Invasive Tunicate Research Articles

Proteome-wide 4-hydroxy-2-nonenal signature of oxidative stress in the marine invasive tunicate Botryllus schlosseri.

Oxidative stress results from environmental challenges that increase in frequency and severity during the Anthropocene. Oxygen radical attack causes lipid peroxidation leading to HNE production. Proteome-wide HNE adduction is highly prevalent in Botryllus schlosseri , a widely distributed, highly invasive, and economically important biofouling ascidian and the first marine species to be analyzed for proteome HNE modification. HNE adduction of specific proteins physiologically sequesters reactive oxygen species, which enhances fitness and resilience during environmental change.

Geography and developmental plasticity shape post-larval thermal tolerance in the golden star tunicate, Botryllus schlosseri

Local adaptation and phenotypic plasticity play key roles in mediating organisms’ ability to respond to spatiotemporal variation in temperature. These two processes often act together to generate latitudinal or elevational clines in acute temperature tolerance. Phenotypic plasticity is also subject to local adaptation, with the expectation that populations inhabiting more variable environments should exhibit greater phenotypic plasticity of thermal tolerance. Here we examine the potential for local adaptation and developmental plasticity of thermal tolerance in the widespread invasive tunicate Botryllus schlosseri. By comparing five populations across a thermal gradient spanning 4.4° of latitude in the northwest Atlantic, we demonstrate that warmer populations south of the Gulf of Maine exhibit significantly increased (∼0.2 °C) post-larval temperature tolerance relative to the colder populations within it. We also show that B. schlosseri post-larvae possess a high degree of developmental plasticity for this trait, shifting their median temperature of survival (LT50) upwards by as much as 0.18 °C per 1 °C increase in environmental temperature. Lastly, we found that populations vary in their degrees of developmental plasticity, with populations that experience more pronounced short-term temperature variability exhibiting greater developmental plasticity, suggesting the local adaptation of developmental plasticity. By comparing the thermal tolerance of populations across space and through time, we demonstrate how geography and developmental plasticity have shaped thermal tolerance in B. schlosseri. These results help inform our understanding of how species are able to adjust their thermal physiology in new environments, including those encountered during invasion and under increasingly novel climate conditions.

Comparison of the Efficiency of Different Eradication Treatments to Minimize the Impacts Caused by the Invasive Tunicate Styela plicata in Mussel Aquaculture.

In 2017, aquaculture producers of the Albufeira lagoon, Portugal, reported an invasion of tunicates that was disrupting mussel production, particularly the tunicate Styela plicata (Lesueur, 1823). A totally effective eradication method still does not exist, particularly for S. plicata, and the effects of the eradication treatments on bivalves' performance are also poorly understood. Our study examined the effectiveness of eradication treatments using three laboratory trials and five treatments (air exposure, freshwater immersion, sodium hypochlorite, hypersaline solution and acetic acid) for S. plicata, as well as their effects on survival and growth of blue mussel Mytilus edulis Linnaeus, 1758. While air exposure and freshwater immersion caused a 27% mortality rate in S. plicata, the acetic acid treatment was the most effective in eliminating this species (>90% mortality). However, a 33-40% mortality rate was registered in mussels. Both species were not affected by the hypersaline treatment in the last trial, but the sodium hypochlorite treatment led to a 57% mortality rate in mussels. Differences in mussels' growth rates were not detected. These trials represent a step forward in responding to the needs of aquaculture producers. However, further studies are needed to investigate the susceptibility of tunicates to treatments according to sexual maturation, as well as to ensure minimum mussel mortality in the most effective treatments, and to better understand the effects on mussel physiological performance in the long-term.

Susceptibility of invasive tunicates Clavelina oblonga to reduced seawater salinities

The intensive colonial tunicate Clavelina oblonga invasion was observed in the summer of 2020 at shellfish farms located on the western and eastern coasts of the Istrian peninsula (NE, Adriatic, Croatia). Aquaculture facilities create an ideal environment for ascidians, therefore, as a possible environmentally friendly eradication measure, laboratory experimental exposure of C. oblonga to different reduced seawater salinities of 37, 30, 20 and 11 was examined in November 2020. During the total observation time (14 days exposure + 14 days of recovery at 37) at all seawater salinities, there was no sexual reproduction of Clavelina specimens and larvae settlement in general. Salinities of 37 and 30 didn’t provoke any significant effect in colonial corpus weight change and morphology. Salinities of 20 and 11 caused disorder in specimens feed uptake and feeding itself which led to change of colour, tissue necrosis and finally body fragmentation. Similar separate laboratory exposure of mussel Mytilus galloprovincialis specimens did not provoke significant differences in meat yield between control 37 salinity (14.0 %) and reduced salinities of 30 (14.2 %), 20 (14.6 %) and 11 (15.3 %) measured at the end of the recovery experiment. Based on our experimental results of C. oblonga salinity susceptibility and field observations (2020–2021), we can assume that field translocation of overgrown mussels and infrastructures to locations of seawater < 20 salinities could be a natural way for its eradication.

