Variation in the impact of non‐native seaweeds along gradients of habitat degradation: a meta‐analysis and an experimental test

Biological invasions are acknowledged among the main drivers of global changes in biodiversity. Despite compelling evidence of species interactions being strongly regulated by environmental conditions, there is a dearth of studies investigating how the effects of non‐native species vary among areas exposed to different anthropogenic pressures. Focusing on marine macroalgae, we performed a meta‐analysis to test whether and how the direction and magnitude of their effects on resident communities and species varies in relation to cumulative anthropogenic impact levels. The relationship between human impact levels and non‐native species impact intensity emerged only for a reduced subset of the response variables examined. Yet, there was a trend for the effects of non‐native species on community biomass and abundance and on species abundance to become less negative at heavily impacted sites. By contrast, the magnitude of negative effects of seaweed on community evenness tended to increase with human impact levels. The hypothesis of decreasing severity of invader’ impacts along a gradient of habitat degradation was also tested experimentally at a regional scale by comparing the effects of the removal of non‐native alga, Caulerpa cylindracea, on resident assemblages among rocky reefs exposed to different anthropogenic pressures. Assemblages at urban and pristine site did not differ when invaded, but did so when C. cylindracea was removed. Our results suggest that, despite the generally weak relationship between human impacts levels and non‐native species impacts, more negative impacts can be expected in less stressful environments (i.e. less degraded or pristine sites), where competitive interactions are presumably the driving force structuring resident communities. Implementing strategies for controlling the establishment of non‐native seaweeds should be, thus, considered a priority for preserving biodiversity in relatively pristine areas. On the other hand, control of invaders at degraded sites could be warranted to lessen their role as propagule sources. Synthesis Local anthropogenic stressors that severely alter biotic and abiotic conditions may underpin context‐dependency in the impacts of biological invasions. We used a meta‐analysis and an experimental test to examine the relationship between cumulative human impacts and ecological impact of non‐native seaweeds on resident assemblages. Our results suggest that more negative impacts of non‐native seaweeds on the abundance and biomass of resident assemblages can be expected in less degraded or pristine sites. Possibly, stronger impacts prevail at pristine sites, where assemblages are mainly structured by biotic interactions. Hence, management efforts should be mostly directed to prevent the establishment and spread of non‐native seaweeds in pristine areas. On the other hand, weak, but positive effects of seaweeds at the most degraded sites add to the ongoing debate on the role of non‐native species in rehabilitation plans.     

Stocked trout have minimal effects on littoral invertebrate assemblages of productive fish‐bearing lakes: a whole‐lake BACI study

1. Rainbow Trout (Oncorhynchus mykiss [Walbaum]) is commonly stocked as a sport fish throughout the world but can have serious negative effects on native species, especially in headwater systems. Productive fish‐bearing lakes represent a frequently stocked yet infrequently studied system, and effects of trout in these systems may differ from those in headwater lakes. 2. We used a Before‐After Control‐Impact (BACI) design to determine how stocked trout affected assemblage‐level and taxon‐level biomass, abundance and average length of littoral invertebrates in a stocked lake relative to three unstocked control lakes in the boreal foothills of Alberta, Canada. Lakes were studied 1 year before and for 2 years after stocking. Because characteristics of productive fish‐bearing lakes should buffer impacts of introduced fish, we predicted that trout would not affect assemblage‐level structure of littoral invertebrates but might reduce the abundance or average length of large‐bodied taxa frequently consumed by trout. 3. Relative to the unstocked control lakes, biomass, but not abundance, of the littoral invertebrate assemblage was affected indirectly by trout through increases of some taxa after trout stocking. At the individual taxon‐level, trout stocking did not affect most (23 of the 27) taxa, with four taxa increasing in abundance or biomass after stocking. Only one taxon, Chironomidae, showed evidence of size‐selective predation by trout, being consumed frequently by trout and decreasing significantly in average length after stocking. 4. Our results contrast with the strong negative effects of trout stocking on invertebrate assemblages commonly reported from headwater lakes. A combination of factors, including large and robust native populations of forage fish, the generalised diet of trout, overwinter aeration, relatively high productivity and dense macrophyte beds, likely works in concert to reduce potentially negative effects of stocked trout in these systems. As such, productive, fish‐bearing lakes may represent a suitable system for trout stocking, especially where native sport fish populations are lacking.     

