Oxidative Damage and Antioxidants as Markers for the Selection of Emersion Hardening Treatments in GreenshellTM Mussel Juveniles (Perna canaliculus).

Transport out of the water is one of the most challenging events for juvenile Perna canaliculus and can be a highly inefficient process, with many juveniles subsequently being lost following extended periods of emersion. Hardening techniques offer a possible method for reducing transport-related stress. In this study, different hardening treatments (short, long and intermittent sub-lethal emersion) were used to prepare ~1.2 mm P.canaliculus for transport (20 h) and subsequent reoxygenation stress during re-immersion (i.e., recovery). The oxidative stress responses, resettlement behaviour, respiration rates and survival of the mussels after transport and during recovery were all assessed. Short emersion (1 h) as a hardening treatment prior to transport did not cause major stress to the mussels, which maintained respiration at control levels, showed significantly stimulated antioxidant defences during recovery, showed greater resettlement behaviour and remained viable after 24 h of recovery. In comparison, the long and intermittent emersion treatments negatively impacted oxidative stress responses and affected the viability of the mussels after 24 h of recovery. This study showed that exposing juvenile P.canaliculus to a mild stress prior to transport may stimulate protective mechanisms, therefore eliciting a hardening response, but care must be taken to avoid overstressing the mussels. Improving the management of stress during the transport of juvenile mussels may be key to minimising mussel losses and increasing harvest production, and biomarkers associated with oxidative stress/antioxidant metabolism could be valuable tools to ensure emersion hardening does not overstress the mussels and reduce survival.

Heterogeneity within and among co-occurring foundation species increases biodiversity.

Habitat heterogeneity is considered a primary causal driver underpinning patterns of diversity, yet the universal role of heterogeneity in structuring biodiversity is unclear due to a lack of coordinated experiments testing its effects across geographic scales and habitat types. Furthermore, key species interactions that can enhance heterogeneity, such as facilitation cascades of foundation species, have been largely overlooked in general biodiversity models. Here, we performed 22 geographically distributed experiments in different ecosystems and biogeographical regions to assess the extent to which variation in biodiversity is explained by three axes of habitat heterogeneity: the amount of habitat, its morphological complexity, and capacity to provide ecological resources (e.g. food) within and between co-occurring foundation species. We show that positive and additive effects across the three axes of heterogeneity are common, providing a compelling mechanistic insight into the universal importance of habitat heterogeneity in promoting biodiversity via cascades of facilitative interactions. Because many aspects of habitat heterogeneity can be controlled through restoration and management interventions, our findings are directly relevant to biodiversity conservation.

Emersion and Relative Humidity Modulate Stress Response and Recovery Dynamics in Juvenile Mussels (Perna canaliculus).

The early stages of intertidal mussels, including the green-lipped mussel, Perna canaliculus, face both direct and indirect environmental threats. Stressors may influence physiological status and, ultimately, survival. An understanding of the nature of stress experienced is critical to inform conservation and aquaculture efforts. Here, we investigated oxidative stress dynamics in juvenile P. canaliculus in relation to emersion duration (1-20 h) and relative humidity (RH, 29-98%) by quantifying oxidative damage (protein carbonyls, lipid hydroperoxides, 8-hydroxydeoxyguanosine) and enzymatic antioxidants (superoxide dismutase, catalase, glutathione peroxidase and reductase). Mussels held in low RH during emersion experienced severe water loss (>70%), high mortality (>80%) and increased oxidative damage (35-45% increase compared to control conditions), while mussels held at high RH were not impacted, even after 20 h of air exposure. Following re-immersion, reoxygenation stress resulted in further increases in damage markers in mussels that had experienced dryer emersion conditions; protective action of antioxidants increased steadily during the 10 h re-immersion period, apparently supporting a reduction in damage markers after 1-5 h of immersion. Clearly, conditions during emersion, as well as duration, substantially influence physiological performance and recovery of juvenile mussels. Successful recruitment to intertidal beds or survival in commercial aquaculture operations may be mediated by the nature of emersion stress experienced by these vulnerable juveniles.

The role of biofouling development in the loss of seed mussels in aquaculture.

The efficient seeding of juvenile mussels is critical to the sustainability and profitability of mussel aquaculture. However, seeding mussels is highly inefficient, with many juveniles being lost in the following few months. One possible cause of these losses could be the development of biofouling assemblages. Therefore, the relationships between biofouling accumulation and losses of juveniles were assessed. Losses of juvenile mussels were initially high (42.9-49.1% over approximately one to two weeks), with lower rates of loss over the following four to five months. Biofouling development followed a successional pattern beginning with colonisation by amphipods, subsequent establishment of macroalgae, and the formation of an assemblage dominated by mussels and sessile invertebrates. However, biofouling development did not play a major role in the loss of juveniles. Rather, large-scale losses of mussels occurred shortly after seeding when biofouling was scant, suggesting alternative causes of loss were in operation.

