Native faunal communities depend on habitat from non-native plants in novel but not in natural ecosystems

Invasive non-native plants are a major driver of native biodiversity loss, yet native biodiversity can sometimes benefit from non-native species. Depending on habitat context, even the same non-native species can have positive and negative effects on biodiversity. Blackberry (Rubus fruticosus aggregate) is a useful model organism to better understand a non-native plant with conflicting impacts on biodiversity. We used a replicated capture-mark-recapture study across 11 consecutive seasons to examine the response of small mammal diversity and abundance to vegetation structure and density associated with non-native blackberry (R. anglocandicans) in native, hybrid and blackberry-dominated novel ecosystems in Australia. Across the three habitat types, increasing blackberry dominance had a positive influence on mammal diversity, while the strength and direction of this influence varied for abundance. At a microhabitat scale within hybrid and native habitat there were no significant differences in diversity, or the abundance of most species, between microhabitats where blackberry was absent versus dominant. In contrast, in novel ecosystems diversity and abundances were very low without blackberry, yet high (comparable to native ecosystems) within blackberry as it provided functionally-analogous vegetation structure and density to the lost native understory. Our results indicate the ecological functions of non-native plant species vary depending on habitat and need to be considered for management. Comparative studies such as ours that apply a standardized approach across a broad range of conditions at the landscape and habitat scale are crucial for guiding land managers on control options for non-native species (remove, reduce or retain and contain) that are context-sensitive and scale-dependent.     

Diversity and Biotic Homogenization of Urban Land-Snail Faunas in Relation to Habitat Types and Macroclimate in 32 Central European Cities

The effects of non-native species invasions on community diversity and biotic homogenization have been described for various taxa in urban environments, but not for land snails. Here we relate the diversity of native and non-native land-snail urban faunas to urban habitat types and macroclimate, and analyse homogenization effects of non-native species across cities and within the main urban habitat types. Land-snail species were recorded in seven 1-ha plots in 32 cities of ten countries of Central Europe and Benelux (224 plots in total). Each plot represented one urban habitat type characterized by different management and a specific disturbance regime. For each plot, we obtained January, July and mean annual temperature and annual precipitation. Snail species were classified into either native or non-native. The effects of habitat type and macroclimate on the number of native and non-native species were analysed using generalized estimating equations; the homogenization effect of non-native species based on the Jaccard similarity index and homogenization index. We recorded 67 native and 20 non-native species. Besides being more numerous, native species also had much higher beta diversity than non-natives. There were significant differences between the studied habitat types in the numbers of native and non-native species, both of which decreased from less to heavily urbanized habitats. Macroclimate was more important for the number of non-native than native species; however in both cases the effect of climate on diversity was overridden by the effect of urban habitat type. This is the first study on urban land snails documenting that non-native land-snail species significantly contribute to homogenization among whole cities, but both the homogenization and diversification effects occur when individual habitat types are compared among cities. This indicates that the spread of non-native snail species may cause biotic homogenization, but it depends on scale and habitat type.     

Pathogens in Crassostrea ariakensis and other Asian oyster species: implications for non-native oyster introduction to Chesapeake Bay

With the drastic decline of eastern oyster Crassostrea virginica populations in the Chesapeake Bay due to over-fishing, diseases and habitat destruction, there is interest in Maryland and Virginia in utilizing the non-native oyster species Crassostrea ariakensis for aquaculture, fishery resource enhancement, and ecological restoration. The International Council for the Exploration of the Sea (ICES) recommends that non-native species be examined for ecological, genetic and disease relationships in the native range prior to a deliberate introduction to a new region. Therefore, a pathogen survey of C. ariakensis and other sympatric oyster species was conducted on samples collected in the PR China, Japan and Korea using molecular diagnostics and histopathology. Molecular assays focused on 2 types of pathogens: protistan parasites in the genus Perkinsus and herpesviruses, both with known impacts on commercially important molluscan species around the world, including Asia. PCR amplification and DNA sequence data from the internal transcribed spacer region of the rRNA gene complex revealed the presence of 2 Perkinsus species not currently found in USA waters: P. olseni and an undescribed species. In addition, 3 genetic strains of molluscan herpesviruses were detected in oysters from several potential C. ariakensis broodstock acquisition sites in Asia. Viral gametocytic hypertrophy, Chlamydia-like organisms, a Steinhausia-like microsporidian, Perkinsus sp., Nematopsis sp., ciliates, and cestodes were also detected by histopathology.     

