Non-native Species and Rates of Spread: Lessons from the Brackish Baltic Sea

The Baltic Sea, a semi-enclosed brackish water region, has been inoculated by non-indigenous species for centuries. Today, much of its biological diversity is of foreign origin (i.e. xenodiversity), intentionally or unintentionally moved by humans over ecological and geographical barriers. As many as 98 introduced species have been recorded in the Baltic Sea and Kattegat. The role and abundance of much of the unique native brackish water fauna of the Baltic Sea are threatened by these non-indigenous species. The rate of primary introductions into the Baltic has increased since the 1950s; the secondary rate of spread of non-indigenous species within the basin varies from 30–480 km/year. We review here the invasion histories of the brown alga Sargassum muticum (introduced in the early 1990s), the mud snail Potamopyrgus antipodarum (1887), the barnacle Balanus improvisus (1844), the polychaetes Marenzelleria viridis (1985) and Polydora redeki (1963), the cladoceran Cercopagis pengoi (1992) and the mysid shrimp Hemimysis anomala (1962). This revised version was published online in July 2006 with corrections to the Cover Date.     

Human Agency in Biological Invasions: Secondary Releases Foster Naturalisation and Population Expansion of Alien Plant Species

The human mediation of biological invasions is still an underestimated phenomenon. This paper attempts to show that introductions on varying spatial scales may strongly foster invasions throughout the whole invasion process. As shown by data from central Europe, invasions frequently result from an interplay of biological and anthropogenic mechanisms. The latter, however, cannot be explained nor predicted by ecological rules. This may be an important reason for the limited predictability of invasions. Initial introductions from a donor to a new range are here distinguished from following secondary releases within the new range. The rate of naturalisation is higher in deliberately introduced plants as compared to accidental introductions. Due to higher numbers of accidental introductions, such species contribute significantly to the pool of naturalised species. Secondary releases of alien species are frequently made over long periods subsequent to the initial introduction. They may mimic demographic and dispersal processes that lead to population growth and range expansion. They also offer a pathway to overcome spatial isolation in species whose propagules are not naturally moved long distances. This even holds for most of Germany's noxious alien plant species. Secondary releases may thus promote invasions even beyond the threshold of naturalisation. In consequence, attempts at prevention should focus on secondary releases as well as on initial introductions. In the last section of the paper, the final invasion stage subsequent to naturalisation is shown as a multi-scale phenomenon. In consequence, the classification of a species as 'invasive' depends on the perspective chosen. Using different biologically or anthropocentrically based approaches leads to sub-sets of alien species that overlap only partially. In conclusion, the term `invasive' should preferably be used in a broader sense to describe the entire invasion process. This revised version was published online in July 2006 with corrections to the Cover Date.     

古尔班通古特沙漠半固定沙丘植物群落物种组成和种群结构

通过野外调查,研究古尔班通古特沙漠半固定沙丘植物群落的物种组成,采用株高、冠幅、体积大小级代替年龄结构,分析了典型灌木种群的生长状态及发展趋势.结果表明:在古尔班通古特沙漠半固定沙丘共调查到23种植物,出现最多的是藜科,6属8种,其次是菊科,5属6种,该沙漠植被组成数目少,群落结构简单.白梭梭主要分布在丘顶,为稳定增长型种群,而梭梭数量少.油蒿是当地工程建设后进行植被恢复而引入的物种,主要分布在迎风坡和丘顶,为增长型年龄结构,有很大的增长潜力,已经影响到原有的物种组成.白皮沙拐枣和蛇麻黄幼苗缺失,幼龄个体少,形成衰退型年龄结构,且白皮沙拐枣的空间分布区域与油蒿重合,未来可能被油蒿替代.     

