Influences on the transport and establishment of exotic bird species: an analysis of the parrots (Psittaciformes) of the world

Most studies of exotic species invasions only consider the factors that affect the establishment of populations following release, yet this is only one step on the invasion pathway. Different factors are likely to influence which species are transported and released. Here, we examine the influence of species traits on the successful transition of species through several stages in the introduction pathway (transport, release, and establishment), using parrots (Aves: Psittaciformes) as a model system. We use a species‐level supertree of parrots to test for phylogenetic auto‐correlation in the introduction process. Our analyses find that different sets of variables are related to the probability that a species enters each stage on the invasion pathway. The availability of individuals for transport and release seems to be most important for passage through these stages, but has no obvious effect on establishment following release. Rather, establishment success is higher for sedentary species, and species with broad diets.     

A neutral terminology to define 'invasive' species

The use of simple terms to articulate ecological concepts can confuse ideological debates and undermine management efforts. This problem is particularly acute in studies of nonindigenous species, which alternatively have been called ‘exotic’, ‘introduced’, ‘invasive’ and ‘naturalised’, among others. Attempts to redefine commonly used terminology have proven difficult because authors are often partial to particular definitions. In an attempt to form a consensus on invasion terminology, we synthesize an invasional framework based on current models that break the invasion process into a series of consecutive, obligatory stages. Unlike previous efforts, we propose a neutral terminology based on this framework. This ‘stage‐based’ terminology can be used to supplement terms with ambiguous meanings (e.g. invasive, introduced, naturalized, weedy, etc.), and thereby improve clarity of future studies. This approach is based on the concept of ‘propagule pressure’ and has the additional benefit of identifying factors affecting the success of species at each stage. Under this framework, invasions can be more objectively understood as biogeographical, rather than taxonomic, phenomena; and author preferences in the use of existing terminology can be addressed. An example of this recommended protocol might be: ‘We examined distribution data to contrast the characteristics of invasive species (stages IVa and V) and noninvasive species (stages III and IVb)’.     

Current and projected future risks of freshwater fish invasions in China

Biological invasions are a primary threat to global biodiversity, supporting mounting calls for the development of early‐warning systems to manage existing and emerging invaders. Here, we evaluated the geographical pattern of invasion risks of currently established and potentially emerging nonnative freshwater fishes in China by jointly considering the threats of introduction and establishment under climate change. Introduction threats were estimated according to proxies of human activities and propagule pressure for two primary pathways (aquaculture or ornamental). Establishment threats for 51 current and 64 potential invaders (based on whether having established or not self‐sustaining populations) were assessed using an ensemble of species distribution models under current (1960–1990) and future [2041–2060 (2050s) and 2061–2080 (2070s)] climate scenarios. Geographical patterns of invasion risk were then assessed by overlaying the threats of introduction and establishment for each species group both in present‐day and in the future. We found that eastern China displayed the highest threat of introduction. By contrast, southeastern and northwestern regions were identified as the most suitable for the establishment of both current and potential invaders. Under a changing climate, 83 out of 115 species displayed an increase in habitat suitability, resulting in an overall increase of 4.8% by 2050s and 7.1% by 2070s in the extent of suitable habitat for nonnative freshwater fishes. Taken together, invasion risk was found to be highest in southeastern China and lowest in the Tibet Plateau. Our research highlights the importance of assessing invasion risk by integrating the threats associated with the introduction and establishment stages. In particular, our findings revealed convergent patterns of invasion risk between current and potential nonnative freshwater fishes under climate change. Geographic patterns in hotspots of existing and emerging invasions provide critical insights to guide the allocation of resources to monitor and control existing and emerging invasions in China.     

Naturalization and invasion of alien plants: concepts and definitions

Abstract.  Much confusion exists in the English-language literature on plant invasions concerning the terms 'naturalized' and 'invasive' and their associated concepts. Several authors have used these terms in proposing schemes for conceptualizing the sequence of events from introduction to invasion, but often imprecisely, erroneously or in contradictory ways. This greatly complicates the formulation of robust generalizations in invasion ecology.
Based on an extensive and critical survey of the literature we defined a minimum set of key terms related to a graphic scheme which conceptualizes the naturalization/invasion process. Introduction means that the plant (or its propagule) has been transported by humans across a major geographical barrier. Naturalization starts when abiotic and biotic barriers to survival are surmounted and when various barriers to regular reproduction are overcome. Invasion further requires that introduced plants produce reproductive offspring in areas distant from sites of introduction (approximate scales: > 100 m over < 50 years for taxa spreading by seeds and other propagules; > 6 m/3 years for taxa spreading by roots, rhizomes, stolons or creeping stems). Taxa that can cope with the abiotic environment and biota in the general area may invade disturbed, seminatural communities. Invasion of successionally mature, undisturbed communities usually requires that the alien taxon overcomes a different category of barriers.
We propose that the term 'invasive' should be used without any inference to environmental or economic impact. Terms like 'pests' and 'weeds' are suitable labels for the 50–80% of invaders that have harmful effects. About 10% of invasive plants that change the character, condition, form, or nature of ecosystems over substantial areas may be termed 'transformers'.     

