The advantages and disadvantages of being introduced

Introduced species, those dispersed outside their natural ranges by humans, now cause almost all biological invasions, i.e., entry of organisms into habitats with negative effects on organisms already there. Knowing whether introduction tends to give organisms specific ecological advantages or disadvantages in their new habitats could help understand and control invasions. Even if no specific species traits are associated with introduction, introduced species might out-compete native ones just because the pool of introduced species is very large (“global competition hypothesis”). Especially in the case of intentional introduction, high initial propagule pressure might further increase the chance of establishment, and repeated introductions from different source populations might increase the fitness of introduced species through hybridization. Intentional introduction screens species for usefulness to humans and so might select for rapid growth and reproduction or carry species to suitable habitats, all which could promote invasiveness. However, trade offs between growth and tolerance might make introduced species vulnerable to extreme climatic events and cause some invasions to be transient (“reckless invader hypothesis”). Unintentional introduction may screen for species associated with human-disturbed habitats, and human disturbance of their new habitats may make these species more invasive. Introduction and natural long-distance dispersal both imply that species have neither undergone adaptation in their new habitats nor been adapted to by other species there. These two characteristics are the basis for many well-known hypotheses about invasion, including the “biotic resistance”, “enemy release”, “evolution of increased competitive ability” and “novel weapon” hypotheses, each of which has been shown to help explain some invasions. To the extent that biotic resistance depends upon local adaption by native species, altering selection pressures could reduce resistance and promote invasion (“local adaptation hypothesis”), and restoring natural regimes could reverse this effect.     

Number and weight of seeds and reproductive strategies of herbaceous plants

The paper brings data set of seed reproduction of about 500 wild herbaceous species of Central Europe and presents a number of produced seeds in a new way. Number of seeds — reproductive capacity of a population (RCP) was defined as a number of seeds produced by one species per 1 m² at its one-hundred-per-cent cover per one season. About 23% of seed weight variability and about 26% of variability in RCP were explainable on a family level. The trade-off between RCP and seed weight was confirmed both within families and also among families. Both characteristics had higher variability on a family level and on lower taxonomic levels than on subclass and class levels. Species with an annual life cycle have a larger number of seeds than perennials. Geophytes and species without lateral spread have a tendency to produce a large amount of relatively weighty seeds in comparison to other life forms, and to species with vegetative lateral spread. Species with seeds dispersed by wind usually have a large number of lightweight seeds. Multivariate analyses confirmed some tendencies of reproductive traits which correspond to the definition of C-, S-, R- strategies by Grime — “verification” of the RCP as ecological trait. The R- strategy was well distinguished from the C- and S- strategies by reproductive traits, whereas C- and S- strategies are very similar to each other. Species with insect pollination agent often correspond to C- strategy, wind agent to S- strategy, and species with self-pollination to R- strategy.     

The invasiveness of an introduced species does not predict its impact

Inconsistent use of terminology plagues the study and management of biological invasions. The term “invasive” has been used to describe inter alia (1) any introduced non-indigenous species; (2) introduced species that spread rapidly in a new region; and (3) introduced species that have harmful environmental impacts, particularly on native species. The second definition in various forms is more commonly used by ecologists, while the third definition is pervasive in policy papers and legislation. We tested the relationship between the invasiveness of an introduced species and its impact on native biodiversity. We quantified a species’ invasiveness by both its rate of establishment and its rate of spread, while its impact was assigned a categorical ranking based on the documented effects of the invader on native species populations. We found no correlations between these variables for introduced plants, mammals, fishes, invertebrates, amphibians and reptiles, suggesting that the mechanisms of invasion and impact are not strongly linked. Our results support the view that the term “invasive” should not be used to connote negative environmental impact.     