Species Richness and Patterns of Invasion in Plants, Birds, and Fishes in the United States*

We quantified broad-scale patterns of species richness and species density (mean # species/km²) for native and non-indigenous plants, birds, and fishes in the continental USA and Hawaii. We hypothesized that the species density of native and non-indigenous taxa would generally decrease in northern latitudes and higher elevations following declines in potential evapotranspiration, mean temperature, and precipitation. County data on plants (n = 3004 counties) and birds (n=3074 counties), and drainage (6 HUC) data on fishes (n = 328 drainages) showed that the densities of native and non-indigenous species were strongly positively correlated for plant species (r = 0.86, P < 0.0001), bird species (r = 0.93, P<0.0001), and fish species (r = 0.41, P<0.0001). Multiple regression models showed that the densities of native plant and bird species could be strongly predicted (adj. R² = 0.66 in both models) at county levels, but fish species densities were less predictable at drainage levels (adj. R² = 0.31, P<0.0001). Similarly, non-indigenous plant and bird species densities were strongly predictable (adj. R² = 0.84 and 0.91 respectively), but non-indigenous fish species density was less predictable (adj. R² = 0.38). County level hotspots of native and non-indigenous plants, birds, and fishes were located in low elevation areas close to the coast with high precipitation and productivity (vegetation carbon). We show that (1) native species richness can be moderately well predicted with abiotic factors; (2) human populations have tended to settle in areas rich in native species; and (3) the richness and density of non-indigenous plant, bird, and fish species can be accurately predicted from biotic and abiotic factors largely because they are positively correlated to native species densities. We conclude that while humans facilitate the initial establishment, invasions of non-indigenous species, the spread and subsequent distributions of non-indigenous species may be controlled largely by environmental factors. The U.S. Government’s right to retain a non-exclusive, royalty-free licence in and to any copyright is acknowledged.     

Species introduced from nearby sources have a more homogenizing effect than species from distant sources: evidence from plants and fishes in the USA

Biotic homogenization is occurring in many biota as widespread introduced species are replacing unique native species. Although efforts to document homogenization have increased, no studies have explicitly compared the homogenizing effects of species introduced from distant areas to the homogenizing effects of species introduced from more proximate areas. The author analysed three data sets, at different scales and in different taxa, that distinguish species introduced from distant sources (e.g. outside the US) from species introduced from less distant sources (e.g. within the United States). These data include: plant introductions among eight major US cities and fish introductions among 12 US states and among 10 watersheds from New York state. The authors found that, for all data sets, species introduced from less distant sources (within the US) have a greater homogenizing effect on community composition than species from more distant sources (outside the US). In agreement with other studies, the author also find that, in terms of absolute numbers, introductions from nearby sources are far more frequent than introduction of species from distant sources. While tentative, these findings point out the importance of considering species introduced from nearby areas (e.g., extralimital native species) when discussing biotic homogenization from human activities.     

Patterns of Plant Invasions: A Case Example in Native Species Hotspots and Rare Habitats

Land managers require landscape-scale information on where exotic plant species have successfully established, to better guide research, control, and restoration efforts. We evaluated the vulnerability of various habitats to invasion by exotic plant species in a 100,000 ha area in the southeast corner of Grand Staircase-Escalante National Monument, Utah. For the 97 0.1-ha plots in 11 vegetation types, exotic species richness (log₁₀) was strongly negatively correlated to the cover of cryptobiotic soil crusts (r = −0.47, P < 0.001), and positively correlated to native species richness (r = 0.22, P < 0.03), native species cover (r = 0.23, P < 0.05), and total nitrogen in the soil (r = 0.40, P < 0.001). Exotic species cover was strongly positively correlated to exotic species richness (r = 0.68, P < 0.001). Only 6 of 97 plots did not contain at least one exotic species. Exotic species richness was particularly high in locally rare, mesic vegetation types and nitrogen rich soils. Dry, upland plots (n = 51) had less than half of the exotic species richness and cover compared to plots (n = 45) in washes and lowland depressions that collect water intermittently. Plots dominated by trees had significantly greater native and exotic species richness compared to plots dominated by shrubs. For the 97 plots combined, 33% of the variance in exotic species richness could be explained by a positive relationship with total plant cover, and negative relationships with the cover of cryptobiotic crusts and bare ground. There are several reasons for concern: (1) Exotic plant species are invading hot spots of native plant diversity and rare/unique habitats. (2) The foliar cover of exotic species was greatest in habitats that had been invaded by several exotic species.(3) Continued disturbance of fragile cryptobiotic crusts by livestock, people, and vehicles may facilitate the further invasion of exotic plant species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of Flow Restoration and Exotic Species Removal on Recovery of Native Fish: Lessons from a Dam Decommissioning