Non-native ecosystem engineer alters estuarine communities

Many ecosystems are created by the presence of ecosystem engineers that play an important role in determining species' abundance and species composition. Additionally, a mosaic environment of engineered and non-engineered habitats has been shown to increase biodiversity. Non-native ecosystem engineers can be introduced into environments that do not contain or have lost species that form biogenic habitat, resulting in dramatic impacts upon native communities. Yet, little is known about how non-native ecosystem engineers interact with natives and other non-natives already present in the environment, specifically whether non-native ecosystem engineers facilitate other non-natives, and whether they increase habitat heterogeneity and alter the diversity, abundance, and distribution of benthic species. Through sampling and experimental removal of reefs, we examine the effects of a non-native reef-building tubeworm, Ficopomatus enigmaticus, on community composition in the central Californian estuary, Elkhorn Slough. Tubeworm reefs host significantly greater abundances of many non-native polychaetes and amphipods, particularly the amphipods Monocorophium insidiosum and Melita nitida, compared to nearby mudflats. Infaunal assemblages under F. enigmaticus reefs and around reef's edges show very low abundance and taxonomic diversity. Once reefs are removed, the newly exposed mudflat is colonized by opportunistic non-native species, such as M. insidiosum and the polychaete Streblospio benedicti, making removal of reefs a questionable strategy for control. These results show that provision of habitat by a non-native ecosystem engineer may be a mechanism for invasional meltdown in Elkhorn Slough, and that reefs increase spatial heterogeneity in the abundance and composition of benthic communities.     

A Framework for Evaluating Heterogeneity and Landscape-Level Impacts of Non-native Aquatic Species

Non-native species are a major component of global environmental change, and aquatic systems are especially vulnerable to non-native species impacts. Much of the research on aquatic non-native species impact has occurred at the local or site level. In reality, non-native species impacts play out across multiple spatial scales on heterogeneous landscapes. How can we ‘scale up’ our understanding of site-level impacts to the broader landscape scale? To address this disconnect, we synthesize our current understanding of key components of landscape-scale non-native species impacts: geographic range, abundance, and local impacts. Most aquatic non-native species have small ranges, while a few have large ranges. However, aquatic non-native species are often far from saturated on landscapes, and occurrence records are often woefully incomplete. Aquatic non-native species are often at low abundances where they are present, reaching high abundance in a small number of locations. Finally, local-scale impact can be estimated from abundance, but this requires knowledge of the abundance–impact relationship. Considering these multiple components enables understanding of non-native species impacts at broader spatial scales. Although the landscape-level impacts of aquatic non-native species may be high, the spatial distribution of site-level impacts is uneven, and highly impacted sites may be relatively uncommon. This heterogeneity in impacts provides an opportunity to optimize and prioritize non-native species management and prevention efforts.     

Engineering role models: do non-human species have the answers?

A shift from traditional engineering approaches to ecologically-based techniques will require changing societal values regarding ‘how and what’ is defined as engineering and design. Non-human species offer many ecological engineering examples that are often beneficial to ecosystem function and other biota. For example, organisms known as ‘ecosystem engineers’ build, modify, and destroy habitat in their quest for food and survival. Similarly, ‘keystone species’ have greater impacts on community or ecosystem function than would be predicted from their abundance. The capacity of these types of organisms to affect ecosystems is great. They exert controlling influences over ecosystems and communities by altering resource allocation, creating habitats and modifying relative competitive advantages.Species’ effects in ecosystems, although context-dependent, can be evaluated as ‘beneficial’ or ‘detrimental’. The evaluation depends on whether effects on other species or ecosystem function are more or less desirable from a given perspective. Organisms with beneficial impacts facilitate the presence of other species, employ efficient nutrient cycling, and are sometimes characterized by specific mutualisms. In contrast, many cases of detrimental engineering are found from introduced (i.e., exotic) species and are characterized by a loss of species richness, a lack of nutrient retention and the degradation of ecosystem integrity. Species’ impacts on ecosystems and community traits have been quantified in ecological studies and can be used similarly to understand, design and model human engineering structures and impacts on the landscape. Emulation of species with beneficial impacts on ecosystems can provide powerful guidance to the goals of ecological engineering. Using role model organisms that have desirable effects on species diversity and ecosystem function will be important in developing alternatives to traditional engineering practices.     