Herbivores and the success of exotic plants: a phylogenetically controlled experiment

In a field experiment with 30 locally occurring old‐field plant species grown in a common garden, we found that non‐native plants suffer levels of attack (leaf herbivory) equal to or greater than levels suffered by congeneric native plants. This phylogenetically controlled analysis is in striking contrast to the recent findings from surveys of exotic organisms, and suggests that even if ‘enemy release’ does accompany the invasion process, this may not be an important mechanism of invasion, particularly for plants with close relatives in the recipient flora.     

外来种隐蔽入侵： 类型及影响

隐蔽入侵（cryptic invasion）是指在形态上与土著种（或早期建群种）不能或难以区分的外来种在人们未觉察的状态下成功入侵的过程。人们对这类入侵方式往往视而不觉。本文综述了外来种隐蔽入侵的类型以及生态影响。隐蔽入侵的类型主要包括外来姊妹种形式的隐蔽入侵、 不同遗传支系的隐蔽入侵、 不同地理种群的隐蔽入侵以及“返传入”。其中, “返传入”目前还是一种假说。由于这类入侵外来种更容易与土著种（或早期建群种）杂交或基因渗透, 因而可对入侵种自身或土著种产生深远的生态影响。鉴于隐蔽入侵现象的广泛性, 建议进一步加强该方面的研究。     

Measuring floristic homogenization by non-native plants in North America

Aim To determine if non‐native plant species are homogenizing species composition among widely dispersed plant communities. Location Twenty localities in North America. Methods Species lists among localities were compared to measure the influence of non‐native species richness at each locality on the Jaccard Index (JI) of similarity between localities. Results After removing the effects of distance, because shared native species decreased with distance, three (nonexclusive) lines of evidence indicate that non‐native species promote homogenization. First, pairs of sites with a high combined total of non‐native species tend to have higher similarity than those with a low total of non‐natives. Second, for a given distance, more non‐native than native species tended to be shared among localities. Third, whereas most of the site comparisons with high total non‐native richness have a non‐native/native JI ratio greater than 1 (often much greater), only half of the comparisons with low total non‐native richness have a ratio greater than one. Main conclusions These findings provide quantitative support for the widely held, but rarely tested, notion that non‐native species tend to homogenize biological communities because they are more commonly shared among communities. Such testing is important as non‐native species could theoretically have no impact or even reduce homogenization among communities, if non‐native colonizers consist of different species pools.     

Plant invasion across space and time: factors affecting nonindigenous species success during four stages of invasion

Invasive nonindigenous plant species (NIPS) threaten native diversity, alter ecosystem processes, and may interact with other components of global environmental change. Here, a general framework is outlined that attempts to connect patterns of plant invasion to processes underlying these patterns at four well-established spatio-temporal stages of the invasion process: transport, colonization, establishment, and landscape spread. At each stage we organize findings and ideas about the filters that limit NIPS success and the interaction of these filters with historical aspects of introduction events, NIPS traits, and ecosystem properties. While it remains difficult to draw conclusions about the risk of invasion across ecosystems, to delineate universal 'invader traits', or to predict large-scale extinctions following invasions, this review highlights the growing body of research that suggests that the success of invasive NIPS is controlled by a series of key processes or filters. These filters are common to all invasion events, and will interact throughout the stages of plant invasion, although the relative importance of a filter may be stage, species or location specific. It is suggested that both research and management programs may benefit from employing multiscale and stage approaches to studying and controlling invasion. We further use the framework to briefly examine potential interactions between climate change and filters that limit NIPS invasion.     