When invasion may not be harmful: niche relations in a lizard assemblage

Some studies have suggested that non‐native species invasions may threaten local diversity by creating homogenized environments. However, many studies have been based on limited or anecdotal data, and/or have failed to consider the influence of habitat modification together with possible influences of non‐native species on native ones. Hemidactylus mabouia (Squamata, Gekkonidae) likely invaded natural environments in Brazil hundreds of years ago. Yet, little is known about whether it affects native lizard fauna. We tested whether H. mabouia negatively influences native lizard species richness and abundance on a regional scale and locally through niche overlap. We analyzed species abundance and richness of nine lizard assemblages, in five of which H. mabouia occurred. We evaluated niche overlap of species in a lizard assemblage with high H. mabouia abundance through null models. Niche axes included spatial use, temporal activity and diet. Although species abundance did not differ among sites with and without the invasive species, the presence of H. mabouia seems constrained to the richer assemblages sampled. We observed significantly higher niche overlap in spatial (?_obs = 0.63; ?_exp = 0.37; P_obs ≥ P_exp = 0.0002) and trophic axes (?_obs = 0.46; ?_exp = 0.17; P_obs ≥ P_exp < 0.001), but not in activity. When we considered all axes (three‐dimensional niche), there was no overlapping among the lizard species. Our findings did not support the hypothesis that this non‐native species negatively influences other sympatric lizard species.     

An invasive fish promotes invasive plants in Minnesota lakes

1. Biological invasions can greatly alter ecological communities, affecting not only the diversity and abundance but also composition of invaded assemblages. This is because invaders’ impacts are mediated by characteristics of resident species: some may be highly sensitive to invader impacts while others are unaffected or even facilitated. In some cases, this can result in invasive species promoting further invasions; in particular, herbivory by introduced animals has been shown to disproportionately harm native plants, which can indirectly benefit non-native plants. Here, we investigated whether such patterns emerged through the effects of an invasive fish species on lake plant communities.
2. Specifically, we tested whether invasion of Minnesota (U.S.A.) lakes by Cyprinus carpio (common carp), an omnivorous, benthivorous fish known to reduce abundance and richness of aquatic plants, differentially affected native versus non-native plant species. We applied statistical models to a large, long-term monitoring dataset (206 macrophyte taxa recorded in 913 lakes over a 20-year time period) to test whether carp altered community composition, to identify which macrophyte species were most sensitive to carp and determine whether species characteristics predicted carp sensitivity, and to characterise consequences of carp invasion on lake-level vegetation attributes.
3. We found that carp exerted strong selective pressure on community composition. Native macrophytes, those with a more aquatic growth form, and those considered less tolerant of disturbance (i.e. higher coefficients of conservatism) were more sensitive to carp. Conversely, no introduced macrophytes exhibited sensitivity to carp and all had higher probabilities of occurrence as carp abundance increased. The net effect of carp invasion was a shift toward less species-rich plant communities characterised by more non-native and disturbance-tolerant species.
4. These results have several implications for conservation and management. First, they reinforce the need to prevent further spread of carp outside of their native range. Where carp have already established, their control should be incorporated into efforts to restore aquatic vegetation; this may be an essential step for recovering particular plant species of high conservation importance. Furthermore, reducing carp abundance could have ancillary benefits of reducing dominance by invasive plant species. Lastly, where carp cannot be eliminated, managers should target native macrophytes that are relatively tolerant of carp in shoreline plantings and other revegetation efforts.

     

Multi‐trophic impacts of an invasive aquatic plant

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Introduced birds in urban remnant vegetation: Does remnant size really matter?