Secondary spread of invasive species: historic patterns and underlying mechanisms of the continuing invasion of the European rockweed Fucus serratus in eastern North America

Post-establishment spread of invasive species is a major determinant of their impact, but the spatial pattern and temporal rhythm of secondary spread are often poorly known or understood. Here we examine the spread of the European rockweed Fucus serratus over 1,500 km of shoreline after its initial discovery in Pictou, Nova Scotia (Canada) in 1868. Building upon earlier periodic surveys, we document the current distribution of this invader and provide a historic analysis of the invasion, including the integration of the recently-detected multiple introductions from Europe. The initial spread was rapid in the southern Gulf of St. Lawrence (6.0 km year⁻¹), likely due to both natural spread in a favorable environment and the extensive regional shipping traffic at the time which linked Pictou to several key ports in eastern Canada and northeast USA. Later spread, especially along the Atlantic coast of Nova Scotia, was slower (recently 0.9 km year⁻¹), but included several major jumps (~50–500 km), undoubtedly due to human-mediated transport. Although the spatial extent of the invasion has increased over the past 140 years, it has experienced several remarkable events: apparent local extinctions of northern satellite populations, a major retraction (100–150 km) along the northwestern limits of its continuous range, and stalled invasions at several points and times during its southward progression. Distributional data suggest that this invader is excluding congeners in shallow zones and possibly kelp species in deeper zones. Genetic data from one contact zone showed hybridization with native F. distichus but no evidence for introgression in migrating F. serratus. Hybridization and several other reproductive traits likely contribute to the competitive dominance of the invader in this environment.     

Invasion speeds for structured populations in fluctuating environments

We live in a time where climate models predict future increases in environmental variability and biological invasions are becoming increasingly frequent. A key to developing effective responses to biological invasions in increasingly variable environments will be estimates of their rates of spatial spread and the associated uncertainty of these estimates. Using stochastic, stage-structured, integrodifference equation models, we show analytically that invasion speeds are asymptotically normally distributed with a variance that decreases in time. We apply our methods to a simple juvenile–adult model with stochastic variation in reproduction and an illustrative example with published data for the perennial herb, Calathea ovandensis. These examples buttressed by additional analysis reveal that increased variability in vital rates simultaneously slow down invasions yet generate greater uncertainty about rates of spatial spread. Moreover, while temporal autocorrelations in vital rates inflate variability in invasion speeds, the effect of these autocorrelations on the average invasion speed can be positive or negative depending on life history traits and how well vital rates “remember” the past.     

Modelling invasibility in endogenously oscillating tree populations: timing of invasion matters

The timing of introduction of a new species into an ecosystem can be critical in determining the invasibility (i.e. the sensitivity to invasion) of a resident population. Here, we use an individual-based model to test how (1) the type of competition (symmetric versus asymmetric) and (2) seed masting influence the success of invasion by producing oscillatory dynamics in resident tree populations. We focus on a case where two species (one resident, one invader introduced at low density) do not differ in terms of competitive abilities. By varying the time of introduction of the invader, we show that oscillations in the resident population favour invasion, by creating “invasibility windows” during which resource is available for the invader due to transiently depressed resident population density. We discuss this result in the context of current knowledge on forest dynamics and invasions, emphasizing the importance of variability in population dynamics.     

The roles of body size and phylogeny in fast and slow life histories

Species’ life histories are often classified on a continuum from “fast” to “slow”, yet there is no consistently used definition of this continuum. For example, some researchers include body mass as one of the traits defining the continuum, others factor it out by analysing body-mass residuals, a third group performs both of these analyses and uses the terms “fast” and “slow” in both ways, while still others do not mention body mass at all. Our analysis of European and North American freshwater fish, mammals, and birds (N = 2,288 species) shows the fundamental differences between life-history patterns of raw data and of body-mass residuals. Specifically, in fish and mammals, the number of traits defining the continuum decreases if body-mass residuals are analysed. In birds, the continuum is defined by a different set of traits if body mass is factored out. Our study also exposes important dissimilarities among the three taxonomic groups analysed. For example, while mammals and birds with a “slow” life history have a low fecundity, the opposite is true for fish. We conclude that our understanding of life histories will improve if differences between patterns of raw data and of body-mass residuals are acknowledged, as well as differences among taxonomic groups, instead of using the “fast–slow continuum” too indiscriminately for any covarying traits that appear to suit the idea.     