Secondary foundation species enhance biodiversity.

It has long been recognized that primary foundation species (FS), such as trees and seagrasses, enhance biodiversity. Among the species facilitated are secondary FS, including mistletoes and epiphytes. Case studies have demonstrated that secondary FS can further modify habitat-associated organisms ('inhabitants'), but their net effects remain unknown. Here we assess how inhabitants, globally, are affected by secondary FS. We extracted and calculated 2,187 abundance and 397 richness Hedges' g effect sizes from 91 and 50 publications, respectively. A weighted meta-analysis revealed that secondary FS significantly enhanced the abundance and richness of inhabitants compared to the primary FS alone. This indirect facilitation arising through sequential habitat formation was consistent across environmental and experimental conditions. Complementary unweighted analyses on log response ratios revealed that the magnitude of these effects was similar to the global average strength of direct facilitation from primary foundation species and greater than the average strength of trophic cascades, a widely recognized type of indirect facilitation arising through sequential consumption. The finding that secondary FS enhance the abundance and richness of inhabitants has important implications for understanding the mechanisms that regulate biodiversity. Integrating secondary FS into conservation practice will improve our ability to protect biodiversity and ecosystem function.

A review of three decades of research on the invasive kelp Undaria pinnatifida in Australasia: An assessment of its success, impacts and status as one of the world's worst invaders.

Marine invasive macroalgae can have severe local-scale impacts on ecological communities. The kelp Undaria pinnatifida is one of the most successful marine invasive species worldwide, and is widely regarded as one of the worst. Here, we review research on Undaria in Australasia, where the kelp is established throughout much of New Zealand and south-eastern Australia. The presence of Undaria for at least three decades in these locations makes Australasia one of the longest-invaded bioregions globally, and a valuable case study for considering Undaria's invasion success and associated impacts. In Australasia, Undaria has primarily invaded open spaces, turf communities, and gaps in native canopies within a relatively narrow elevation band on rocky shores. Despite its high biomass, Undaria has relatively few direct impacts on native species, and can increase community-wide attributes such as primary productivity and the provision of biogenic habitat. Therefore, Australasian Undaria research provides an example of a decoupling between the success and impact of an invasive species. Undaria will most likely continue to spread along thousands of kilometres of rocky coastline in temperate Australasia, due to its tolerance to large variations in temperature, ability to exploit disturbances to local communities, and the continued transfer among regions via vessel movements and aquaculture activities. However, the spread of Undaria remains difficult to manage as eradication is challenging and seldom successful. Therefore, understanding potential invasion pathways, maintaining native canopy-forming species that limit Undaria success, and effectively managing anthropogenic vectors of Undaria spread, should be key management priorities.

A sixth-level habitat cascade increases biodiversity in an intertidal estuary.

Many studies have documented habitat cascades where two co-occurring habitat-forming species control biodiversity. However, more than two habitat-formers could theoretically co-occur. We here documented a sixth-level habitat cascade from the Avon-Heathcote Estuary, New Zealand, by correlating counts of attached inhabitants to the size and accumulated biomass of their biogenic hosts. These data revealed predictable sequences of habitat-formation (=attachment space). First, the bivalve Austrovenus provided habitat for green seaweeds (Ulva) that provided habitat for trochid snails in a typical estuarine habitat cascade. However, the trochids also provided habitat for the nonnative bryozoan Conopeum that provided habitat for the red seaweed Gigartina that provided habitat for more trochids, thereby resetting the sequence of the habitat cascade, theoretically in perpetuity. Austrovenus is here the basal habitat-former that controls this "long" cascade. The strength of facilitation increased with seaweed frond size, accumulated seaweed biomass, accumulated shell biomass but less with shell size. We also found that Ulva attached to all habitat-formers, trochids attached to Ulva and Gigartina, and Conopeum and Gigartina predominately attached to trochids. These "affinities" for different habitat-forming species probably reflect species-specific traits of juveniles and adults. Finally, manipulative experiments confirmed that the amount of seaweed and trochids was important and consistent regulators of the habitat cascade in different estuarine environments. We also interpreted this cascade as a habitat-formation network that describes the likelihood of an inhabitant being found attached to a specific habitat-former. We conclude that the strength of the cascade increased with the amount of higher-order habitat-formers, with differences in form and function between higher and lower-order habitat-formers, and with the affinity of inhabitants for higher-order habitat-formers. We suggest that long habitat cascades are common where species traits allow for physical attachment to other species, such as in marine benthic systems and old forest.