Propagule Pressure: A Null Model for Biological Invasions

Invasion ecology has been criticised for its lack of general principles. To explore this criticism, we conducted a meta-analysis that examined characteristics of invasiveness (i.e. the ability of species to establish in, spread to, or become abundant in novel communities) and invasibility (i.e. the susceptibility of habitats to the establishment or proliferation of invaders). There were few consistencies among invasiveness characteristics (3 of 13): established and abundant invaders generally occupy similar habitats as native species, while abundant species tend to be less affected by enemies; germination success and reproductive output were significantly positively associated with invasiveness when results from both stages (establishment/spread and abundance/impact) were combined. Two of six invasibility characteristics were also significant: communities experiencing more disturbance and with higher resource availability sustained greater establishment and proliferation of invaders. We also found that even though ‘propagule pressure’ was considered in only ~29% of studies, it was a significant predictor of both invasiveness and invasibility (55 of 64 total cases). Given that nonindigenous species are likely introduced non-randomly, we contend that ‘propagule biases’ may confound current paradigms in invasion ecology. Examples of patterns that could be confounded by propagule biases include characteristics of good invaders and susceptible habitats, release from enemies, evolution of ‘invasiveness’, and invasional meltdown. We conclude that propagule pressure should serve as the basis of a null model for studies of biological invasions when inferring process from patterns of invasion. An erratum to this article can be found at     

Habitat niche breadth predicts invasiveness in solitary ascidians

A major focus of invasion biology is understanding the traits associated with introduction success. Most studies assess these traits in the invaded region, while only few compare nonindigenous species to the pool of potential invaders in their native region. We focused on the niche breadth hypothesis, commonly evoked but seldom tested, which states that generalist species are more likely to become introduced as they are capable of thriving under a wide set of conditions. Based on the massive introduction of tropical species into the Mediterranean via the Suez Canal (Lessepsian migration), we defined ascidians in the Red Sea as the pool of potential invaders. We constructed unique settlement plates, each representing six different niches, to assess ascidian niche breadth, and deployed them in similar habitats in the native and invaded regions. For each species found on plates, we evaluated its abundance, relative abundance across successional stages, and niche breadth, and then compared (1) species in the Red Sea known to have been introduced into the Mediterranean (Lessepsian species) and those not known from the Mediterranean (non‐Lessepsian); and (2) nonindigenous and indigenous species in the Mediterranean. Lessepsian species identified on plates in the Red Sea demonstrated wider niche breadth than non‐Lessepsian species, supporting the niche breadth hypothesis within the native region. No differences were found between Lessepsian and non‐Lessepsian species in species abundance and successional stages. In the Mediterranean, nonindigenous species numerically dominated the settlement plates. This precluded robust comparisons of niche breadth between nonindigenous and indigenous species in the invaded region. In conclusion, using Red Sea ascidians as the pool of potential invaders, we found clear evidence supporting the niche breadth hypothesis in the native region. We suggest that such patterns may often be obscured when conducting trait‐based studies in the invaded regions alone. Our findings indicate that quantifying the niche breadth of species in their native regions will improve estimates of invasiveness potential.     

Estuarine and scalar patterns of invasion in the soft-bottom benthic communities of the San Francisco Estuary

The spatial patterns of nonindigenous species in seven subtidal soft-bottom communities in the San Francisco Estuary were quantified. Sixty nonindigenous species were found out of the 533 taxa enumerated (11%). Patterns of invasion across the communities were evaluated using a suite of invasion metrics based on the abundance or species richness of nonindigenous species. Patterns of invasion along the estuarine gradient varied with the invasion metric used, and the ecological interpretation of the metrics is discussed. Overall, the estuarine transition community located in the estuarine turbidity maximum zone (mean 5 practical salinity unit (psu)), main estuarine community (mean 16 psu), and marine muddy community (mean 28 psu) were more invaded than two fresh-brackish communities (mean < 1 psu) and a marine sandy community (mean 27 psu). Nonindigenous species were numerically dominant over much of the Estuary, making up more than 90% of the individuals in two communities. The percentage of the total species composed of nonindigenous species increased at smaller spatial scales: 11% at the estuary (gamma) scale, 21% at the community (alpha) scale, and 42% at the grab (point) scale. Wider spatial distributions of nonindigenous species and a relatively greater percentage of rare native species may have resulted in this pattern. Because of this scale dependency, comparisons among sites need to be made at the same spatial scale. Native species were positively correlated with nonindigenous species in several of the communities, presumably due to similar responses to small-scale differences in habitat quality. The rate of invasion into the soft-bottom communities of the San Francisco Estuary appears to have increased over the last one to two decades and many of the new introductions have become numerically dominant.     

Invasive Australian crayfish Cherax quadricarinatus in the Sanyati Basin of Lake Kariba: a preliminary survey

The invasion of Cherax quadricarinatus, the Australian redclaw crayfish, in the Sanyati Basin of Lake Kariba, Zimbabwe, is reported. A total of 79 crayfish were caught at 10 out of 12 sampling sites in the Sanyati Basin in November–December 2012. The average catch per unit effort (CPUE) varied from 1.1 to 4 crayfish per trap per night, carapace length ranged from 29 to 93.5 mm, and weight ranged from 4 to 196.2 g. Most crayfish were between 50 and 59.9 mm carapace length. Males (average 82.6 g) were significantly heavier than females (37.2 g) and males were larger in carapace length, carapace width, chela length and chela width. A feral population of C. quadricarinatus is now established in the Sanyati Basin. Possible modes of dispersal within the basin are discussed.     