Flow diversion and invasive species are two major threats to freshwater ecosystems, threats that restoration efforts attempt to redress. Yet, few restoration projects monitor whether removal of these threats improve target characteristics of the ecosystem. Fewer still have an appropriate experimental design from which causal inferences can be drawn as to the relative merits of removing exotic fish, restoring flow, or both. We used a dam decommissioning in Fossil Creek, Arizona, to compare responses of native fish to exotic fish removal and flow restoration, using a before‐after‐control‐impact design with three impact treatments: flow restoration alone where exotics had not been present, flow restoration and exotic fish removal, and flow restoration where exotics remain and a control reach that was unaffected by restoration actions. We show that removal of exotic fish dramatically increased native fish abundance. Flow restoration also increased native fish abundance, but the effect was smaller than that from removing exotics. Flow restoration had no effect where exotic fish remained, although it may have had other benefits to the ecosystem. The cost to restore flow ($12 million) was considerably higher than that to eradicate exotics ($1.1 million). The long‐term influence of flow restoration could increase, as travertine dams grow and re‐shape the creek increasing habitat for native fish. But in the 2‐year period considered here, the return on investment for extirpating exotics far exceeded that from flow restoration. Projects aimed to restore native fish by restoring flow should also consider the additional investment required to eradicate exotic fish.     

Recovery of Plant Species Richness and Composition in an Abandoned Forest Settlement Area in Kenya

Cultivation of annual crops in the initial stage of reforestation has been commonly practiced in the tropics. In recent decades, however, cultivation of such areas has been discontinued, resulting in widespread abandoned settlements. In this article we used a former forest village settlement in Kenya, which had been cleared, cultivated and then abandoned, to study how natural vegetation recovers after such disturbances. Species richness, abundance, and composition of tree seedlings, saplings, adult trees, shrubs, and herbs were recorded in different zones, from a heavily degraded zone in the center of the settlement, through less disturbed transition zones (TZs), and in the surrounding secondary forest (SF). Species richness and abundance of tree seedlings, saplings, and adult trees increased gradually from the heavily degraded zone to the SF, whereas shrub and herb richness were the same for TZs and SF and abundance was lowest in the SF. Total species richness was highest in the SF. Some pioneer tree species were highly associated with the TZs, whereas sub‐canopy tree species were associated with the SF. A group of tree species were not particularly associated with any of the four zones. Thus, these species might have good potential as restoration species. The results of our study contribute to the knowledge of natural regeneration in general, and of individual species characterizing the different stages of recovery of abandoned settlements in particular. Such information is urgently needed in designing ecologically sound management strategies for restoring abandoned forest settlements in tropical areas.     

Effects of Species Richness on Resident and Target Species Components in a Prairie Restoration

The ecological role of biodiversity in achieving successful restoration has been little explored in restoration ecology. We tested the prediction that we are more likely to create persistent, species‐rich plant communities by increasing the number of species sown, and, to some degree, by varying functional group representation, in experimental prairie plantings. There were 12 treatments consisting of 1‐, 2‐, 3‐, 4‐, 8‐, 12‐, and 16‐species mixtures of native perennials representing four functional groups (C₄ grasses, C₃ grasses, nitrogen‐fixing species, and late‐flowering composites) that predominate within Central Plains tallgrass prairies. In 2000, species were seeded into square plots (6 × 6 m), with five replicates per treatment, on former agricultural land. Annually, we measured total species richness and evenness, target species richness and cover, and richness and cover of resident species (i.e., those emerging from the seed bank). Both target species richness and rate of establishment of target communities were highest in the most species‐rich mixtures, but there was no additional benefit for treatments that contained more than eight species. Richness of resident species did not vary with target species richness; however, cover by resident species was lower in the higher target species treatments. Our results, indicating that establishment of species‐rich prairie mimics can be enhanced by starting with larger numbers of species at the outset, have implications for grassland restoration in which community biodiversity creation and maintenance are key goals.     