Context dependence of marine ecosystem engineer invasion impacts on benthic ecosystem functioning

Introduced ecosystem engineers can severely modify the functioning on invaded systems. Species-level effects on ecosystem functioning (EF) are context dependent, but the effects of introduced ecosystem engineers are frequently assessed through single-location studies. The present work aimed to identify sources of context-dependence that can regulate the impacts of invasive ecosystem engineers on ecosystem functioning. As model systems, four locations where the bivalve Ruditapes philippinarum (Adams and Reeve) has been introduced were investigated, providing variability in habitat characteristics and community composition. As a measure of ecosystem engineering, the relative contribution of this species to community bioturbation potential was quantified at each site. The relevance of bioturbation to the local establishment of the mixing depth of marine sediments (used as a proxy for EF) was quantified in order to determine the potential for impact of the introduced species at each site. We found that R. philippinarum is one of the most important bioturbators within analysed communities, but the relative importance of this contribution at the community level depended on local species composition. The net contribution of bioturbation to the establishment of sediment mixing depths varied across sites depending on the presence of structuring vegetation, sediment granulometry and compaction. The effects of vegetation on sediment mixing were previously unreported. These findings indicate that the species composition of invaded communities, and the habitat characteristics of invaded systems, are important modulators of the impacts of introduced species on ecosystem functioning. A framework that encompasses these aspects for the prediction of the functional impacts of invasive ecosystem engineers is suggested, supporting a multi-site approach to invasive ecology studies concerned with ecosystem functioning.     

Ecosystem engineering effects on species diversity across ecosystems: a meta‐analysis

Ecosystem engineering is increasingly recognized as a relevant ecological driver of diversity and community composition. Although engineering impacts on the biota can vary from negative to positive, and from trivial to enormous, patterns and causes of variation in the magnitude of engineering effects across ecosystems and engineer types remain largely unknown. To elucidate the above patterns, we conducted a meta‐analysis of 122 studies which explored effects of animal ecosystem engineers on species richness of other organisms in the community. The analysis revealed that the overall effect of ecosystem engineers on diversity is positive and corresponds to a 25% increase in species richness, indicating that ecosystem engineering is a facilitative process globally. Engineering effects were stronger in the tropics than at higher latitudes, likely because new or modified habitats provided by engineers in the tropics may help minimize competition and predation pressures on resident species. Within aquatic environments, engineering impacts were stronger in marine ecosystems (rocky shores) than in streams. In terrestrial ecosystems, engineers displayed stronger positive effects in arid environments (e.g. deserts). Ecosystem engineers that create new habitats or microhabitats had stronger effects than those that modify habitats or cause bioturbation. Invertebrate engineers and those with lower engineering persistence (<1 year) affected species richness more than vertebrate engineers which persisted for >1 year. Invertebrate species richness was particularly responsive to engineering impacts. This study is the first attempt to build an integrative framework of engineering effects on species diversity; it highlights the importance of considering latitude, habitat, engineering functional group, taxon and persistence of their effects in future theoretical and empirical studies.     

Anthropogenic water conditions amplify predatory impact of the non-native Oriental river prawn Macrobrachium nipponense

Biological Invasions - Investigating non-native species’ impacts on recipient communities in the context of local habitat conditions is vital to understanding their potential invasiveness....     

