, we review approaches by which population dynamic and eco‐evolutionary models could be developed to test hypotheses regarding the dynamics and persistence of partially migratory meta‐populations given diverse forms of seasonal environmental variation and change, and to forecast system‐specific dynamics. To demonstrate one such approach, we use an evolutionary individual‐based model to illustrate that multiple forms of partial migration can readily co‐exist in a simple spatially structured landscape. Finally, we summarise recent empirical studies that demonstrate key components of demographic structure in partial migration, and demonstrate diverse associations with reproduction and survival. We thereby identify key theoretical and empirical knowledge gaps that remain, and consider multiple complementary approaches by which these gaps can be filled in order to elucidate population dynamic and eco‐evolutionary responses to spatio‐temporal seasonal environmental variation and change.     

Population-specific demography and invasion potential in medfly

Biological invasions are constantly gaining recognition as a significant component of global change. The Mediterranean fruit fly (medfly) constitutes an ideal model species for the study of biological invasions due to its (1) almost cosmopolitan geographic distribution, (2) huge economic importance, and (3) well-documented invasion history. Under a common garden experimental set up, we tested the hypothesis that medfly populations obtained from six global regions [Africa (Kenya), Pacific (Hawaii), Central America (Guatemala), South America (Brazil), Extra-Mediterranean (Portugal), and Mediterranean (Greece)] have diverged in important immature life-history traits such as preadult survival and developmental times. We also tested the hypothesis that medfly populations from the above regions exhibit different population growth rates. For this purpose, data on the life history of immatures were combined with adult survival and reproduction data derived from an earlier study in order to calculate population parameters for the above six populations. Our results clearly show that medfly populations worldwide exhibit significant differences in preadult survival, developmental rates of immatures and important population parameters such as the intrinsic rate of increase. Therefore, geographically isolated medfly populations may share different invasion potential, since population growth rates could influence basic population processes that operate mostly during the last two stages of an invasion event, such as establishment and spread. Our findings provide valuable information for designing population suppression measures and managing invasiveness of medfly populations worldwide.     

Seasonal changes and population dynamics of the ctenophore Mnemiopsis leidyi after its first year of invasion in the Kiel Fjord, Western Baltic Sea

We analyzed the seasonal variations of the ctenophore Mnemiopsis leidyi weekly collected since its first record in the western Baltic Sea in October 2006. The distribution pattern together with the seasonal dynamics and population outbreaks in late summer 2007 indicate recent successfully establishment of M. leidyi in this area. Seasonal changes showed two periods of high reproductive activity characterized by a population structure dominated by small size classes, followed by an increase of larger ones. These results further revealed that the bulk of the population remains in deep layers during the periods of low population density, whereas it appeared situated in upper layers during the proliferation of the species. We further emphasized the strength of the population outbreaks, which can reach abundances >10-fold higher in time periods shorter than a week. The predatory impact this species may have in pelagic ecosystems warns on the importance of its recent range of expansion.     

The effect of forest fragments on abundance,diversity and species composition of predatory ants in sugarcane fields

Habitat loss and fragmentation have gradually caused loss of diversity and consequently the decline of ecological services. This study aimed to evaluate the effect of tropical forest fragments as natural habitats (river valley fragments and plateau fragments) on the community of predatory and omnivorous ants in nearby sugarcane fields. Twenty fields adjacent to these fragments were selected and evaluated one (dry season) and four months (rainy season) after harvest. In each field, ants were sampled in five linear plots (10 m inside the fragment, 0 m (field path between field and fragment), 5 m, 50 m and 100 m inside the crop fields). Each plot comprised ten sardine baits in a row parallel to the field edge. Species richness and frequency of ant species decreased with increasing distance from the forest fragments. Inside fields, species richness and frequency were higher during the period of vegetative growth (rainy season) than after harvest (dry season). Ant communities of sugarcane fields and forest fragments were more similar later in the season than directly after sugarcane harvest suggesting recolonization of the fields from the fragments. Several ant species were limited to forest fragments after harvest but occurred later in the season also in sugarcane fields confirming the potential contribution of fragments to the recolonization process and therefore to biological control of sugarcane-dominated pest insects.