Invading species in the Eel River, California: successes, failures, and relationships with resident species

We examined invasions of non-native fishes into the Eel River, California. At least 16 species of fish have been introduced into the drainage which originally supported 12–14 fish species. Our study was prompted by the unauthorized introduction in 1979 of Sacramento squawfish, Ptychocheilus grandis, a large predatory cyprinid. From 1986 to 1990, we conducted growth and diet studies of squawfish, conducted intensive surveys of the distribution and habitat associations of both native and introduced species, and examined the nature of species-habitat and interspecies relationships. We found no evidence for increased growth or expanded feeding habits, compared to native populations, of Sacramento squawfish as they invaded the Eel River drainage. Ten of the introduced species were well established, with four species limited to a reservoir and six species established in streams. The success or failure of introductions of stream species appeared to be a function of the ability of a species to survive the fluctuating, highly seasonal, flow regime. The present mixture of native and exotic species has not formed stable fish assemblages but it seems likely that four habitat-associated assemblages will develop. The overall effect of the successful species introductions has been to assemble a group of species, with some exceptions, that are native to and occur together in many California streams. The assemblages now forming are similar to those found in other California streams. The assemblage characterized by squawfish and suckers is likely to be resistant to invasion, in the absence of human caused habitat modifications.     

Sustainability,Health, and the Human Population

A sustainable human population (e.g., range, density, and total numbers) is essential to health and in management. The notion of sustainability applies to all species and ecosystems and to the biosphere. Sustainability involves the health not only of individual humans, but also of ecosystems and other species. Thus, sustainability of the human population is important because of the wealth of factors involved: both the elements of systems it affects and those that contribute to its size. In this article, I address the sustainability of the human population on the basis of the argument that other species serve as examples of sustainability at the species level—an example of an application of systemic management that simultaneously accounts for complexity and achieves measurable health for individuals, species, and ecosystems. I conclude that the human population is two to four orders of magnitude larger than is optimally sustainable when compared with the populations of other mammalian species of similar body size and that this is a significant contributor to health problems for our species, other species, and ecosystems—a systemic pathology.     

Effects of Herbicide on the Invasive Biennial Alliaria petiolata (Garlic Mustard) and Initial Responses of Native Plants in a Southwestern Ohio Forest

Restoration often includes control of invasive plants, but little is known about how native plant communities respond to this control. The biennial Alliaria petiolata (M. Bieb.) Cavara and Grande (garlic mustard) is one of the most prevalent invasive plants in forests of eastern North America. We investigated the effects of the herbicide Round‐up (glyphosate) on Alliaria and the response of the forest floor plant community to the herbicide and the subsequent decline of Alliaria. In an old‐growth Acer–Fagus stand and a second‐growth Liriodendron‐dominated stand in Hueston Woods State Nature Preserve, Ohio, United States, we spot applied Round‐up in November 2000 and 2001 in 25 1 × 1–m plots and maintained 25 control plots. Herbicide decreased Alliaria density in both stands and reduced the density of other species in leaf during treatment (mostly exotic winter annuals) in the old‐growth stand. Treatment did not affect the initial density of the Alliaria cohort that germinated in the spring of 2001, but decreased the 2002 cohort. Community differences were found in the old‐growth stand after Alliaria reduction, specifically greater cover of spring ephemerals in the herbicide treatment. In the second‐growth stand, herbicide treatment increased reproduction of the late‐summer perennial, Phryma leptostachya. These results indicate that glyphosate reduces Alliaria without negatively impacting native species and that some native species respond positively to a single‐year reduction in this invasive biennial.     

Calculating Climate Effects on Birds and Mammals: Impacts on Biodiversity, Conservation, Population Parameters, and Global Community Structure

This paper describes how climate variation in time and spacecan constrain community structure on a global scale. We explorebody size scaling and the energetic consequences in terms ofabsorbed mass and energy and expended mass and energy. We explainhow morphology, specific physiological properties, and temperaturedependent behaviors are key variables that link individual energeticsto population dynamics and community structure. This paper describes an integrated basic principles model formammal energetics and extends the model to bird energetics.The model additions include molar balance models for the lungsand gut. The gut model couples food ingested to respiratorygas exchanges and evaporative water loss from the respiratorysystem. We incorporate a novel thermoregulatory model that yieldsmetabolic calculations as a function of temperature. The calculationsmimic empirical data without regression. We explore the differencesin the quality of insulation between hair and feathers withour porous media model for insulation. For mammals ranging in size from mice to elephants we show thatcalculated metabolic costs are in agreement with experimentaldata. We also demonstrate how we can do the same for birds rangingin size from hummingbirds to ostriches. We show the impact ofchanging posture and changing air temperatures on energeticcosts for birds and mammals. We demonstrate how optimal bodysize that maximizes the potential for growth and reproductionchanges with changing climatic conditions and with diet quality.Climate and diet may play important roles in constraining communitystructure (collection of functional types of different bodysizes) at local and global scales. Thus, multiple functionaltypes may coexist in a locality in part because of the temporaland spatial variation in climate and seasonal food variation.We illustrate how the models can be applied in a conservationand biodiversity context to a rare and endangered species ofparrot, the Orange-bellied Parrot of Australia and Tasmania.     

