External morphology explains the success of biological invasions

Biological invasions have become major players in the current biodiversity crisis, but realistic tools to predict which species will establish successful populations are still unavailable. Here we present a novel approach that requires only a morphometric characterisation of the species. Using fish invasions of the Mediterranean, we show that the abundance of non‐indigenous fishes correlates with the location and relative size of occupied morphological space within the receiving pool of species. Those invaders that established abundant populations tended to be added outside or at the margins of the receiving morphospace, whereas non‐indigenous species morphologically similar to resident ones failed to develop large populations or even to establish themselves, probably because the available ecological niches were already occupied. Accepting that morphology is a proxy for a species' ecological position in a community, our findings are consistent with ideas advanced since Darwin's naturalisation hypothesis and provide a new warning signal to identify invaders and to recognise vulnerable communities.     

Understanding the long-term effects of species invasions

We describe here the ecological and evolutionary processes that modulate the effects of invasive species over time, and argue that such processes are so widespread and important that ecologists should adopt a long-term perspective on the effects of invasive species. These processes (including evolution, shifts in species composition, accumulation of materials and interactions with abiotic variables) can increase, decrease, or qualitatively change the impacts of an invader through time. However, most studies of the effects of invasive species have been brief and lack a temporal context; 40% of recent studies did not even state the amount of time that had passed since the invasion. Ecologists need theory and empirical data to enable prediction, understanding and management of the acute and chronic effects of species invasions.     

Seagrass feeding choices and digestive strategies of the herbivorous fish Sarpa salpa

This is the first study investigating the plant–herbivore interaction between Sarpa salpa, which has overgrazed seagrass transplants in Portugal, and the seagrasses Cymodocea nodosa, Zostera marina and Zostera noltii, which have been considered for restoration. When offered the choice between the three seagrasses in outdoor tanks, adult S. salpa clearly preferred Z. noltii. Testing the seagrasses separately, mean ± s.d. feeding rates ranged from 21 ± 11 g seagrass fresh mass kg^?1 fish mass day^?1 for Z. marina to 32 ± 9 g seagrass fresh mass kg^?1 fish mass day^?1 for C. nodosa and 40 ± 11 g seagrass fresh mass kg^?1 fish mass day^?1 for Z. noltii (temperature = 16° C). Food‐processing rate in S. salpa did not differ between seagrasses, and there was no evidence of a regulation of processing rate according to food intake. Seagrasses differed substantially in nitrogen content and C:N, with C. nodosa containing the highest nitrogen content and lowest C:N (2·5 ± 0·1% and 14·0 ± 1·0), followed by Z. noltii (2·1 ± 0·1% and 17·0 ± 1·0) and Z. marina (1·4 ± 0·1% and 26·0 ± 2·0). Food‐processing rate in S. salpa and the nutritional value of the seagrasses were not correlated with the observed feeding preference and rate. The study suggests that C. nodosa and Z. marina are less at risk of overgrazing by S. salpa and might thus be preferable to Z. noltii for seagrass restoration in areas with noticeable abundances of this fish.     

Importance of feeding ecology to the reproductive success of Blackbirds Turdus merula nesting in rural habitats

The feeding ecology of Blackbirds Turdus merula breeding in contiguous woodland and farmland habitats was studied over three years. The aim of the study was to investigate how reproductive success was influenced by nestling diet and the provisioning rates of parents feeding nestlings. Parental provisioning rates increased with brood size, and consequently individual nestlings were no lighter in larger broods. None of the environmental factors measured had strong effects on parental provisioning rate. The nestling diet was dominated by caterpillars and earthworms, the former occurring in a short period in the middle of the breeding season. The availability of earthworms was higher in woodland and was dependent on rainfall in farmland. Nestling mass and provisioning rates were marginally higher under predominantly earthworm diets. Nestling mass increased with rainfall in farmland only, and was higher in farmland than in woodland or woodland-edge, although it is doubtful whether this result is of any significance for fledgling survival. Overall, Blackbirds were able to provision their nestlings adequately throughout the breeding season across a range of conditions. There was no evidence to suggest that reproductive success was constrained by aspects of feeding ecology within the natural range of brood size.     