Abstract Introduced birds are a pervasive and dominant element of urban ecosystems. We examined the richness and relative abundance of introduced bird species in small (1–5 ha) medium (6–15 ha) and large (>15 ha) remnants of native vegetation within an urban matrix. Transects were surveyed during breeding and non‐breeding seasons. There was a significant relationship between introduced species richness and remnant size with larger remnants supporting more introduced species. There was no significant difference in relative abundance of introduced species in remnants of different sizes. Introduced species, as a proportion of the relative abundance of the total avifauna (native and introduced species), did not vary significantly between remnants of differing sizes. There were significant differences in the composition of introduced bird species between the different remnant sizes, with large remnants supporting significantly different assemblages than medium and small remnants. Other variables also have substantial effects on the abundance of introduced bird species. The lack of significant differences in abundance between remnant sizes suggests they were all equally susceptible to invasion. No patches in the urban matrix are likely to be unaffected by introduced species. The effective long‐term control of introduced bird species is difficult and resources may be better spent managing habitat in a way which renders it less suitable for introduced species (e.g. reducing areas of disturbed ground and weed dominated areas).     

Relationships between the spread of Caulerpa filiformis and fish communities on temperate rocky reefs

The previously sub‐dominant native marine macrophyte Caulerpa filiformis is now dominant on many sub‐tidal rocky reefs in New South Wales (NSW), Australia and is expanding its distribution. As C. filiformis is highly chemically defended and structurally different to co‐occurring habitat‐forming macrophytes, two key attributes that govern fish assemblages, we hypothesized that fish assemblages, particularly herbivorous fishes, would be different at sites where C. filiformis occurred from where it was previously absent and within sites, fish community structure would be correlated to the cover of C. filiformis. We investigated these hypotheses by determining reef‐associated fish assemblage attributes (assemblage structure, species richness, total abundance, Shannon‐Weiner diversity, abundance of herbivorous species) along transects within sites where C. filiformis was present and absent. Surprisingly, despite large patches and very high densities of C. filiformis on the reefs we sampled, at larger spatial scales (i.e., among sites) no fish assemblage metrics differed between sites with large stands of C. filiformis and sites without the alga. Moreover the abundance of one dominant herbivore, the rock cale Aplodactylus lophodon, was greater at sites within large beds of C. filiformis. At smaller spatial scales, however, i.e. within sites where C. filiformis was present, fish assemblages did vary as a function of C. filiformis cover along transects, although this was not consistent across sampling times. Overall, our results suggest that the potential effects of the spread of this alga on faunal communities warrants further investigation.     

Spatially variable response of native fish assemblages to discharge,predators and habitat characteristics in an arid‐land river

1. Fish assemblages and habitats were sampled annually at fixed sites in three tributaries of the Gila River catchment over a 21‐year span that included prolonged low‐ and high‐flow periods. Model selection was used to evaluate responses of seven native fishes with variable ecological traits (four small‐bodied cyprinids, one large‐bodied cyprinid, and two large‐bodied catostomids) to mean annual discharge and predacious non‐native fishes across the three sites. We also compared habitat use and overlap of native and non‐native fishes to identify potential for negative interactions among species. 2. Assemblage structure (species abundance and richness) and recruitment of native species was strongly and primarily affected by mean annual discharge and secondarily by location and densities of non‐native predators (mainly the centrarchid Micropterus dolomieui). 3. Densities of age‐0 catostomids and small‐bodied cyprinids were positively associated with discharge, and this pattern was strongest in the tributary with the lowest densities of non‐native predators. Absence or extreme low abundance of natives during low‐flow years was most pronounced at the sites where non‐native predators were comparatively common. Densities of adults of large‐bodied native species also varied by site, but often were positively associated with densities of non‐native predators. 4. Spatially variable responses of native fish assemblages indicated that the persistence of native fishes could be jeopardized if key habitats were lost or flow regimes unnaturally altered, particularly during low‐flow conditions when recruitment of native fishes is low and predation by non‐natives is high. Large‐bodied species may be less vulnerable to multiple years of poor conditions because adults are able to avoid predation by non‐natives and thus can rely on occasional high discharge years for successful recruitment. 5. As in other arid‐land streams, native fish assemblages of the Gila River Basin continue to decline. Our results indicate that conservation requires specific knowledge and consideration of physical influences as well as life‐history attributes of native and non‐native fishes.     