Ecological Invasion,Roughened Fronts,and a Competitor’s Extreme Advance: Integrating Stochastic Spatial-Growth Models

Both community ecology and conservation biology seek further understanding of factors governing the advance of an invasive species. We model biological invasion as an individual-based, stochastic process on a two-dimensional landscape. An ecologically superior invader and a resident species compete for space preemptively. Our general model includes the basic contact process and a variant of the Eden model as special cases. We employ the concept of a “roughened” front to quantify effects of discreteness and stochasticity on invasion; we emphasize the probability distribution of the front-runner’s relative position. That is, we analyze the location of the most advanced invader as the extreme deviation about the front’s mean position. We find that a class of models with different assumptions about neighborhood interactions exhibits universal characteristics. That is, key features of the invasion dynamics span a class of models, independently of locally detailed demographic rules. Our results integrate theories of invasive spatial growth and generate novel hypotheses linking habitat or landscape size (length of the invading front) to invasion velocity, and to the relative position of the most advanced invader.     

Testing population-level mechanisms of invasion by a mobile vertebrate: a simple conceptual framework for salmonids in streams

Invasion biology suffers from a lack of the ability to predict the outcome of particular invasions because of reliance on verbal models and lack of rigorous experimental data at the appropriate scale. More progress is likely to be made by considering invasions as population-level phenomena and initially focusing on specific taxa or particular categories of invasions. To this end, we propose a simple conceptual framework to motivate studies of invasion by salmonids (salmon, trout, grayling, and whitefish) in streams that emphasizes population-level mechanisms affecting native species and promoting spread by the invader. Specifically, the only direct mechanisms by which the abundance of the native species can decline are through biotic interactions which cause decreased reproductive rates or survival at specific life stages, net emigration, debilitating or fatal diseases introduced by the invader, or a combination of these factors. Conversely, abundance of the invader must increase by local reproduction, high survival, net immigration, or a combination of these factors. Review of existing salmonid invasion literature suggests that future studies could be improved by using manipulative field experiments at a spatial and temporal scale appropriate to address population-level processes, characterizing how movement affects the establishment and spread of an invader, and including abiotic context in experimental designs. Using the example of brook trout (Salvelinus fontinalis) invasion into streams containing native Colorado River cutthroat trout (Oncorhynchus clarki pleuriticus) in the central Rocky Mountains (USA), we demonstrate how the framework can be used to design a manipulative field experiment to test for population-level mechanisms causing ecological effects and promoting invasion success. Experiments of this type will give invasion ecologists a useful example of how a taxon-specific invasion framework can improve the ability to predict ecological effects, and provide fishery biologists with the quantitative foundation necessary to better manage stream salmonid invasions.     

Detecting life history trade-offs: measuring energy stores in “capital” breeders reveals costs of reproduction

Life history trade-offs should be detectable as negative correlations between the relevant traits (e.g. reproductive output versus energy storage), but may be masked by variation in resource levels among individuals. One way to detect underlying trade-offs, at least in organisms that rely on stored energy for reproduction (“capital breeders”), may be to monitor an individual's energy stores before and after reproduction. We analysed energy stores and reproductive output in Eulamprus tympanum, a viviparous scincid lizard that stores energy for reproduction in its tail. One predicted trade-off (that between the size and number of offspring in a litter) is consistently observed, and is detectable with minimal information. Another predicted trade-off (that between offspring size and subsequent energy reserves) is not apparent in our data, perhaps because of constraints imposed by correlations among other traits. Finally, trade-offs between reproductive output and subsequent energy stores are evident in this species, but are only detectable with information on the extent of pre-reproductive as well as post-reproductive energy stores. For “capital breeders”, non-destructive measurement of pre- and post-reproductive energy stores may greatly enhance our ability to detect significant life history trade-offs. Received: 10 July 1996 / Accepted: 16 January 1997     

Ecological explanations for successful invasion of exotic plants

The intentional introduction of exotic species can increase the level of local biodiversity, enrich people’s material lives, and bring significant social and economic benefits that are also the symbols of human progress. However, along with the frequent intercourse among countries and regions, the frequency of uncontrolled crossregional migration of species is increased and there is a lack of scientific management strategy for the intentional introduction of exotic species. Exotic species invasion, which is behind habitat fragmentation, has become the second largest threatening factor to the maintenance of the global-scale level of biological diversity. Exotic species invasion can destroy the structure of an ecosystem, disturb the economic life of a society, and do harm to human health. In this paper, the authors review some of the ecological explanations for issues such as “what causes or mechanisms have led to the successful invasion of exotic species”, including the “ideal weeds characteristics”, “biodiversity resistance hypothesis”, “enemies release hypothesis”, “evolution of increased competitive ability hypothesis”, “niche opportunity hypothesis”, and “novel weapon hypothesis”. The authors also analyze and evaluate the background and theoretical basis of the hypotheses, providing explanations for some phenomena, as well as the deficiencies of these explanations.     