Paradox lost: genetic diversity and the success of aquatic invasions

There is mounting evidence that reduced genetic diversity in invasive populations is not as commonplace as expected. Recent studies indicate that high propagule vectors, such as ballast water and shellfish transplantations, and multiple introductions contribute to the elimination of founder effects in the majority of successful aquatic invasions. Multiple introductions, in particular, can promote range expansion of introduced populations through both genetic and demographic mechanisms. Closely related to vectors and corridors of introduction, propagule pressure can play an important role in determining the genetic outcome of introduction events. Even low-diversity introductions have numerous means of avoiding the negative impact of diversity loss. The interaction of high propagule vectors and multiple introductions reveal important patterns associated with invasion success and deserve closer scrutiny.     

Eco-immunology of fish invasions: the role of MHC variation

The relationship between invaders and the pathogens encountered in their new environment can have a large effect on invasion success. Invaders can become free from their natural pathogens and reallocate costly immune resources to growth and reproduction, thereby increasing invasion success. Release from enemies and relaxation of selective pressures could render newly founded populations more variable at immune-related genes, such as the major histocompatibility complex (MHC), particularly when they have different origins. Using rainbow and brown trout, two of the world’s most successful fish invaders, we tested the general hypothesis that invaders should display high intrapopulation immunogenetic diversity and interpopulation divergence, due to the interplay between genetic drift and successive waves of genetically divergent introductions. We analysed genetic diversity and signatures of selection at the MHC class II β immune-related locus. In both species, MHC diversity (allelic richness and heterozygosity) for southern hemisphere populations was similar to values reported for populations at their native range. However, MHC functional diversity was limited, and population immunogenetic structuring weaker than that observed using neutral markers. Depleted MHC functional diversity could reflect a decrease in immune response, immune-related assortative mating or selection for resistance to newly encountered parasites. Given that the role of MHC diversity in the survival of these populations remains unclear, depleted functional diversity of invasive salmonids could compromise their long-term persistence. A better understanding of the eco-immunology of invaders may help in managing and preventing the impact of biological invasions, a major cause of loss of biodiversity worldwide.     

Importance of fungi in the diet of Gammarus pulex and Asellus aquaticus I: feeding strategies

The importance of fungi in the trophic biology of the freshwater detritivores Gammarus pulex and Asellus aquaticus was investigated. Inspection of leaves used in feeding trials indicated that whereas A. aquaticus scrapes at the leaf surface, G. pulex bites through the leaf material. Both species discriminated between fungal mycelia, fungally colonized and uncolonized leaf material but, although A. aquaticus selectively consumed fungal mycelia, G. pulex fed preferentially on leaf material. Fungi appear to be an important food source for A. aquaticus and selection of food material was positively correlated with fungal biomass. In contrast, for G. pulex, fungi appear to be more important as modifiers of leaf material. However, no significant correlations were found between food preference and any of the leaf modifications measured.     

Behavioral aspects of biological invasions of alien fish species

The role of the behavior of animals determining the success of invasion to new areas and habitats is analyzed. The majority of examples are related to fish whose distribution attracts increasing attention from both scientists and persons involved in applied fisheries. The contribution of migratory, feeding, defensive, reproductive, and social behavior in “the invasive potential” of the species is considered. The success of the colonization of new biotopes by the species and expansion of its area may not depend so much on its properties as the ability to cover long distances, plasticity in mastering new resources, and stability to pressure from new competitors, predators, and parasites. Some properties of animals are stressed whose role in invasion may be important and is still to be demonstrated. Among them, a special interest for behaviorists is the ability of animals towards learning and towards “innovation activity”, the ability to switch rapidly from a cooperative mode of life to an individual mode of life, and to efficiently explore and to exploit heterogeneous microhabitats.     