THE INFLUENCE OF SUBMERGED MACROPHYTES ON SEDIMENTARY DIATOM ASSEMBLAGES1

Submerged macrophytes are a central component of lake ecosystems; however, little is known regarding their long‐term response to environmental change. We have examined the potential of diatoms as indicators of past macrophyte biomass. We first sampled periphyton to determine whether habitat was a predictor of diatom assemblage. We then sampled 41 lakes in Quebec, Canada, to evaluate whether whole‐lake submerged macrophyte biomass (BiomEpiV) influenced surface sediment diatom assemblages. A multivariate regression tree (MRT) was used to construct a semiquantitative model to reconstruct past macrophyte biomass. We determined that periphytic diatom assemblages on macrophytes were significantly different from those on wood and rocks (ANOSIM R = 0.63, P < 0.01). A redundancy analysis (RDA) of the 41‐lake data set identified BiomEpiV as a significant (P < 0.05) variable in structuring sedimentary diatom assemblages. The MRT analysis classified the lakes into three groups. These groups were (A) high‐macrophyte, nutrient‐limited lakes (BiomEpiV ≥525 μg · L^?1; total phosphorus [TP] <35 μg · L^?1; 23 lakes); (B) low‐macrophyte, nutrient‐limited lakes (BiomEpiV <525 μg · L^?1; TP <35 μg · L^?1; 12 lakes); and (C) eutrophic lakes (TP ≥35 μg · L^?1; six lakes). A semiquantitative model correctly predicted the MRT group of the lake 71% of the time (P < 0.001). These results suggest that submerged macrophytes have a significant influence on diatom community structure and that sedimentary diatom assemblages can be used to infer past macrophyte abundance.     

Inferring past zooplanktivorous fish and macrophyte density in a shallow lake: application of a new regression tree model

1. Eutrophication has a profound effect on the biological structure and function of shallow lakes, altering the composition and abundance of submerged macrophyte and fish assemblages. Relatively little is known, however, about decadal to centennial‐scale change in these important aspects of shallow lake ecology. 2. Established palaeolimnological inference models are limited to reconstructing a single variable. As macrophyte and zooplanktivorous fish abundance exert dual and interacting controls on cladoceran assemblages a single variable inference model may contain significant error. To obviate this problem, we applied a new cladoceran‐based multivariate regression tree (MRT) model to cladoceran subfossil assemblages from dated cores from a small shallow lake (Felbrigg Lake, U.K.) to assess long‐term change in fish and submerged macrophyte abundance. Plant macrofossil, chironomid and mollusc subfossil assemblages were also analysed to track changes in biological structure and function and to evaluate the inferences of the MRT model. 3. Over the 200+ year period covered by the sediment cores, there was good agreement in the timing and nature of ecological change reflected by the plant macrofossil, mollusc, chironomid and cladoceran data. The sediment sequence was divided into three dated zones: c. 1797–1890, c. 1890–1954 and c. 1954–present. Prior to 1890 plant‐associated mollusc, cladoceran and chironomid assemblages indicated a species‐rich macrophyte community; a scenario confirmed by the plant macrofossil data. From c. 1890 to 1954 macrophyte‐associated species of all three invertebrate groups remained abundant but the proportion of pelagic cladocerans rose. Post‐1954 mollusc and chironomid assemblages changed to sediment associated detrital feeders and the proportion of pelagic cladoceran taxa increased further. 4. The cladoceran‐based MRT model indicated a long period of stability, c. 1790–1927, characterised by abundant submerged macrophytes and zooplanktivorous fish. From c. 1927 to 1980, the MRT model inferred a decline in zooplanktivorous fish density (ZF) but relative stability in August macrophyte abundance. From 1980 to 2000, an increase in zooplanktivorous fish was inferred tallying well with available data on the fish population (since the 1970s), which indicated extirpation of perch in the 1970s and a subsequent increase in the rudd population. The model inferred little change in August macrophyte abundance until post‐c. 1980 at which point it indicated a decline. The surface sediment assemblage was placed in MRT group A, where submerged plants are absent or very rare in late summer in good agreement with current conditions at the site. 5. The MRT model, applied here for the first time, appears to have successfully tracked changes in macrophyte abundance and ZF over the last 200 years at Felbrigg Lake. The inferences agreed with historical observations on the fish community and the supporting palaeolimnological data. Given that multiple structuring forces shape most biological communities, the application of a model capable of allowing for this represents a significant advance in palaeolimnology.     