Epifaunal and algal assemblages associated with the native Chondrus crispus (Stackhouse) and the non-native Grateloupia turuturu (Yamada) in eastern Long Island Sound

Although the spread of non-native algae is rapidly escalating, relatively few ecological studies have been done to gauge the impacts incurred to native flora and fauna. A reduction in the dominance of a native habitat-forming macroalga due to the replacement by an introduced species can have adverse effects on the community. The non-native red alga Grateloupia turuturu, first reported in Rhode Island, USA in 1994, has since extended its southern range into eastern Long Island Sound. This large alga has the potential to impact coastal communities by altering the floristic composition important to associated flora and fauna. A comparison of algal and epifaunal assemblages was made during 2006 and 2007 between native and non-native algal communities dominated by either G. turuturu or the native, Chondrus crispus at two sites in Long Island Sound. We found that within Grateloupia-dominated habitat, there was a large decrease in overall macrophyte biomass as compared to native habitat. We also found that habitat dominated by the non-native alga reduced species richness and total abundance of invertebrates relative to nearby habitats dominated by C. crispus. In addition, we found that the dominant mesofaunal species, important to higher trophic level consumers, had greatly reduced densities in communities dominated by the non-native alga.     

Disappearing refuges in time and space: how environmental change threatens species coexistence

Understanding the impacts of environmental changes on species survival is a major challenge in ecological research, especially when shifting from single- to multispecies foci. Here, we apply a spatially explicit two-species simulation model to analyze the effects of geographic range shifting and habitat isolation on different coexistence mechanisms. The model explicitly considers dispersal, local competition, and growth on a single resource. Results highlight that both range shifting and habitat isolation severely impact coexistence. However, the strength of these impacts depends on the underlying coexistence mechanisms. Neutrally coexisting species are particularly sensitive to habitat isolation, while stabilized coexistence through overcompensatory density regulation is much more sensitive to range shifting. We conclude that, at the community level, the response to environmental change sensitively depends on the underlying coexistence mechanisms. This suggests that predictions and management recommendations should consider differences between neutral versus stabilized community structures whenever possible.     

Metacommunities of spiders in grassland habitat fragments of an agricultural landscape

Arthropod communities in fragmented agricultural landscapes depend on local processes and the interactions between communities in the habitat islands. We aimed to study metacommunity structure of spiders, a group that is known for high dispersal power, local niche partitioning and for engaging in species interactions. While living in fragmented habitats could lead to nestedness, other ecological traits of spiders might equally lead to patterns dominated either by species interactions or habitat filtering. We asked, which community pattern will prevail in a typical agricultural landscape with isolated patches of semi-natural habitats. Such a situation was studied by sampling spiders in 28 grassland locations in a Hungarian agricultural landscape. We used the elements of metacommunity structure (EMS) framework to distinguish between alternative patterns that reveal community organization. The EMS analysis indicated coherent species ranges, high turnover and boundary clumping, suggesting Clementsian community organization. The greatest variation in species composition was explained by local habitat characteristics, indicating habitat filtering. The influence of dispersal could be detected by the significant effect of landscape composition, which was strongest at 500 m. We conclude that dispersal allows spiders to respond coherently to the environment, creating similar communities in similar habitats. Consistent habitat differences, such as species rich versus species poor vegetation, lead to recognisably different, recurrent communities. These characteristics make spiders a predictable and diverse source of natural enemies in agricultural landscapes. Sensitivity to habitat composition at medium distances warns us that landscape homogenization may alter these metacommunity processes.     

Ecology of invasive mosquitoes: effects on resident species and on human health

Investigations of biological invasions focus on patterns and processes that are related to introduction, establishment, spread and impacts of introduced species. This review focuses on the ecological interactions operating during invasions by the most prominent group of insect vectors of disease, mosquitoes. First, we review characteristics of non-native mosquito species that have established viable populations, and those invasive species that have spread widely and had major impacts, testing whether biotic characteristics are associated with the transition from established non-native to invasive. Second, we review the roles of interspecific competition, apparent competition, predation, intraguild predation and climatic limitation as causes of impacts on residents or as barriers to invasion. We concentrate on the best-studied invasive mosquito, Aedes albopictus, evaluating the application of basic ecological theory to invasions by Aedes albopictus. We develop a model based on observations of Aedes albopictus for effects of resource competition and predation as barriers to invasion, evaluating which community and ecosystem characteristics favour invasion. Third, we evaluate the ways in which invasive mosquitoes have contributed to outbreaks of human and animal disease, considering specifically whether invasive mosquitoes create novel health threats, or modify disease transmission for existing pathogen-host systems.     

