Graeme F. Clark

The Role of Propagule Pressure in Invasion Success

One of the core goals of invasion biology is the identification of factors that increase the risk of establishment success of non-native species. Historically, marine invasions have been investigated through observational studies and surveys (Cohen and Carlton 1998; Ruiz et al. 2000). These have guided ecologists towards the processes most relevant to invasion, but researchers are becoming increasingly aware of the limitations of observational studies alone. It is clear that different factors may influence invasion success at different stages of the invasion process (Kolar and Lodge 2001) and a major challenge is to quantify the relative importance of these factors. Understanding the intricacies of invasion dynamics requires a rigorous approach, in which potentially important factors can be controlled, manipulated and tested (Ruiz et al. 2000). Particularly strong calls have been made for the inclusion of propagule pressure or invader supply into our models, experiments and surveys, and this chapter reviews recent progress in elucidating the role of propagule pressure on invasion success in marine ecosystems.

Polychaete Richness and Abundance Enhanced in Anthropogenically Modified Estuaries Despite High Concentrations of Toxic Contaminants

Ecological communities are increasingly exposed to multiple chemical and physical stressors, but distinguishing anthropogenic impacts from other environmental drivers remains challenging. Rarely are multiple stressors investigated in replicated studies over large spatial scales (>1000 kms) or supported with manipulations that are necessary to interpret ecological patterns. We measured the composition of sediment infaunal communities in relation to anthropogenic and natural stressors at multiple sites within seven estuaries. We observed increases in the richness and abundance of polychaete worms in heavily modified estuaries with severe metal contamination, but no changes in the diversity or abundance of other taxa. Estuaries in which toxic contaminants were elevated also showed evidence of organic enrichment. We hypothesised that the observed response of polychaetes was not a ‘positive’ response to toxic contamination or a reduction in biotic competition, but due to high levels of nutrients in heavily modified estuaries driving productivity in the water column and enriching the sediment over large spatial scales. We deployed defaunated field-collected sediments from the surveyed estuaries in a small scale experiment, but observed no effects of sediment characteristics (toxic or enriching). Furthermore, invertebrate recruitment instead reflected the low diversity and abundance observed during field surveys of this relatively ‘pristine’ estuary. This suggests that differences observed in the survey are not a direct consequence of sediment characteristics (even severe metal contamination) but are related to parameters that covary with estuary modification such as enhanced productivity from nutrient inputs and the diversity of the local species pool. This has implications for the interpretation of diversity measures in large-scale monitoring studies in which the observed patterns may be strongly influenced by many factors that covary with anthropogenic modification.

Invasion Expansion: Time since introduction best predicts global ranges of marine invaders.

Strategies for managing biological invasions are often based on the premise that characteristics of invading species and the invaded environment are key predictors of the invader’s distribution. Yet, for either biological traits or environmental characteristics to explain distribution, adequate time must have elapsed for species to spread to all potential habitats. We compiled and analyzed a database of natural history and ecological traits of 138 coastal marine invertebrate species, the environmental conditions at sites to which they have been introduced, and their date of first introduction. We found that time since introduction explained the largest fraction (20%) of the variability in non-native range size, while traits of the species and environmental variables had significant, but minimal, influence on non-native range size. The positive relationship between time since introduction and range size indicates that non-native marine invertebrate species are not at equilibrium and are still spreading, posing a major challenge for management of coastal ecosystems.

Elevated nutrients change bacterial community composition and connectivity: high throughput sequencing of young marine biofilms

Abstract Biofilms are integral to many marine processes but their formation and function may be affected by anthropogenic inputs that alter environmental conditions, including fertilisers that increase nutrients. Density composition and connectivity of biofilms developed in situ (under ambient and elevated nutrients) were compared using 454-pyrosequencing of the 16S gene. Elevated nutrients shifted community composition from bacteria involved in higher processes (eg Pseudoalteromonas spp. invertebrate recruitment) towards more nutrient-tolerant bacterial species (eg Terendinibacter sp.). This may enable the persistence of biofilm communities by increasing resistance to nutrient inputs. A core biofilm microbiome was identified (predominantly Alteromonadales and Oceanospirillales) and revealed shifts in abundances of core microbes that could indicate enrichment by fertilisers. Fertiliser decreased density and connectivity within biofilms indicating that associations were disrupted perhaps via changes to energetic allocations within the core microbiome. Density composition and connectivity changes suggest nutrients can affect the stability and function of these important marine communities.

Assessing national biodiversity trends for rocky and coral reefs through the integration of citizen science and scientific monitoring programs.

Abstract Reporting progress against targets for international biodiversity agreements is hindered by a shortage of suitable biodiversity data. We describe a cost-effective system involving Reef Life Survey citizen scientists in the systematic collection of quantitative data covering multiple phyla that can underpin numerous marine biodiversity indicators at high spatial and temporal resolution. We then summarize the findings of a continental- and decadal-scale State of the Environment assessment for rocky and coral reefs based on indicators of ecosystem state relating to fishing, ocean warming, and invasive species and describing the distribution of threatened species. Fishing impacts are widespread, whereas substantial warming-related change affected some regions between 2005 and 2015. Invasive species are concentrated near harbors in southeastern Australia, and the threatened-species index is highest for the Great Australian Bight and Tasman Sea. Our approach can be applied globally to improve reporting against biodiversity targets and enhance public and policymakers’ understanding of marine biodiversity trends.