Turning back the tide? Local-scale impacts of climate change may have positive effects by restoring natural riverine habitat and reducing invasive fish density

1. Global biodiversity is increasingly threatened by habitat loss, climate change, and biological invasion. However, predictions of impacts on native fauna are hampered by an inadequate knowledge of how these factors interact and how climate change will affect the distribution, abundance, and behaviour of both native and invasive species, not least as most predictions are based on the long-term effects of temperature alone.
2. Here, we present a case study illustrating how local-scale climate change impacts (increased temperature, reduced rainfall, shifts in peak rainfall) affected the hydrology of a channelised lowland European river (reduced flow, reduction in flood events, increased siltation, macrophyte growth), allowing native fish species to recolonise the bankside zone and reducing the density of invasive round goby Neogobius melanostomus by removing its preferred habitat.
3. While most studies predict long-term negative impacts on global fish populations, some suggest potential direct and indirect benefits at a local scale. We are of the opinion that, at a local scale, climate change impacts on fish will be more nuanced and complex than long-term predictions suggest, resulting in both positive and negative effects, with consolation prizes in the face of larger losses. While impacts on fish will differ between regions and/or continents, depending on the specific impacts of climate change, identification of positive effects will be essential in clarifying long-range forecasts and identifying management procedures for mitigating overall negative impacts.

     

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.     

The negative ecological impacts of a globally introduced species decrease with time since introduction

While there is a long‐history of biological invasions and their ecological impacts have been widely demonstrated across taxa and ecosystems, our knowledge on the temporal dynamic of these impacts remains extremely limited. Using a meta‐analytic approach, we investigated how the ecological impacts of non‐native brown trout (Salmo trutta), a model species with a 170‐year‐long and well‐documented history of intentional introductions across the globe, vary with time since introduction. We first observed significant negative ecological impacts immediately after the species introduction. Second, we found that the negative ecological impacts decrease with time since introduction and that the average ecological impacts become nonsignificant more than one century after introduction. This pattern was consistent across other ecological contexts (i.e., geographical location, levels of biological organization, and methodological approach). However, overall negative ecological impacts were more pronounced at the individual and population levels and in experimental studies. While the mechanisms leading to this decrease remain to be determined, our results indicate that rapid response of native organisms (e.g. adaptation, but also local extinction) may play an important role in this dynamic. Changes in native species traits and local extinction can have important conservation implications. Therefore, we argue that the decline of the negative ecological impacts over time should not be used as an argument to neglect the negative impacts of biological invasions.     

The simultaneous inference of zooplanktivorous fish and macrophyte density from sub‐fossil cladoceran assemblages: a multivariate regression tree approach

1. Quantitative palaeolimnology has traditionally sought to quantify species‐environment relationships to use alterations in biological assemblages to reflect past environmental change. Transfer functions have used regression techniques, such as weighted averaging, to define taxon optima and tolerance for a single chemical or biological variable. 2. Cladoceran assemblages and their sub‐fossil remains in shallow lakes are shaped by a combination of interacting factors. Partial constrained ordination of sub‐fossil cladoceran assemblages from 39 shallow lakes (29 in Norfolk, U.K. and 10 in Denmark) indicated that both zooplanktivorous fish (ZF) density and submerged macrophyte abundance significantly influenced community composition. These dual structuring forces precluded the use of a transfer function as one of the key assumptions of this approach was not met, namely that environmental variables apart from the variable being modelled have negligible influence on species distribution or that there is a linear relationship between the two. Separate transfer functions for ZF and macrophyte abundance were developed but had poor performance diagnostics with low bootstrapped r², high root mean square error of prediction (RMSEP) and large bias. 3. To obviate the problem of multiple structuring forces a multivariate regression tree (MRT) was employed, which allows for more than one explanatory variable within a model. The MRT analysis defined six groups with discrete ranges of ZF and macrophyte densities. The technique identified critical values or ‘break points’ in ZF and macrophyte abundances which result in significant alterations in the sub‐fossil cladoceran assemblage. In addition, the MRT groups had different summer mean values for chlorophyll‐a, Secchi depth, total phosphorus and nitrate‐nitrogen. 4. The predictive abilities of the model were assessed by comparing the observed versus predicted MRT group membership. In general group membership was reliably predicted, suggesting sub‐fossil cladoceran assemblages reliably reflect ZF and macrophyte density in shallow lakes. For a relatively small number of sites there were differences between the observed and predicted MRT group membership. These failures of prediction may result, at least in part, from the disparity of the time period represented by the environmental data and the surface sediment cladoceran assemblage.     