Invasiveness and homogenization: synergism of wide dispersal and high local abundance

Aim To determine whether invasive and locally abundant non‐native species have a more homogenizing effect on plant communities than non‐invasive and less abundant non‐native species. Location California and Florida counties, conservation areas in the USA, and eight US cities. Methods Species lists among counties, conservation areas and cities were compared to see whether invasive and abundant non‐native species increased the Jaccard index of similarity between localities beyond any increases caused by non‐invasive and less abundant non‐native species. Results For all comparisons, we found that invasive non‐native species have a significantly greater homogenizing effect than non‐invasive non‐native species. For the US conservation areas, we found that locally abundant invasive species tend to be more widespread and more widely shared than less abundant invasive species. There is also a positive relationship between homogenization by invasive species and the magnitude of human disturbance. Main conclusions Invasive non‐native species tend to be disproportionately shared among communities relative to non‐invasive non‐native species. This effect is enhanced by human disturbance, as measured by the ratio of non‐native to native species. There is a synergism between abundance and geographical range which enhances the homogenizing effects of abundant species. Invasive species, with wide ecological niches, are more widely shared among communities and more locally abundant. Abundant invasive species are thus more spatially homogenizing, and more ecologically dominant (functionally homogenizing). Also, ‘perceived homogenization’ is probably greater than homogenization measured by the increase in shared species. The abundant species typically seen by the casual observer in a biological community are probably more commonly shared between communities than less common species. Studies that lack abundance data and measure homogenization only on the basis of shared species, which includes most homogenization studies to date, probably underestimate the homogenizing impacts of non‐native species as perceived by people.     

Population survey and management strategies of free-roaming dogs (Canis familiaris) on Saipan,Commonwealth of the Northern Mariana Islands

The island of Saipan, Commonwealth of the Northern Mariana Islands, is populated by free-roaming dogs who were introduced during World War II. The local nonhuman animal control agency manages this population; however, the demographic information and public perception of this population remain unknown. To characterize the free-roaming dog population, an island-wide survey on Saipan was conducted. Photographic, mark-recapture data were used to estimate the population size per land type. Age, sex, sociality, and behavior per land type were documented and associations between these variables were tested using G tests. The effects of land type and urbanization on dog abundance were also analyzed. Lastly, in-person surveys provided data on the public perception of free-roaming dogs. Four main findings are reported: (a) The population size of free-roaming dogs on Saipan is estimated at 21,316. (b) Most dogs encountered were sentry adults. (c) Dog abundance increased with urbanization level and was highest in urban areas. (d) The public perceived free-roaming dogs as a health concern and suggested the implementation of leash laws and sheltering initiatives. This article discusses potential and existing population management strategies for free-roaming dogs on Saipan.     

Crossing the (Wallace) line: local abundance and distribution of mammals across biogeographic barriers

Past and ongoing vertebrate introductions threaten to rearrange ecological communities in the Indo‐Malay Archipelago, one of Earth's most biodiverse regions. But the consequences of these translocations are difficult to predict. We compared local abundance and distributions in four tropical mammal lineages that have crossed from Asia to Wallacea or New Guinea. The local abundance of macaques (Macaca spp.), which naturally crossed Wallace's Line, was higher in Sulawesi (east of the line; mean = 3.7 individuals per camera station, 95% CI = 2.2: 5.1) than in Borneo (west of the line; mean = 1.1, CI = 0.8: 1.4), but the local abundance of Malay civets (Viverra tangalunga), Rusa deer, and Sus pigs was similar in their native ranges and where they had been introduced by humans east of Wallace's Line. Proximity to rivers increased Malay Civet local abundance and decreased the local abundance of pigs in parts of their introduced ranges (Maluku and New Guinea, respectively), while having no effect on local abundance in their native ranges (Borneo) or other areas where they have been introduced (Sulawesi). That local abundance was higher east of Wallace's Line in just one of four mammal lineages is consistent with findings from plant invasions, where most species have similar abundance in their native and introduced ranges. However, species’ ecology may change as they enter new communities, for example, their patterns of abundance at local scales. This could make it difficult to predict community structure in the face of ongoing species introductions.     