塔里木盆地鱼类入侵及区系演变趋势

鱼类入侵是塔里木盆地土著鱼类濒危的关键性影响因素之一,结合历史文献数据及近年调查数据,系统综述了塔里木盆地鱼类入侵现状,并就鱼类入侵引起的鱼类区系均一化问题进行了分析。结果显示,该地区记录鱼类63种,其中土著种19种,入侵种44种。尽管入侵鱼类提升了塔里木盆地鱼类群落属级及科级的多样性水平,但相应的G-F多样性指数却明显下降,表明该区鱼类群落从物种组成简单、分化明显的特点演变为物种成分复杂、分化贫乏的特点。入侵种中源于长江和额尔齐斯河分别有30种(68.2%)和7种(15.9%),源于美欧和亚洲其他区域的有7种(15.9%)。入侵鱼类的进入引起塔里木盆地与其相关区域鱼类区系相似性显著上升(P0.01),并导致该区鱼类区系从单一的中亚高原山区复合体演变为多区系混合体。进一步利用鱼类个体生态矩阵分析方法研究显示,外来种在塔里木盆地适应性广泛,能够占据水域各种栖息生境,显著压缩了土著种的栖息空间,进而通过捕食、食物竞争及其他作用途径严重危及土著鱼类种群的续存,提示须采取有力措施限制入侵种的进一步扩散及对濒危土著物种迅速开展人工保育工作。     

Conceptualising the interactive effects of climate change and biological invasions on subarctic freshwater fish

Climate change and species invasions represent key threats to global biodiversity. Subarctic freshwaters are sentinels for understanding both stressors because the effects of climate change are disproportionately strong at high latitudes and invasion of temperate species is prevalent. Here, we summarize the environmental effects of climate change and illustrate the ecological responses of freshwater fishes to these effects, spanning individual, population, community and ecosystem levels. Climate change is modifying hydrological cycles across atmospheric, terrestrial and aquatic components of subarctic ecosystems, causing increases in ambient water temperature and nutrient availability. These changes affect the individual behavior, habitat use, growth and metabolism, alter population spawning and recruitment dynamics, leading to changes in species abundance and distribution, modify food web structure, trophic interactions and energy flow within communities and change the sources, quantity and quality of energy and nutrients in ecosystems. Increases in temperature and its variability in aquatic environments underpin many ecological responses; however, altered hydrological regimes, increasing nutrient inputs and shortened ice cover are also important drivers of climate change effects and likely contribute to context‐dependent responses. Species invasions are a complex aspect of the ecology of climate change because the phenomena of invasion are both an effect and a driver of the ecological consequences of climate change. Using subarctic freshwaters as an example, we illustrate how climate change can alter three distinct aspects of species invasions: (1) the vulnerability of ecosystems to be invaded, (2) the potential for species to spread and invade new habitats, and (3) the subsequent ecological effects of invaders. We identify three fundamental knowledge gaps focused on the need to determine (1) how environmental and landscape characteristics influence the ecological impact of climate change, (2) the separate and combined effects of climate and non‐native invading species and (3) the underlying ecological processes or mechanisms responsible for changes in patterns of biodiversity.     

The effect of alternative feeding strategies on the feeding motivation of broiler breeder pullets

Due to genetic selection for fast growth and high breast meat yield, commercial strains of broiler chickens and broiler breeders are predisposed to high feed intake; however, feeding broiler breeders ad libitum impairs their health and reproductive performance. Broiler breeders are feed-restricted throughout rearing to maintain health and performance, yet feed restriction results in hunger, feeding frustration and lack of satiety. The objective of this study was to examine the effect of alternative feeding strategies, including feed additives (separately or combined) and a fixed non-daily feeding schedule, on the feeding motivation and welfare of broiler breeders during rearing. At 3 weeks of age, 180 Ross 308 breeder pullets were allocated to 90 cages and fed with one of five isocaloric treatments: (1) daily control diet (control), (2) daily calcium propionate diet (CaP), (3) daily soybean hull diet (SBH), (4) daily alternative diet (alternative: CaP + SBH) and (5) 4/3 control diet (four on-feed days and three non-consecutive off-feed days per week). The CaP diet included calcium propionate at 1.4% from 3 to 6 weeks of age, and at 3.2% from 7 to 12 weeks of age, and the SBH diet contained soybean hulls included at 40%. The alternative diet included both soybean hulls and calcium propionate at the same inclusion rate as the SBH and CaP diets, respectively. Pullets were weighed and scored for feather coverage every week. A feed intake test was conducted at 3, 4, 8, 10 and 11 weeks of age for 10 min during on- and off-feed days. At 12 weeks of age, feather samples were analysed for fault bars. Data were analysed using linear mixed regression models, with cage nested in the models and age as a repeated measure. At 4 weeks of age, pullets fed soybean hull-enriched diets (SBH and alternative diets) and those on the 4/3 schedule had lower feed intake than control pullets (P = 0.02). Feathers from pullets fed the SBH diet had fewer fault bars than those fed the CaP diet (P = 0.04). The results indicated that the inclusion of soybean hulls (alone or combined with calcium propionate) and a 4/3 feeding schedule can reduce feeding motivation of broiler breeders during early rearing.     