Habitat specificity of butterflies along urban environmental gradients in Tama City,Tokyo

Butterfly assemblages were monitored by transect counts in a riverine area along Tamagawa River (RIV), a residential area on the plain (RES1), a residential area on the hill (RES2), the core area of the city (COR), the Tama Experimental Station of Forestry and Forest Products Research Institute (TES) and Tokyo Metropolitan Sakuragaoka Park (MSP) in Tama City, Tokyo, in 2005. The butterfly assemblages in forest‐dominated TES and MSP were more species‐rich than those in the other areas. The assemblage in the grassland‐dominated RIV was characterized by the highest abundance of individuals. Species richness and abundance were lowest in COR. Ordination of the areas by detrended correspondence analysis placed RIV, MSP and TES in increasing order of scores along axis 1, and RES1, RES2 and COR had higher scores along axis 2 than RIV, TES and MSP. In axis 1, grassland species had low scores and forest species high scores; the two groups were generally separated, coinciding with Tanaka's classification. However, the scores for two “forest species”, Papilio xuthus and Ypthima argus, were low and those for two “grassland species”, Potanthus flavus and Anthocharis scolymus, were high, challenging the validity of the classification. Most species recorded in this study were “seminatural type”, with relatively few “natural type” species, according to Sunose's classification. Most “urban type” species occurred in several different habitats and were not specific to highly human‐dominated RES1, RES2 and COR.     

How often are invasion-induced ecological impacts missed?

Managers and policy makers depend on empirical research to guide and support biosecurity measures that mitigate introduced species’ impacts. Research contributing to this knowledge base generally uses null hypothesis significance testing to determine the significance of data patterns. However, reliance on traditional statistical significance testing methods, combined with small effect and sample size and large variability inherent to many impact studies, may obscure effects on native species, communities or ecosystems. This may result in false certainty of no impact. We investigated potential Type II error rates and effect sizes for 31 non-significant empirical evaluations of impact for introduced algal and crustacean species. We found low power consistently led to acceptance of Type II errors at rates 5.6–19 times greater than Type I errors (despite moderate to large effect sizes). Our results suggest that introduced species for which impact studies have statistically non-significant outcomes (often interpreted as “no impact”) may potentially have large impacts that are missed due to small sample or effect sizes and/or high variation. This alarming willingness to “miss” impacts has severe implications for conservation efforts, including under-managing species’ impacts and discounting the costs of Type II errors.     

Invasive plants negatively impact native,but not exotic,animals

Despite our growing understanding of the impacts of invasive plants on ecosystem structure and function, important gaps remain, including whether native and exotic species respond differently to plant invasion. This would elucidate basic ecological interactions and inform management. We performed a meta‐analytic review of the effects of invasive plants on native and exotic resident animals. We found that invasive plants reduced the abundance of native, but not exotic, animals. This varied by animal phyla, with invasive plants reducing the abundance of native annelids and chordates, but not mollusks or arthropods. We found dissimilar impacts among “wet” and “dry” ecosystems, but not among animal trophic levels. Additionally, the impact of invasive plants increased over time, but this did not vary with animal nativity. Our review found that no studies considered resident nativity differences, and most did not identify animals to species. We call for more rigorous studies of invaded community impacts across taxa, and most importantly, explicit consideration of resident biogeographic origin. We provide an important first insight into how native and exotic species respond differently to invasion, the consequences of which may facilitate cascading trophic disruptions further exacerbating global change consequences to ecosystem structure and function.     

