Facilitation of an invader by a native habitat-former increases along interacting gradients of environmental stress.

Native habitat-forming species can facilitate invasion by reducing environmental stress or consumer pressure. However, the intensity of one stressor along a local gradient may differ when expanding the scale of observation to encompass major variations in background environmental conditions. In this study, we determined how facilitation of the invasive porcelain crab, Petrolisthes elongatus, by the native tube-forming serpulid, Galeolaria caespitosa, varied with environmental gradients at local (tidal height) and larger (wave exposure) spatial scales. G. caespitosa constructs a complex calcareous matrix on the underside of intertidal boulders and we predicted that its positive effects on P. elongatus density would increase in intensity with shore height and be stronger at wave-sheltered than wave-exposed locations. To test these predictions, we conducted two experiments. First, we determined the effects of serpulid presence (boulders with live or dead serpulid matrix vs. bare boulders) at six shore heights that covered the intertidal distribution of P. elongatus. Second, we determined the effects of serpulid presence (present vs. absent), shore height (high vs. low) and wave exposure (sheltered vs. exposed) on crabs across six locations within the invaded range in northern Tasmania, Australia. In Experiment 1, the presence of serpulids (either dead or alive) enhanced P. elongatus densities at all shore heights, with facilitation intensity (as determined by a relative interaction index; RII) tending to increase with shore height. In Experiment 2, serpulids facilitated P. elongatus across shore heights and wave exposures, although crab densities were lower at high shore levels of wave-sheltered locations. However, the intensity of crab facilitation by serpulids was greater on wave-sheltered than on wave-exposed shores, but only at the high shore level. This study demonstrates that local effects of native habitat-formers on invasive species are dependent on prevailing environmental conditions at larger spatial scales and that, under more stressful conditions, invaders become increasingly reliant on positive interactions with native habitat-formers. Increased strength of local-scale facilitation by native species, dampening broader scale variations in environmental stressors, could enhance the ability of invasive species to establish self-sustaining populations in the invaded range.

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 vs. 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.

Carry over effects of nutrient addition on the recovery of an invasive seaweed from the winter die-back.

Nutrient enrichment of coastal waters can enhance the invasibility and regrowth of non-native species. The invasive alga Caulerpa cylindracea has two distinct phases: a well-studied fast-growing summer phase, and a winter latent phase. To investigate the effects of nutrient enrichment on the regrowth of the seaweed after the winter resting-phase, a manipulative experiment was carried out in intertidal rockpools in the North-western Mediterranean. Nutrients were supplied under different temporal regimes: press (constant release from January to May), winter pulse (January to March) and spring pulse (March to May). Independently from the temporal characteristics of their addition, nutrients accelerated the re-growth of C. cylindracea after the winter die-back, resulting in increased percentage covers at the peak of the growing season. Nutrient addition did not influence the number and length of fronds and the biomass. Native components of the algal community did not respond to nutrient additions. Our results show that nutrient supply can favour the spread of C. cylindracea even when occurring at a time of the year at which the seaweed is not actively growing.