Predicting Coral Species Richness: The Effect of Input Variables,Diversity and Scale

Coral reefs are facing a biodiversity crisis due to increasing human impacts, consequently, one third of reef-building corals have an elevated risk of extinction. Logistic challenges prevent broad-scale species-level monitoring of hard corals; hence it has become critical that effective proxy indicators of species richness are established. This study tests how accurately three potential proxy indicators (generic richness on belt transects, generic richness on point-intercept transects and percent live hard coral cover on point-intercept transects) predict coral species richness at three different locations and two analytical scales. Generic richness (measured on a belt transect) was found to be the most effective predictor variable, with significant positive linear relationships across locations and scales. Percent live hard coral cover consistently performed poorly as an indicator of coral species richness. This study advances the practical framework for optimizing coral reef monitoring programs and empirically demonstrates that generic richness offers an effective way to predict coral species richness with a moderate level of precision. While the accuracy of species richness estimates will decrease in communities dominated by species-rich genera (e.g. Acropora), generic richness provides a useful measure of phylogenetic diversity and incorporating this metric into monitoring programs will increase the likelihood that changes in coral species diversity can be detected.     

Effects of Environmental Heterogeneity and Disturbance on the Native and Non-native Flora of Desert Springs

Vegetation often is used as a decision variable for conservation and resource management. Because time and money are limited, it is useful to identify predictable relationships between measures of vegetation diversity or status, the physical environment, and disturbance; native and non-native plants may have different functional responses. Working towards development of effective, practical strategies for management and ecological restoration in the Spring Mountains, an isolated mountain range in the eastern Mojave Desert (Nevada, USA) that is a focus of regional conservation planning, we examined whether native and non-native assemblages of spring-associated perennial plants have predictable relationships with elevation, springbrook length, and various land uses. We also tested whether elevation, springbrook length, and overall disturbance were associated with the degree of predictability of local species presence and absence. Consistent with work in other systems, species richness and cover of native plants tended to decrease as intensity of disturbance increased, whereas species richness (but not cover) of non-native plants tended to peak with intermediate disturbance. Our results may suggest that invasions of non-native plants at springs in the Spring Mountains are relatively recent, and that rapid restoration and management actions may help protect ecological processes and viability of native plants. Ability to predict the order in which individual species are likely to be extirpated from or colonize springs was limited, perhaps reflecting considerable environmental heterogeneity among springs.     

The worldwide transfer of ants: geographical distribution and ecological invasions

Aim This is the first comprehensive account of the biogeography of ants transferred and at least temporarily established outside their native habitat. Location Using museum and literature records, I established the distributions of transferred ant species. Methods I used taxonomic and functional groups to assess how geographical spread as a transferred species is affected by taxonomy and life history. Results 147 ant species in forty-nine genera have been recorded outside of their native habitat. The proportion of transferred ants is similar to the number of genera and species in each subfamily. The species-rich subfamily Myrmicinae contains nearly 50% of all transferred species, while many of the species-poor subfamilies have absolutely no transferred species. A disproportionate high number of transferred ants originate from the Neotropical and Oriental biogeographic regions. The Pacific Islands are the recipients of the most transferred ant species. Most transferred ants belong to the CRYPTIC, OPPORTUNIST, and GENERALIZED MYRMICINE functional groups, while there are no recorded transfers of army ants or leaf-cutting ants. Both invasive and human commensal ‘tramp’ ant species are nonrandom subsets of transferred ants. Main conclusions ‘Tramp’ species and invasive species tend to have widespread geographical distributions, and share life history characteristics including queen number, nest structure, and foraging behaviour. Combining observations of functional groups and biogeography may lead to a better understanding of the factors contributing to the spread of transferred species.