Fish species introductions provide novel insights into the patterns and drivers of phylogenetic structure in freshwaters

Despite long-standing interest of terrestrial ecologists, freshwater ecosystems are a fertile, yet unappreciated, testing ground for applying community phylogenetics to uncover mechanisms of species assembly. We quantify phylogenetic clustering and overdispersion of native and non-native fishes of a large river basin in the American Southwest to test for the mechanisms (environmental filtering versus competitive exclusion) and spatial scales influencing community structure. Contrary to expectations, non-native species were phylogenetically clustered and related to natural environmental conditions, whereas native species were not phylogenetically structured, likely reflecting human-related changes to the basin. The species that are most invasive (in terms of ecological impacts) tended to be the most phylogenetically divergent from natives across watersheds, but not within watersheds, supporting the hypothesis that Darwin''s naturalization conundrum is driven by the spatial scale. Phylogenetic distinctiveness may facilitate non-native establishment at regional scales, but environmental filtering restricts local membership to closely related species with physiological tolerances for current environments. By contrast, native species may have been phylogenetically clustered in historical times, but species loss from contemporary populations by anthropogenic activities has likely shaped the phylogenetic signal. Our study implies that fundamental mechanisms of community assembly have changed, with fundamental consequences for the biogeography of both native and non-native species.     

Evaluating the Risk of Establishing a Self-Sustaining Population of Non-Native Oysters Through Large-Scale Aquaculture in Chesapeake Bay

A multistep invasive species methodology was used to evaluate the probability that the deployment of sterile non-native oysters (Crassostrea ariakensis, called Asian oyster) in field-based aquaculture could give rise to a reproductive population in Chesapeake Bay. Several pathways that could potentially lead to the unintentional release of reproductive C. ariakensis from aquaculture were identified. For the four quantifiable pathways, a probability was estimated for each step based on knowledge of ecological rates. A conservative estimate for the number of reproductive C. ariakensis that could arise over one spawning season at a hypothetical aquaculture site was estimated. A statistical expansion of this probability estimated that it is highly likely that the cultivation of putatively sterile C. ariakensis would initiate a reproductive population in Chesapeake Bay over a 10-year time span. The potential benefits of restoration actions involving non-native species in estuarine ecosystems must be weighed against potential ecological risks. A probabilistic approach can provide a useful method for summarizing the risk of an unintended introduction that may occur despite a reasonable and presumably “safe” approach for realizing benefits of a non-native species in aquaculture. The results presented here have important implications for future proposals involving the commercial use of non-native species in natural ecosystems.     

Ecological periodic tables: in principle and practice

“Science is organized knowledge.” Immanuel Kant (1724–1804) Ecological periodic tables are an information organizing system with categorical habitat types as elements and predictably recurring (periodic) properties of a target biotic community, such as its relative species richness, abundance and biomass, as attributes. Ecological periodic tables are founded on the ecological tenet that habitats structure biotic communities and its corollary that habitats are templets for ecological strategies. They are a durable, open and flexible system that accommodates all operationally defined habitat types and biotic communities for which the periodicity of habitat usage patterns by a biotic community have been empirically substantiated. Discovering quantitative, periodic habitat usage patterns requires quantitative, representative, unbiased sampling of a biotic community across habitat types at ecologically relevant temporal and spatial scales. Like chemical periodic tables, the Linnaean system of classification and the Hertzsprung–Russell diagram in chemistry, biology and astronomy, respectively, ecological periodic tables are simple, easy to understand, exceptionally useful and they foster the expansion of scientific understanding, inquiry and theory.     