Allometric Effects of <Emphasis Type="Italic">Agriophyllum squarrosum</Emphasis> in Response to Soil Nutrients,Water, and Population Density in the Horqin Sandy Land of China

We monitored the allometric effects for greenhouse-grown Agriophyllum squarrosum plants in response to variations in population density and the availability of soil nutrients and water. Biomass allocations were size-dependent. The plasticity of roots, stems, leaves, and reproductive effort was “true” in response to changes in nutrient content. At a low level of soil minerals, plants allocated more resources to the development of roots and reproductive organs than to leaves, but data for stem allocations were consistent for tradeoffs between the effects of nutrients and plant size. The plasticities of leaf allocation and reproductive effort were “true” whereas those of root and stem allocations were “apparent” in response to fluctuations in soil water, being a function of plant size. Decreasing soil water content was associated with higher leaf allocation and lower reproductive effort. Except for this “apparent” plasticity of leaf allocation, none was detected with population density on biomass allocation. Architectural traits were determinants of the latter. For roots, the determining trait was the ratio of plant height to total biomass; for stems and reproduction, plant height; and for leaves, the ratio of branch numbers to plant height.     

Freshwater crayfish invasions: former crayfish invader Galician crayfish hands title “invasive” over to new invader spiny-cheek crayfish

Biological invasions are a major threat to global biodiversity. Invasive freshwater crayfish in that context are especially prominent for their negative effects on both ecosystem integrity and native crayfish. However, some systems may have supported a crayfish species not originally native to the system without perceivable negative consequences for the ecosystem while other invasive crayfish species may constitute a major threat to ecosystem stability. Here I present an example how two crayfish, the spiny-cheek and the Galician crayfish both by researchers and governmental agencies considered non-native differ in their threats to the native ecosystem. Whereas the spiny-cheek crayfish is a recent potentially disease-transmitting and still spreading invader with high local densities the Galician crayfish might be part of the lake’s fauna since several hundred years, appears in lower densities and is unlikely to be a vector of disease. Therefore, regardless of the Galician crayfish’s actual date of introduction it is thus a rather “old and integrated” invader, which is now being faced and itself potentially threatened by the emergence of a “new and dangerous” invader: the spiny-cheek crayfish. This also exemplifies that in the face of often insufficient scientific information about dates of species introductions care should be taken in postulating species as invasive and dangerous without any form of risk assessment for their impact on the ecosystem.     

Growth and reproduction in higher plants I. Theoretical analysis by mathematical models

To analyse the whole life of higher plants, an attempt was made to describe their growth and reproduction by mathematical models based on the elements determining matter production and economy of the matter. A plant body was regarded as a compound system of two parts; “productive part” and “reproductive part”. A parameter (reproductive index) was introduced to connect these two parts, and a set of the mathematical models describing the quantitative growth of these two parts were established. Two basic patterns of reproduction in higher plants were distinguished into “D-reproduction” and “I-reproduction”. The state of matter production of the mother plant determined an initial size of the daughter plant in theD-reproduction, while, in theI-reproduction, it did not determine the initial size of the daughter, but determined the number of propagules. The model of each reproduction pattern was also constructed. A formula determining the initial size of a plant in a given generation was constructed as the model of theD-reproduction. The model for theI-reproduction described the number of propagules produced in a given generation. Some aspects of the plant life, e.g. the optimum reproductive index, the switch-over time from the vegetative to the reproductive growth phase, the seed number, types of expansive reproduction, were theoretically analysed and discussed under these mathematical models.     