Coastal urban lighting has ecological consequences for multiple trophic levels under the sea

Urban land and seascapes are increasingly exposed to artificial lighting at night (ALAN), which is a significant source of light pollution. A broad range of ecological effects are associated with ALAN, but the changes to ecological processes remain largely unstudied. Predation is a key ecological process that structures assemblages and responds to natural cycles of light and dark. We investigated the effect of ALAN on fish predatory behaviour, and sessile invertebrate prey assemblages. Over 21days fish and sessile assemblages were exposed to 3 light treatments (Day, Night and ALAN). An array of LED spotlights was installed under a wharf to create the ALAN treatments. We used GoPro cameras to film during the day and ALAN treatments, and a Dual frequency IDentification SONar (DIDSON) to film during the night treatments. Fish were most abundant during unlit nights, but were also relatively sedentary. Predatory behaviour was greatest during the day and under ALAN than at night, suggesting that fish are using structures for non-feeding purposes (e.g. shelter) at night, but artificial light dramatically increases their predatory behaviour. Altered predator behaviour corresponded with structural changes to sessile prey assemblages among the experimental lighting treatments. We demonstrate the direct effects of artificial lighting on fish behaviour and the concomitant indirect effects on sessile assemblage structure. Current and future projected use of artificial lights has the potential to significantly affect predator-prey interactions in marine systems by altering habitat use for both predators and prey. However, developments in lighting technology are a promising avenue for mitigation. This is among the first empirical evidence from the marine system on how ALAN can directly alter predation, a fundamental ecosystem process, and have indirect trophic consequences.

What does impacted look like? High diversity and abundance of epibiota in modified estuaries

Ecosystems modified by human activities are generally predicted to be biologically impoverished. However, much pollution impact theory stems from laboratory or small-scale field studies, and few studies replicate at the level of estuary. Furthermore, assessments are often based on sediment contamination and infauna, and impacts to epibiota (sessile invertebrates and algae) are seldom considered. We surveyed epibiota in six estuaries in south-east Australia. Half the estuaries were relatively pristine, and half were subject to internationally high levels of contamination, urbanisation, and industrialisation. Contrary to predictions, epibiota in modified estuaries had greater coverage and were similarly diverse as those in unmodified estuaries. Change in epibiota community structure was linearly correlated with sediment-bound copper, and the tubeworm Hydroides elegans showed a strong positive correlation with sediment metals. Stressors such as metal contamination can reduce biodiversity and productivity, but others such as nutrient enrichment and resource provision may obscure signals of impact.

The Role of Habitat Complexity in Community Development Is Mediated by Resource Availability

Habitat complexity strongly affects the structure and dynamics of ecological communities, with increased complexity often leading to greater species diversity and abundance. However, habitat complexity changes as communities develop, and some species alter their environment to themselves provide habitat for other species. Most experimental studies manipulate basal substrate complexity, and while the importance of complexity likely changes during community development, few studies have examined the temporal dynamics of this variable. We used two experiments to quantify the importance of basal substrate complexity to sessile marine invertebrate community development through space and time. First, we compared effects of substrate complexity at 70 sites across ten estuaries. Sites differed in recruitment and community development rates, and after three months provided spatial variation in community development stage. Second, we tested for effects of substrate complexity at multiple times at a single site. In both experiments, complexity affected marine sessile invertebrate community composition in the early stages of community development when resource availability was high. Effects of complexity diminished through time as the amount of available space (the primary limiting resource) declined. Our work suggests the presence of a bare-space threshold, at which structural complexity of the basal substrate is overwhelmed by secondary biotic complexity. This threshold will be met at different times depending on local recruitment and growth rates and is likely to vary with productivity gradients.

Contaminant cocktails: Interactive effects of fertiliser and copper paint on marine invertebrate recruitment and mortality.

Understanding interactive effects of contaminants is critical to predict how human activities change ecosystem structure and function. We examined independent and interactive effects of two contaminants (fertiliser and copper paint) on the recruitment, mortality, and total abundance of developing invertebrate communities in the field, 2, 4, 6, and 8 weeks after substrate submersion. Contaminants affected community structure differently, and produced an intermediate community in combination. Fertiliser increased recruitment and decreased mortality of active filter feeders (ascidians and barnacles), while copper paint decreased recruitment and increased mortality of some taxa. Contaminants applied together affected some taxa (e.g. Didemnid ascidians) antagonistically, as fertiliser mitigated adverse effects of copper paint. Recruitment of active filter feeders appears to be indicative of nutrient enrichment, and their increased abundance may reduce elevated nutrients in modified waterways. This study demonstrates the need to consider both independent and interactive effects of contaminants on marine communities in the field.

Responses of ghost crabs to habitat modification of urban sandy beaches

Sandy beaches in highly urbanised areas are subject to a wide range of human impacts. Ghost crabs are a commonly used ecological indicator on sandy beaches, as they are key consumers in these systems and counting burrow openings allows for rapid assessment of population size. This study assessed the pressures of urbanisation on sandy beaches in the highly urbanised estuary of Sydney Harbour. Across 38 beaches, we examined which physical beach properties, management practices and human induced habitat modification best predicted ghost crab distributions. Of all variables measured, the frequency of mechanical beach cleaning was the most important predictor of crab abundance, with low burrow densities at the highest cleaning frequency and the highest densities at beaches cleaned at the intermediate frequency (≤3 times per week). These results indicate that ghost crab populations in Sydney Harbour are more robust to the impacts of urbanisation than previously thought.