Propagules are not all equal: traits of vegetative fragments and disturbance regulate invasion success

Invasion success is regulated by multiple factors. While the roles of disturbance and propagule pressure in regulating the establishment of non-native species are widely acknowledged, that of propagule morphology (a proxy for quality) is poorly known. By means of a multi-factorial field experiment, we tested how the number (5 versus 10) and quality (intact, without fronds or without rhizoids) of fragments of the clonal invasive seaweed, Caulerpa cylindracea, influenced its ability to establish in patches of the native seagrass, Posidonia oceanica, exposed to different intensities of disturbance (0, 50 or 100% reduction in canopy cover). We hypothesized that the ability of fragments to establish would be greater for intact fragments (high quality) and reduced more by frond removal (low quality) than rhizoid removal (intermediate quality). At low propagule pressure or quality, fragment establishment was predicted to increase with increasing disturbance, whereas, at high propagule pressure or quality, it was predicted to be high regardless of disturbance intensity. Disturbance intensity, fragment number and quality had independent effects on C. cylindracea establishment success. Disturbance always facilitated fragment establishment. However, fragments retaining fronds, either intact or deprived of rhizoids, had higher establishment success than fragments deprived of fronds. Increasing propagule number had weak effects on the cover of C. cylindracea. Our results demonstrate that propagule traits enabling the acquisition of resources made available by disturbance can be more important than propagule number in determining the establishment and spread of clonal non-native plants. More generally, our study suggests that propagule quality is a key, yet underexplored, determinant of invasion success

Chapter 8 Comparative Biogeography of Marine Invaders Across Their Native and Introduced Ranges

Biological invasions continue to exert extensive environmental and economic impacts. Understanding why some introduced species become invasive is critical to their management. Determining the mechanisms underpinning invasion success has focussed on aspects of the ecology and physiology of the species in the introduced range. Through the application of biogeographic approaches, however, a growing body of research highlights insights that stem from studying invasion success as a biogeographic issue. In particular, a comparison of both biogeographic regions (i.e., the native and invasive ranges) allows exclusive insight into seven different major biogeographic hypotheses that we identified to explain invader success. These include the enemy release hypothesis, niche shifts, trait differences, the evolution of invasiveness, native allies, environmental matching, and genetic diversity. All imply a difference or gradient between the ranges that may mechanistically explain an invader’s differential performance. This review summarises the support for these seven different theories underpinning the biogeography of marine invasions, and also provides case studies for different theories addressing the comparative biogeography of marine invasions. Additionally, we catalogue the geographic regions of the invasive species used in biogeographic comparisons and the diversity of species, habitats and climate zones examined. Finally, we highlight critical knowledge gaps and suggest future research directions for improving our understanding the processes driving invasion success

Variable direct and indirect effects of a habitat-modifying invasive species on mortality of native fauna

Habitat-modifying invasive species can influence rates of predation on native prey either directly by providing protective structure or indirectly by modifying traits of prey species responding to the habitat. The alga Caulerpa taxifolia is one of the most successful invasive species of shallow-water marine systems globally, often provisioning habitat in areas previously lacking in vegetated structure. We experimentally evaluated the direct effect of Caulerpa to provide refuge for the native clam Anadara trapezia and how this balances with its influence on two trait-mediated indirect interactions that may increase Anadara\u27s susceptibility to predators. Specifically, Caulerpa\u27s alteration of physical and chemical properties of the surrounding water and sediment deteriorate Anadara\u27s condition and predator resistance properties and also cause Anadara, though normally buried, to project from beneath the sediment, exposing it to predators. Our results show that Anadara are somewhat (but not consistently) protected from predators by living among Caulerpa. Shallow burial depth did not counteract this protective effect. However at times of year when predator activity diminishes and conducive environmental conditions develop, negative effects of Caulerpa habitat such as hypoxia and lowered flow may dominate. Under such situations, poor clam condition accentuates Anadara\u27s susceptibility to mortality. Ultimately, a slight and inconsistent positive effect of Caulerpa to protect Anadara from predators is exceeded by the strong negative effect of Caulerpa on clam mortality, which is heightened by clams\u27 weakened condition produced by chronic exposure to Caulerpa. Our results show that invasive habitat-modifying species can affect mortality of native species not simply through obvious positive direct effects of their protective structure, but indirectly through contrasting negative modification of the traits of prey species responding to the habitat

Invasive ecosystem engineer selects for different phenotypes of an associated native species

Invasive habitat-forming ecosystem engineers modify the abiotic environment and thus represent a major perturbation to many ecosystems. Because native species often persist in these invaded habitats but have no shared history with the ecosystem engineer, the engineer may impose novel selective pressure on native species. In this study, we used a phenotypic selection framework to determine whether an invasive habitat-forming ecosystem engineer (the seaweed Caulerpa taxifolia) selects for different phenotypes of a common cooccurring native species (the bivalve Anadara trapezia). Compared to unvegetated habitat, Caulerpa habitat has lower water flow, lower dissolved oxygen, and sediments are more silty and anoxic. We determined the performance consequences of variation in key functional traits that may be affected by these abiotic changes (shell morphology, gill mass, and palp mass) for Anadara transplanted into Caulerpa and unvegetated habitat. Both linear and nonlinear performance gradients in Anadara differed between habitats, and these gradients were stronger in Caulerpa compared to unvegetated sediment. Moreover, in Caulerpa alternate phenotypes performed well, and these phenotypes were different from the dominant phenotype in unvegetated sediment. By demonstrating that phenotype-performance gradients differ between habitats, we have highlighted a role for Caulerpa as an agent of selection on native species