Differential population responses of native and alien rodents to an invasive predator,habitat alteration and plant masting

Invasive species and anthropogenic habitat alteration are major drivers of biodiversity loss. When multiple invasive species occupy different trophic levels, removing an invasive predator might cause unexpected outcomes owing to complex interactions among native and non-native prey. Moreover, external factors such as habitat alteration and resource availability can affect such dynamics. We hypothesized that native and non-native prey respond differently to an invasive predator, habitat alteration and bottom-up effects. To test the hypothesis, we used Bayesian state-space modelling to analyse 8-year data on the spatio-temporal patterns of two endemic rat species and the non-native black rat in response to the continual removal of the invasive small Indian mongoose on Amami Island, Japan. Despite low reproductive potentials, the endemic rats recovered better after mongoose removal than did the black rat. The endemic species appeared to be vulnerable to predation by mongooses, whose eradication increased the abundances of the endemic rats, but not of the black rat. Habitat alteration increased the black rat''s carrying capacity, but decreased those of the endemic species. We propose that spatio-temporal monitoring data from eradication programmes will clarify the underlying ecological impacts of land-use change and invasive species, and will be useful for future habitat management.     

Can Invasive Species Facilitate Native Species? Evidence of How, When, and Why These Impacts Occur

Although the predatory and competitive impacts of biological invasions are well documented, facilitation of native species by non-indigenous species is frequently overlooked. A search through recent ecological literature found that facilitative interactions between invasive and native species occur in a wide range of habitats, can have cascading effects across trophic levels, can re-structure communities, and can lead to evolutionary changes. These are critical findings that, until now, have been mostly absent from analyses of ecological impacts of biological invasions. Here I present evidence for several mechanisms that exemplify how exotic species can facilitate native species. These mechanisms include habitat modification, trophic subsidy, pollination, competitive release, and predatory release. Habitat modification is the most frequently documented mechanism, reflecting the drastic changes generated by the invasion of functionally novel habitat engineers. Further, I predict that facilitative impacts on native species will be most likely when invasive species provide a limiting resource, increase habitat complexity, functionally replace a native species, or ameliorate predation or competition. Finally, three types of facilitation (novel, substitutive, and indirect) define why exotic species can lead to facilitation of native species. It is evident that understanding biological invasions at the community and ecosystem levels will be more accurate if we integrate facilitative interactions into future ecological research. Since facilitative impacts of biological invasions can occur with native endangered species, and can have wide-ranging impacts, these results also have important implications for management, eradication, and restoration.Contribution Number 2293, Bodega Marine Laboratory, University of California at Davis.     

集合群落及其在河流鱼类群聚研究中的应用

集合群落(metacommunity)是指多个潜在相互作用的物种通过它们之间的扩散而连接在一起的一组局域群落,目前已成为斑块生境下生物群落结构、格局和动态的重要理论基础之一。斑块动态、物种排序、群体效应和中性模型等4种理论模型,可用于解释不同情形下集合群落内物种的迁移状况,描述集合群落的动态。可采用群落结构或生态学机制等途径,来阐述所研究的群落是属于哪一种特定的集合群落类型。集合群落可用于研究河流鱼类群聚,解释鱼类的群落结构等问题。另外本文还结合我国水域生态环境及水生生物现状,对今后集合群落的研究作了展望。     

Microhabitat preference constrains invasive spread of non-native natal grass (<Emphasis Type="Italic">Melinis repens</Emphasis>)

Preventing the establishment of a non-native species is critical for ensuring the species does not become invasive, yet most non-native species will have little impact on their environment. Despite this, little is known about what influences whether a species will remain relatively benign, or whether it will cause economic or ecological harm. Understanding a plant’s microhabitat provides insight into the necessary conditions for establishment and the current distribution limitations of a population. We investigated microhabitat preference of the non-native natal grass (Melinis repens (Willd.) Zizka) in Florida scrub using microhabitat sampling to measure vegetation composition. We examined the extent to which microhabitats were associated with natal grass presence and biomass in invaded disturbed scrub and roadside plots using backwards stepwise logistic regression and general linear models to identify significant microhabitat variables. We further compared these plots with those in undisturbed, uninvaded scrub to characterize vegetation across habitat types, and used our model to predict the probability of natal grass invasion in undisturbed scrub. Natal grass preferred microhabitats with high litter volume and distance to shrubs and intermediate cactus, graminoid, and vine cover. Roadside natal grass achieved higher biomass and was less microhabitat limited than disturbed scrub natal grass. We determined that undisturbed scrub plots represent distinct microhabitats that natal grass is unlikely to invade. Microhabitat sampling provides land-managers a non-intrusive technique to assess potential habitat suitability based non-native plant preferences before a costly invasion occurs.