植物氮形态利用策略及对外来植物入侵性的影响

氮是影响外来植物入侵性的重要因素之一, 但相关研究多关注土壤氮水平的效应, 较少考虑氮形态的作用。为从土壤氮形态利用的角度阐释外来植物的入侵机制, 本文在植物氮形态利用策略分析的基础上, 综述了外来植物氮形态利用的偏好性及其对入侵性的影响。植物的氮形态利用策略有偏好性和可塑性两种, 这可能与植物对土壤氮形态特性的长期适应有关; 植物不仅可以对土壤氮形态做出响应, 而且还能改造土壤氮形态, 并对改变后的土壤氮形态做出反馈响应。很多外来植物入侵硝态氮占优势的干扰生境, 偏好硝态氮的外来植物与本地植物竞争硝态氮; 而偏好铵态氮的外来植物通过抑制土壤硝化作用, 营造铵态氮环境, 促进自身生长, 同时抑制偏好硝态氮的本地植物生长。然而, 植物氮形态利用策略不是一成不变的, 而是受多种生物和非生物因素共同作用影响的复杂过程, 今后应加强多因素交互作用对外来入侵植物氮形态利用策略的影响及机制研究, 更好地揭示氮形态利用策略, 尤其是氮形态利用的可塑性与外来植物入侵性的关系。     

Tackling aquatic invasions: risks and opportunities for the aquarium fish industry

The aquarium trade is an important and rapidly growing vector for introduced species in the United States. We examined this vector by surveying pet stores in the San Francisco Bay–Delta region to compile a list of aquarium fish species commonly stocked. We identified which of these species might be able to survive in the Bay–Delta, and investigated store representatives’ knowledge and attitudes about biological invasions. A restrictive analysis using conservative estimates of fish temperature tolerances and environmental conditions found that the local aquarium trade includes 5 fish species that can survive in a temperate system such as the Bay–Delta. Under more inclusive parameters, up to 27 fish species met the criteria for survival in the Bay–Delta. We further explored these results by comparing potential invader incidence between different types of stores. In the more restrictive analysis, three national retail chains stocked significantly more potentially invasive species than independent aquarium stores, but there was no difference in the more inclusive analysis. A significantly higher percentage of fish taxa were easily identifiable and well-labeled in chain stores than in independent stores. Most aquarium store representatives indicated willingness to take action to reduce the threat of trade-related introductions, although chain store employees were more willing to assign responsibility for reducing this threat to the aquarium industry than were independent store employees. Management efforts for this vector should focus on (a) improving labeling and identification of fish species in stores, (b) expanding the often spotty data on fish physiological tolerances, especially for saltwater species, (c) educating customers and store employees about the risks posed by pet release, and (d) providing better options for responsible disposal of unwanted fish. Electronic Supplementary Material   The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Elevated success of multispecies bacterial invasions impacts community composition during ecological succession

Successful microbial invasions are determined by a species’ ability to occupy a niche in the new habitat whilst resisting competitive exclusion by the resident community. Despite the recognised importance of biotic factors in determining the invasiveness of microbial communities, the success and impact of multiple concurrent invaders on the resident community has not been examined. Simultaneous invasions might have synergistic effects, for example if resident species need to exhibit divergent phenotypes to compete with the invasive populations. We used three phylogenetically diverse bacterial species to invade two compositionally distinct communities in a controlled, naturalised in vitro system. By initiating the invader introductions at different stages of succession, we could disentangle the relative importance of resident community structure, invader diversity and time pre‐invasion. Our results indicate that multiple invaders increase overall invasion success, but do not alter the successional trajectory of the whole community.     

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.     

