Sublethal effects on reproduction in native fauna: are females more vulnerable to biological invasion?

Although invasive species are a major threat to survivorship of native species, we know little about their sublethal effects. In soft-sediment marine systems, mat-forming invasive species often have positive effects, facilitating recruitment and enhancing the diversity and abundance of native invertebrates. However, because mat-forming invasive species change the habitat in which they invade, and benthic invertebrates are sensitive to environmental disturbance, important sublethal effects on native species may exist. Using a model marine system we show that the widespread mat-forming invasive alga Caulerpa taxifolia (Vahl) C. Agardh has strong negative effects on the reproductive traits of a native bivalve Anadara trapezia (Deshayes, 1840) (e.g. timing of reproductive development and spawning, and follicle and gamete production) even though the invader has positive effects on recruitment. Moreover, gender specific responses occurred and indicated that females were more susceptible to invasion than males. Our results indicate that sublethal effects of an invasive species on reproductive traits will have severe consequences for fitness of the native species

Morphological traits and density of foundation species modulate a facilitation cascade in Australian mangroves.

Facilitation cascades are critical to the maintenance of biodiversity in a variety of habitats. Through a series of two experiments, we examined how the morphological traits and density of interacting foundation species influence the establishment and persistence of a facilitation cascade in temperate Australian mangrove forests. In this system, mangrove pneumatophores trap the free-living alga, Hormosira banksii, which, in turn, supports dense and diverse assemblages of epifaunal mollusks. The first experiment, which manipulated pneumatophore height and density, revealed that these two traits each had additive negative effects on the establishment, but additive positive effects on the persistence of the cascade. High densities of tall pneumatophores initially served as a physical barrier to algal colonization of pneumatophore plots, but over the longer-term enhanced the retention of algae. The increased algal biomass, in turn, facilitating epifaunal colonization. The second experiment demonstrated that the retention of algae by pneumatophores was influenced more by algal thallus length than vesicle diameter, and this effect occurred independent of pneumatophore height. Our study has extended facilitation theory by showing that the morphological traits and density of basal and intermediary facilitators influence both the establishment and persistence of facilitation cascades. Hence, attempts to use foundation species as a tool for restoration will require an understanding not only of the interactions among these, but also of the key traits that modify interrelationships

Mechanisms influencing the spread of a native marine alga

Like invasive macrophytes, some native macrophytes are spreading rapidly with consequences for community structure. There is evidence that the native alga Caulerpa filiformis is spreading along intertidal rocky shores in New South Wales, Australia, seemingly at the expense of native Sargassum spp. We experimentally investigated the role physical disturbance plays in the spread of C. filiformis and its possible consequences for Sargassum spp. Cleared patches within beds of C. filiformis (Caulerpa habitat) or Sargassum spp. (Sargassum habitat) at multiple sites showed that C. filiformis had significantly higher recruitment (via propagules) into its own habitat. The recruitment of Sargassum spp. to Caulerpa habitat was rare, possibly due in part to sediment accretion within Caulerpa habitat. Diversity of newly recruited epibiotic assemblages within Caulerpa habitat was significantly less than in Sargassum habitat. In addition, more C. filiformis than Sargassum spp. recruited to Sargassum habitat at some sites. On common boundaries between these two macroalgae, the vegetative growth of adjacent C. filiformis into cleared patches was significantly higher than for adjacent Sargassum spp. In both experiments, results were largely independent of the size of disturbance (clearing). Lastly, we used PAM fluorometry to show that the photosynthetic condition of Sargassum spp. fronds adjacent to C. filiformis was generally suppressed relative to those distant from C. filiformis. Thus, physical disturbance, combined with invasive traits (e.g. high levels of recruitment and vegetative growth) most likely facilitate the spread of C. filiformis, with the ramifications being lower epibiotic diversity and possibly reduced photosynthetic condition of co-occurring native macrophytes. © 2014 Zhang et al