Resemblance in phylogenetic diversity among ecological assemblages

Questions: How can a resemblance (similarity or dissimilarity) measure be formulated to include information on both the evolutionary relationships and abundances of organisms, and how does it compare to measures lacking such information? Methods: We extend the family of Phylogenetic Diversity (PD) measures to include a generalized method for calculating pair‐wise resemblance of ecological assemblages. Building on previous work, we calculate the matching/mismatching components of the 2 × 2 contingency table so as to incorporate information on both phylogeny and abundance. We refer to the class of measures so defined as “PD resemblance” and use the term “SD resemblance” for the traditional class of measures based on species diversity alone. As an illustration, we employ data on the diversity and stem density of shrubs of Toohey Forest, Australia, to compare PD resemblance to its SD resemblance equivalent for both incidence and abundance data. Results: While highly correlated, PD resemblance consistently measures assemblages as more similar than does SD resemblance, and tends to “smooth out” the otherwise skewed and truncated distribution of pair‐wise resemblance indices of our high‐turnover data set, resulting in nMDS ordinations with lower stress. Randomization of species distributions across assemblages indicates that phylogeny has made a significant contribution to the ordination pattern. Conclusions: PD resemblance measures, in addition to providing an evolutionary perspective, have great potential to improve distance‐based analyses of community patterns, particularly if species responses to ecological gradients are unimodal and phylogenetically conserved.     

Aquatic macrophytes, macroinvertebrate associations and water levels in a lowland Tasmanian river

Aquatic macrophytes are a common habitat for macroinvertebrates and may occupy depth zones in the littoral region of lowland rivers. Studies have indicated that different species of macrophyte typically support different assemblages, abundances and numbers of species of macroinvertebrates. This has often been attributed to differences in the dissectedness of stems and leaves of the macrophytes, resulting in differences in the surface area and/or the number of microhabitats available to invertebrates. I set out to measure the abundance and taxonomic richness and to describe the macroinvertebrate assemblages associated with three species of aquatic macrophyte in a pool in the Macquarie River, Tasmania and to examine responses of these variables to changes in water levels over summer. The macrophyte species sampled wereMyriophyllum simulans/variifolium, Triglochin procera} and Eleocharis sphacelata, each one differing in the dissectedness of its stems and leaves and its location in the littoral zone. Whereas the greatest abundance of macroinvertebrates was found associated in all months (i.e. at all water levels) with the structurally complex and shallowest macrophyte species, Myriophyllum, the number of taxa associated with this species was in several cases lower than for the structurally simpler and deeper water Triglochin and Eleocharis. While water depth and total plant biomass of samples were often correlated with invertebrate abundance and richness, these relationships were different for each macrophyte species. Of the nine most common invertebrate taxa collected from all samples, the abundances of more than half showed consistent differences among macrophyte species across months, two showed differences among macrophytes, but with an interaction with month and two showed no differences among macrophytes. There were major differences in the invertebrate assemblages associated with each macrophyte species in any one month, however, there was also a large turnover of taxa associated with the species of macrophytes from one month to the next. Changes in water level and concomitant changes in environmental variables are suggested as factors influencing the invertebrate fauna in the littoral zone of the pool of the Macquarie River. It is thus important for river managers to be aware that species of macroinvertebrates are not evenly distributed across species of macrophyte and that water levels and their influence on macrophytes as invertebrate habitat may play an integral part in determining the abundance, richness and assemblage of invertebrates in rivers.     

Native species richness buffers invader impact in undisturbed but not disturbed grassland assemblages

Many systems are prone to both exotic plant invasion and frequent natural disturbances. Native species richness can buffer the effects of invasion or disturbance when imposed in isolation, but it is largely unknown whether richness provides substantial resistance against invader impact in the face of disturbance. We experimentally examined how disturbance (drought/burning) influenced the impact of three exotic invaders (Centaurea stoebe, Linaria dalmatica, or Potentilla recta) on native abundance across a gradient of species richness, using previously constructed grassland assemblages. We found that invaders had higher cover in experimentally disturbed plots than in undisturbed plots across all levels of native species richness. Although exotic species varied in cover, all three invaders had significant impacts on native cover in disturbed plots. Regardless of disturbance, however, invader cover diminished with increasing richness. Invader impacts on native cover also diminished at higher richness levels, but only in undisturbed plots. In disturbed plots, invaders strongly impacted native cover across all richness levels, as disturbance favoured invaders over native species. By examining these ecological processes concurrently, we found that disturbance exacerbated invader impacts on native abundance. Although diversity provided a buffering effect against invader impact without disturbance, the combination of invasion and disturbance markedly depressed native abundance, even in high richness assemblages.