Monitoring the feeding activity of individual fish with a demand feeding system

Feeding activity of individual Arctic charr ( Salvelinus alpinus ) was recorded automatically for 29 days using a demand feeding system. Each of three groups of 15 fish was kept in 1 m³ tanks, containing brackish water at 10° C. Individual biting activity was continuously monitored using a PIT-tag (Passive Integrated Transponders) system with unique individual codes. The accuracies of the bite detection system were 91, 93.1 and 99.5% respectively, in the three tanks. In all tanks, most of the individuals (12–14) bit on the releasing trigger a few times during the first 3 days. Thereafter, one or two individuals per tank accounted for almost all of the biting activity. This pronounced shift in bite-number distribution among individuals was probably due to the development of a dominance hierarchy, in which the dominant individuals monopolized the trigger. Growth rates appeared to be highest among high ranking fish. The implications of using demand feeding and PIT-tag devices in feeding studies are discussed.     

Different feeding strategies in Antarctic scavenging amphipods and their implications for colonisation success in times of retreating glaciers

Background

Scavenger guilds are composed of a variety of species, co-existing in the same habitat and sharing the same niche in the food web. Niche partitioning among them can manifest in different feeding strategies, e.g. during carcass feeding. In the bentho-pelagic realm of the Southern Ocean, scavenging amphipods (Lysianassoidea) are ubiquitous and occupy a central role in decomposition processes. Here we address the question whether scavenging lysianassoid amphipods employ different feeding strategies during carcass feeding, and whether synergistic feeding activities may influence carcass decomposition. To this end, we compared the relatively large species Waldeckia obesa with the small species Cheirimedon femoratus, Hippomedon kergueleni, and Orchomenella rotundifrons during fish carcass feeding (Notothenia spp.). The experimental approach combined ex situ feeding experiments, behavioural observations, and scanning electron microscopic analyses of mandibles. Furthermore, we aimed to detect ecological drivers for distribution patterns of scavenging amphipods in the Antarctic coastal ecosystems of Potter Cove. In Potter Cove, the climate-driven rapid retreat of the Fourcade Glacier is causing various environmental changes including the provision of new marine habitats to colonise. While in the newly ice-free areas fish are rare, macroalgae have already colonised hard substrates. Assuming that a temporal dietary switch may increase the colonisation success of the most abundant lysianassoids C. femoratus and H. kergueleni, we aimed to determine their consumption rates (g food x g amphipods^?1 x day^?1) and preferences of macroalgae and fish.

Results

We detected two functional groups with different feeding strategies among scavenging amphipods during carcass feeding: carcass ‘opener’ and ‘squeezer’. Synergistic effects between these groups were not statistically verified under the conditions tested. C. femoratus switched its diet when fish was not available by consuming macroalgae (about 0.2 day^?1) but preferred fish by feeding up to 80% of its own mass daily. Contrary, H. kergueleni rejected macroalgae entirely and consumed fish with a maximal rate of 0.8 day^?1.

Conclusion

This study reveals functional groups in scavenging shallow-water amphipods and provides new information on coastal intraguild niche partitioning. We conclude that the dietary flexibility of C. femoratus is a potential ecological driver and central to its success in the colonisation of newly available ice-free Antarctic coastal habitats.

     

Effect of saline water on early success of amphidromous fish

Organisms that migrate between rivers and the sea inevitably pass through estuarine habitats. Despite the potential importance of salinity and temperature fluctuations for metabolic adaptation, little is known about the impact of environmental changes in estuaries on the survival of residents. Ayu (Plecoglossus altivelis) is a migratory fish that inhabits estuarine brackish water in its early life stages. The recent decline in the abundance of populations ascending into rivers is of concern for local biodiversity. The present study aims to elucidate the ecological processes that determine the early success of Ayu larvae under variable environmental conditions. The effects of salinity and water temperature on the endogenous growth of newly hatched larvae from the same brood were examined experimentally based on morphological and metabolic characteristics. High salinity and high water temperature together appeared to require more energy for larval osmoregulation, resulting in the acceleration of yolk depletion and reduced growth of the notochord. Increasing the osmoregulative cost during the yolk-sac stage resulted in the faster induction of a state of starvation. Seawater is considered to have an adverse effect on the survival of newly hatched larvae, as it lowers the efficiency of foraging and predator avoidance due to an energetic tradeoff. More attention should be paid to the significance of estuarine environments involving brackish waters to ensure the early survival of amphidromous fish such as Ayu.