Disturbance-mediated facilitation by an intertidal ecosystem engineer

Ecosystem engineers facilitate communities by providing a structural habitat that reduces abiotic stress or predation pressure for associated species. However, disturbance may damage or move the engineer to a more stressful environment, possibly increasing the importance of facilitation for associated communities. In this study, we determined how disturbance to intertidal boulders (i.e., flipping) and the subsequent movement of a structural ecosystem engineer, the tube-forming serpulid worm Galeolaria caespitosa, from the bottom (natural state, low abiotic stress) to the top (disturbed state, high abiotic stress) surface of boulders influenced the importance of facilitation for intertidal communities across two intertidal zones. Theory predicts stronger relative facilitation should occur in the harsher environments of the top of boulders and the high intertidal zone. To test this prediction, we experimentally positioned boulders with the serpulids either face up or face down for 12 months in low and high zones in an intertidal boulder field. There were very different communities associated with the different boulders and serpulids had the strongest facilitative effects on the more stressful top surface of boulders with approximately double the species richness compared to boulders lacking serpulids. Moreover, within the serpulid matrix itself there was also approximately double the species richness (both zones) and abundance (high zone only) of small invertebrates on the top of boulders compared to the bottom. The high relative facilitation on the top of boulders reflected a large reduction in temperature by the serpulid matrix on that surface (up to 10°C) highlighting a key role for modification of the abiotic environment in determining the community-wide facilitation. This study has demonstrated that disturbance and subsequent movement of an ecosystem engineer to a more stressful environment increased the importance of facilitation and allowed species to persist that would otherwise be unable to survive in that environment

Incorporating marine macrophytes in plant–soil feedbacks: Emerging evidence and opportunities to advance the field

Plants can change the biotic and abiotic characteristics of soil, which can in turn affect the growth of plants. For example, changes in below-ground microbial composition by one plant can affect the relative performance of a second plant, in turn affecting the outcome of plant–plant interactions, invasive species success, species abundance distributions, successional processes and plant community composition and diversity. The effects of plant–soil feedbacks on population, community and ecosystem processes in coastal and marine soft-sediment systems have received relatively little attention, yet several recent examples offer compelling evidence for their importance. Marine soft-sediment systems offer an ideal opportunity to test key theoretical predictions for when plant–soil feedbacks may contribute to species coexistence and influence community composition, including the role of phylogenetic distance and relatedness, the potential for intraspecific feedbacks, the importance of environmental context, the influence of climate change and the implications for ecosystem function. Synthesis. There is a documented disconnect between marine and terrestrial research that slows the advancement of generalisable theory. We suggest that bridging this gap in our understanding of plant–soil feedbacks may provide a unifying framework underlying plant community structure in both terrestrial and marine ecosystems

Sublethal effects of a rapidly spreading native alga on a key herbivore

Multiple anthropogenic stressors are causing a global decline in foundation species, including macrophytes, often resulting in the expansion of functionally different, more stressor-tolerant macrophytes. Previously subdominant species may experience further positive demographic feedback if they are exposed to weaker plant–herbivore interactions, possibly via decreased palatability or being structurally different from the species they are replacing. However, the consequences of the spread of opportunistic macrophytes for the local distribution and life history of herbivores are unknown. The green alga, Caulerpa filiformis, previously a subdominant macrophyte on low intertidal-shallow subtidal rock shores, is becoming locally more abundant and has spread into warmer waters across the coast of New South Wales, Australia. In this study, we measured (a) the distribution and abundance of a key consumer, the sea urchin Heliocidaris erythrogramma, across a seascape at sites where C. filiformis has become dominant, (b) performed behavioral field experiments to test the role of habitat selection in determining the local distribution of H. erythrogramma, and (c) consumer experiments to test differential palatability between previously dominant higher quality species like Ecklonia radiata and Sargassum sp. and C. filiformis and the physiological consequences of consuming it. At all sites, urchin densities were positively correlated with distance away from C. filiformis beds, and they actively moved away from beds. Feeding experiments showed that, while urchins consumed C. filiformis, sometimes in equal amounts to higher quality algae, there were strong sublethal consequences associated with C. filiformis consumption, mainly on reproductive potential (gonad size). Specifically, the gonad size of urchins that fed on C. filiformis was equivalent to that in starved urchins. There was also a tendency for urchin mortality to be greater when fed C. filiformis. Overall, strong negative effects on herbivore life-history traits and potentially their survivorship may establish further positive feedback on C. filiformis abundance that contributes to its spread and may mediate shifts from top-down to bottom-up control at locations where C. filiformis has become dominant

Positive and Negative Species Interactions Shape Recruitment Patterns of a Range Expanding Native Alga