Studying invasion: have we missed the boat?

Invasive species, and the ensuing homogenization of the world's biota, form a global problem with consequences ranging from the decline and extirpation of native species to threats to human health. The magnitude of this issue demands a thorough understanding of the invasion process, which consists of three main stages: initial dispersal, establishment of self‐sustaining populations, and spread. To assess the relative distribution of research effort among these stages, we conducted a literature review using 873 articles published in 23 major journals over the past 10 years. Of the 873 papers, only 96 (11.0%) studied initial dispersal, and only half of these (6.2% of the total) were empirical. As the first stage in a contingent process, we argue that initial dispersal is the best stage during which to direct management efforts. In addition, initial dispersal has direct relevance for fundamental ecological questions regarding community assembly and metacommunity dynamics. In so far that answering these questions and preventing invasion are goals of ecologists, the disparity in research effort noted here suggests that ecologists need to expand their efforts to include more research on initial dispersal.     

Secondary invasion: When invasion success is contingent on other invaders altering the properties of recipient ecosystems

Positive interactions between exotic species may increase ecosystem‐level impacts and potentially facilitate the entry and spread of other exotic species. Invader‐facilitated invasion success—”secondary invasion”—is a key conceptual aspect of the well‐known invasional meltdown hypothesis, but remains poorly defined and empirically underexplored. Drawing from heuristic models and published empirical studies, we explore this form of “secondary invasion” and discuss the phenomenon within the recognized conceptual framework of the determinants of invasion success. The term “secondary invasion” has been used haphazardly in the literature to refer to multiple invasion phenomena, most of which have other more accepted titles. Our usage of the term secondary invasion is akin to “invader‐facilitated invasion,” which we define as the phenomenon in which the invasion success of one exotic species is contingent on the presence, influence, and impacts of one or more other exotic species. We present case studies of secondary invasion whereby primary invaders facilitate the entry or establishment of exotic species into communities where they were previously excluded from becoming invasive. Our synthesis, discussion, and conceptual framework of this type of secondary invasion provides a useful reference to better explain how invasive species can alter key properties of recipient ecosystems that can ultimately determine the invasion success of other species. This study increases our appreciation for complex interactions following invasion and highlights the impacts of invasive species themselves as possible determinants of invasion success. We anticipate that highlighting “secondary invasion” in this way will enable studies reporting similar phenomena to be identified and linked through consistent terminology.     

Fish communities of estuarine salt marshes of eastern North America,and comparisons with temperate estuaries of other continents

Twenty eastern North American estuarine/saltmarsh locations, for which publishedinventories of inhabiting fishes wereavailable, were selected for study. Thegeographic range of systems extended from southTexas on the west; North River, FL. on thesouth; to Prince Edward Island, Canada on theeast; and James and Hudson Bays, Canada at thenorth. A total of 237(±) species offishes were associated with these systems. Lifehistory groupings included: permanent residents(9.3%); marine nursery species (17.7%);diadromous fishes (5.5%); marine transients(52.3%); and freshwater transients (15.2%).The most widely distributed species wereprimarily permanent residents and marinenursery species. The fraction of residentspecies did not vary significantly over thelatitudinal range, but the fraction of marinemigratory (MN + MT) species decreased withlatitude. An increasing fraction of diadromous(anadromous) species may compensate for thedecreasing fraction of marine migratoryspecies. Permanent residents, marine nurseryspecies, and marine transients all showedextended periods of spawning. Permanentresident species showed the widest ranges ofambient salinity tolerances or of ambientsalinity ranges of occupied habitats, withmarine nursery species second. Patterns withrespect to ambient temperature tolerances wereassociated with geographic ranges rather thanlife history groups. A general profile wasproduced of characteristics of species thatlive as permanent residents in salt marshestuaries of eastern North America.Species and family relationships of fishes froma group of 17 and another of 25 estuarieslocated along the west coast of Europe werecompared with the eastern North American group.European estuaries showed higher speciesrichness per system than did those of easternNorth America. Family representations ofeastern North American and western Europeanestuaries were compared with a series ofestuaries located in southwestern Australia andSouth Africa. This showed significant overlapin family representation, with two speciesbeing common among the four continents.