Chapter 8 Comparative Biogeography of Marine Invaders Across Their Native and Introduced Ranges

Biological invasions continue to exert extensive environmental and economic impacts. Understanding why some introduced species become invasive is critical to their management. Determining the mechanisms underpinning invasion success has focussed on aspects of the ecology and physiology of the species in the introduced range. Through the application of biogeographic approaches, however, a growing body of research highlights insights that stem from studying invasion success as a biogeographic issue. In particular, a comparison of both biogeographic regions (i.e., the native and invasive ranges) allows exclusive insight into seven different major biogeographic hypotheses that we identified to explain invader success. These include the enemy release hypothesis, niche shifts, trait differences, the evolution of invasiveness, native allies, environmental matching, and genetic diversity. All imply a difference or gradient between the ranges that may mechanistically explain an invader’s differential performance. This review summarises the support for these seven different theories underpinning the biogeography of marine invasions, and also provides case studies for different theories addressing the comparative biogeography of marine invasions. Additionally, we catalogue the geographic regions of the invasive species used in biogeographic comparisons and the diversity of species, habitats and climate zones examined. Finally, we highlight critical knowledge gaps and suggest future research directions for improving our understanding the processes driving invasion success

Propagule composition regulates the success of an invasive seaweed across a heterogeneous seascape

Abstract Propagule pressure is acknowledged as a key determinant of invasion success. Nonetheless, the role of morphological or physiological attributes of propagules (i.e. their quality) in regulating invader establishment has been little explored. In particular, no study has investigated how the presence of propagules differing in quality within an inoculum influences establishment across heterogeneous landscapes. We experimentally tested the hypothesis that the quality (+Fronds+Rhizoids; +Fronds−Rhizoids; −Fronds+Rhizoids) and the diversity (1, 2 and 3 fragment types) of vegetative fragments of the seaweed Caulerpa taxifolia determine their establishment success across seascapes consisting of bare sediments and patches of the seagrass Zostera muelleri exposed to different disturbance intensities (control, seagrass canopy clipping and total removal). After 6 weeks, seaweed biomass, stolon and frond length, frond and rhizoid number were generally greater in unvegetated habitats (bare sediments and total seagrass removal) than full or reduced seagrass canopies. The type and the diversity of types of fragments inoculated had significant effects on the final biomass and morphological features of C. taxifolia only in vegetated habitats. In control plots, inocula of fragments retaining both fronds and rhizoids achieved higher biomass, developed longer stolons and more fronds. In canopy clipping plots, mixed inocula of +Fronds+Rhizoids and −Fronds+Rhizoids fragments had the greatest biomass and stolon length. Synthesis. Assessing how propagules differing in quality perform in different habitats might be not sufficient to draw a comprehensive picture of invasion risk, as their establishment can be modulated by both negative and positive interactions among them. Propagule composition should be, therefore, considered as a further dimension of propagule pressure. Our results also suggest that the relevance of specific propagule traits for invader establishment decreases from intact to degraded habitats. Considering propagule size in terms of amount of competent propagules, rather than an absolute measure, would refine our ability of predicting invasion risk across habitats differing in biotic or abiotic conditions

Differences in soft-sediment macrobenthic assemblages invaded by Caulerpa taxifolia compared to uninvaded habitats

Caulerpa taxifolia is a habitat-forming green alga that has invaded several temperate regions worldwide. Although C. taxifolia covers large areas of soft-sediment habitat, little is known about its effects on soft-sediment invertebrate assemblages. We compared soft-sediment macroinvertebrate assemblages in 2 estuaries in southeastern Australia invaded by C. taxifolia to examine 2 main predictions: (1) areas covered with C. taxifolia will have different assemblages compared to unvegetated sediment because infauna are inhibited but epifauna are facilitated, and (2) areas with C. taxifolia will have different assemblages compared to those with native seagrasses (Halophila ovalis and Zostera capricorni) because infauna are inhibited but epifauna are not. Multidimensional scaling and ANOSIM showed differences in invertebrate assemblages between all habitats. In C. taxifolia, infauna were less abundant and epifauna were more abundant compared to unvegetated sediment. However, when compared to native seagrasses, epifauna in C. taxifolia were more abundant than in H. ovalis in one estuary but less abundant than in Z. capricorni in another estuary, while infauna in C. taxifolia were less abundant than in both seagrass species. The consistently low infaunal abundance in C. taxifolia, irrespective of infaunal feeding mode, suggests C. taxifolia impacts infauna generally. Examination of environmental factors potentially responsible for the low abundance of infauna indicated that differences in redox potential (and associated chemical changes) may explain patterns in abundance of infauna among habitats. Our findings indicate that invasion by C. taxifolia causes important changes to soft-sediment macroinvertebrate assemblages and suggest that infauna may be particularly vulnerable to invasion because of changes to sediment chemistry