A species’ ability to spread is in part governed by the extent to which recipient habitats either resist, tolerate or promote the species’ recruitment. In disturbed marine environments, there is a general trend for the loss of algal canopies, thought to resist invasion, toward algal turf or rock barrens habitat. This study tested whether the spread of the range-expanding native macroalga Caulerpa filiformis was resisted by algal canopies but facilitated by algal turf or barrens habitats. Large-scale field surveys generally supported the predicted recruitment patterns, with C. filiformis recruits being most abundant on turf (but not barrens) and absent under kelp canopies. However, a discrepancy existed between different structural forms of turf, with the positive association holding true only for geniculate corallines, not filamentous turf. Secondly, a laboratory experiment tested whether the physical structure and/or the sediment-trapping properties of coralline turf influenced the recruitment success of C. filiformis. Whilst the structural complexity of turf aided overall recruitment performance (i.e., increased rhizoid production, attachment speed, success, and strength), a positive influence of turf-derived sediment on recruits’ growth was less obvious, at least over 10 days. The high morphological plasticity of C. filiformis propagules resulted in possible benefits of faster or stronger attachment of more developed propagules being only temporary, and that recruitment may be regulated in accordance with habitat preference. Finally, a field experiment confirmed the observed positive role of turf and the negative influence of algal canopies in the short-term, however, adverse environmental conditions in the longer-term resulted in the loss of most fragments. In conclusion, this study demonstrated the importance of both positive and negative species interactions for the recruitment success of a native alga, suggesting that a shift from kelp to turf algae can initiate further community change.</jats:p

Reduced performance of native infauna following recruitment to a habitat-forming invasive marine alga

Despite well-documented negative impacts of invasive species on native biota, evidence for the facilitation of native organisms, particularly by habitat-forming invasive species, is increasing. However, most of these studies are conducted at the population or community level, and we know little about the individual fitness consequences of recruitment to habitat-forming invasive species and, consequently, whether recruitment to these habitats is adaptive. We determined the consequences of recruitment to the invasive green alga Caulerpa taxifolia on the native soft-sediment bivalve Anadara trapezia and nearby unvegetated sediment. Initially, we documented the growth and survivorship of A. trapezia following a natural recruitment event, to which recruitment to C. taxifolia was very high. After 12 months, few clams remained in either habitat, and those that remained showed little growth. Experimental manipulations of recruits demonstrated that all performance measures (survivorship, growth and condition) were significantly reduced in C. taxifolia sediments compared to unvegetated sediments. Exploration of potential mechanisms responsible for the reduced performance in C. taxifolia sediments showed that water flow and water column dissolved oxygen (DO) were significantly reduced under the canopy of C. taxifolia and that sediment anoxia was significantly higher and sediment sulphides greater in C. taxifolia sediments. However, phytoplankton abundance (an indicator of food supply) was significantly higher in C. taxifolia sediments than in unvegetated ones. Our results demonstrate that recruitment of native species to habitat-forming invasive species can reduce growth, condition and survivorship and that studies conducted at the community level may lead to erroneous conclusions about the impacts of invaders and should include studies on life-history traits, particularly juveniles. © 2008 Springer-Verlag

Unusual but consistent latitudinal patterns in macroalgal habitats and their invertebrate communities across two countries

Aim: The physical characteristics of biogenic habitats and environmental conditions are important determinants of biodiversity, yet their relative importance can change across spatial scales. We aimed to understand how relationships between the physical characteristics of macroalgal habitats and their invertebrate communities varied across spatial scales and whether general ecological patterns occurred across two countries. Location: Eighteen sites across the temperate east coasts of Australia (over 1,300 km) and New Zealand (over 1,000 km), with the latitudinal gradient in the two countries overlapping by 6.73 decimal degrees. Time period: January to early April 2012. Major taxa studied: Three intertidal macroalgal habitats in each country and the invertebrate communities within them. Methods: We measured variation in patch- and individual-level characteristics of macroalgal habitats and their invertebrate communities. Patterns in macroalgal characteristics and communities were compared across latitude, and at smaller spatial scales, and correlated with 26 abiotic environmental variables using multiple multivariate analyses. Results: Separately, macroalgal habitat characteristics and communities showed unusual but consistent nonlinear latitudinal patterns, with greater similarity among sites at the edges of the sampled distribution (i.e., north and south) than at centrally located sites. Macroalgal characteristics did not correlate with a particular set of environmental variables; however, communities were structured by sea surface temperature at the country scale and by macroalgal habitat type and biomass within countries. Anthropogenic variables were also important and may have contributed to the unusual nonlinear patterns observed between macroalgal characteristics and communities across latitude. Main conclusions: Our results support other studies showing that large-scale patterns can emerge from systems where there is high local-scale variability. The results show that communities within macroalgal habitats respond to both the physical characteristics of the habitat and external environmental conditions (e.g., temperature), suggesting that local-scale environmental factors, including anthropogenic stressors, may modulate environmental gradients over larger scales