Grandmothers,hunters and human life history

This paper critiques the competing “Grandmother Hypothesis” and “Embodied Capital Theory” as evolutionary explanations of the peculiarities of human life history traits. Instead, I argue that the correct explanation for human life history probably involves elements of both hypotheses: long male developmental periods and lives probably evolved due to group selection for male hunting via increased female fertility, and female long lives due to the differential contribution women’s complex foraging skills made to their children and grandchildren’s nutritional status within groups provisioned by male hunting.     

<Emphasis Type="Italic">PRUNUS</Emphasis>: a spatially explicit demographic model to study plant invasions in stochastic,heterogeneous environments

To model the invasion of Prunus serotina invasion within a real forest landscape we built a spatially explicit, non-linear Markov chain which incorporated a stage-structured population matrix and dispersal functions. Sensitivity analyses were subsequently conducted to identify key processes controlling the spatial spread of the invader, testing the hypothesis that the landscape invasion patterns are driven in the most part by disturbance patterns, local demographical processes controlling propagule pressure, habitat suitability, and long-distance dispersal. When offspring emigration was considered as a density-dependent phenomenon, local demographic factors generated invasion patterns at larger spatial scales through three factors: adult longevity; adult fecundity; and the intensity of self-thinning during stand development. Three other factors acted at the landscape scale: habitat quality, which determined the proportion of the landscape mosaic which was potentially invasible; disturbances, which determined when suitable habitats became temporarily invasible; and the existence of long distance dispersal events, which determined how far from the existing source populations new founder populations could be created. As a flexible “all-in-one” model, PRUNUS offers perspectives for generalization to other plant invasions, and the study of interactions between key processes at multiple spatial scales.     

Coexistence and Spread of Competitors in Heterogeneous Landscapes

Competition between species is ubiquitous in nature and therefore widely studied in ecology through experiment and theory. One of the central questions is under which conditions a (rare) invader can establish itself in a landscape dominated by a resident species at carrying capacity. Applying the same question with the roles of the invader and resident reversed leads to the principle that “mutual invasibility implies coexistence.” A related but different question is how fast a locally introduced invader spreads into a landscape (with or without competing resident), provided it can invade. We explore some aspects of these questions in a deterministic, spatially explicit model for two competing species with discrete non-overlapping generations in a patchy periodic environment. We obtain threshold values for fragmentation levels and dispersal distances that allow for mutual invasion and coexistence even if the non-spatial competition model predicts competitive exclusion. We obtain exact results when dispersal is governed by a Laplace kernel. Using the average dispersal success, we develop a mathematical framework to obtain approximate results that are independent of the exact dispersal patterns, and we show numerically that these approximations are very accurate.     

Pleuroxus denticulatus (Crustacea: Anomopoda: Chydoridae) a new invader in the Danube Basin

Pleuroxus denticulatus Birge, 1879 is a new invader from Western to Central Europe. Its occurrence in a Danube inundation (Slovak region) probably corresponds with the junction of the Danube and Rhine Rivers. The species was found among ‘the hard macrophytes’ of flowing arms and artificial ponds. A large variability was recorded in the number of valval denticles and in postabdominal denticulation. It is evident that part of the N. American populations is different from European populations. P. denticulatus seems to represent two sibling species. Both are characterised by an angular distal postabdominal corner, bearing 3–4 denticles, and by valval denticles on the postero-ventral corner. This revised version was published online in July 2006 with corrections to the Cover Date.     

The QTL analysis on maternal and endosperm genome and their environmental interactions for characters of cooking quality in rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L<Emphasis Type="Italic">.</Emphasis>)

Investigations to identify quantitative trait loci (QTLs) governing cooking quality traits including amylose content, gel consistency and gelatinization temperature (expressed by the alkali spread value) were conducted using a set of 241 RIL populations derived from an elite hybrid cross of “Zhenshan 97” × “Minghui 63” and their reciprocal backcrosses BC₁F₁ and BC₂F₁ populations in two environments. QTLs and QTL × environment interactions were analyzed by using the genetic model with endosperm and maternal effects and environmental interaction effects on quantitative traits of seed in cereal crops. The results suggested that a total of seven QTLs were associated with cooking quality of rice, which were subsequently mapped to chromosomes 1, 4 and 6. Six of these QTLs were also found to have environmental interaction effects.