Complementary genomic and epigenomic adaptation to environmental heterogeneity.

While adaptation is commonly thought to result from selection on DNA sequence-based variation, recent studies have highlighted an analogous epigenetic component as well. However, the relative roles of these mechanisms in facilitating population persistence under environmental heterogeneity remain unclear. To address the underlying genetic and epigenetic mechanisms and their relationship during environmental adaptation, we screened the genomes and epigenomes of nine global populations of a predominately sessile marine invasive tunicate, Botryllus schlosseri. We detected clear population differentiation at the genetic and epigenetic levels. Patterns of genetic and epigenetic structure were significantly influenced by local environmental variables. Among these variables, minimum annual sea surface temperature was identified as the top explanatory variable for both genetic and epigenetic variation. However, patterns of population structure driven by genetic and epigenetic variation were somewhat distinct, suggesting possible autonomy of epigenetic variation. We found both shared and specific genes and biological pathways among genetic and epigenetic loci associated with environmental factors, consistent with complementary and independent contributions of genetic and epigenetic variation to environmental adaptation in this system. Collectively, these mechanisms may facilitate population persistence under environmental change and sustain successful invasions across novel environments.

Environmental DNA Based Surveillance for the Highly Invasive Carpet Sea Squirt Didemnum vexillum: A Targeted Single-Species Approach

The presence and diversity of marine non-native species, the number of new invasions, and the impact on native communities and habitats are important metrics used to assess the health of marine ecosystems. Monitoring for marine non-native species, using traditional approaches such as rapid assessment surveys (RASs), requires taxonomic expertise and may still fail to detect rare or inconspicuous species. This study reports a validation process for a quantitative PCR (qPCR) assay based on the cytochrome oxidase 1 gene, designed to detect highly invasive tunicate Didemnum vexillum by targeting environmental DNA (eDNA) present in water samples. The D. vexillum qPCR assay showed high sensitivity, with the threshold limit of detection (LOD) and modeled LOD3 (based on triplicate qPCR reactions) estimated as 9.187 and 1.117 copies reaction–1, respectively and the limit of quantification (LOQ) was calculated as 18 copies reaction–1. Analyses of water samples collected from selected Pacific oyster farms and recreational marinas in Scotland showed 100% concordance between the historical data on presence of D. vexillum from RASs and detection of D. vexillum eDNA. Consistency of detection of D. vexillum eDNA among different sampling points within each infected sampling site varied, ranging between 100% positive throughout the site to some sampling points testing “negative” or only as “suspected” for D. vexillum. Sites with lower within-site detection consistency included sites with a low density of D. vexillum as reported by RASs or were sites undergoing D. vexillum management. The present pilot monitoring program demonstrates the potential to generate important data on presence of D. vexillum. This program will be scaled up across large geographic regions and used in the first instance to focus and target the traditional RASs to D. vexillum eDNA-positive sites in a cost-effective way, with an aim to verify the species presence by visual observation and direct Sanger sequencing of positive qPCR products.

Genetic diversity and relatedness in aquaculture and marina populations of the invasive tunicate Didemnum vexillum in the British Isles

Introductions of invasive, non-native species in the marine environment are increasing as human activity within coastal areas rises. Genetic datasets are useful tools to identify source populations, track routes of invasions, and illuminate the role of genetic variation in the establishment and subsequent spread of novel introductions. Here, a microsatellite dataset is used to estimate the genetic diversity and population structure of 7 introduced Didemnum vexillum populations in Britain and Ireland, 4 of which are associated with aquaculture and 3 with marinas. Genetic differentiation observed between these populations indicates human-mediated transport as the main mechanism underlying the population structure of D. vexillum in Britain and Ireland. In addition to elucidating patterns of population structure we found that aquaculture sites showed significantly higher genetic diversity (measured as allelic richness) in comparison to the marina sites. We discuss these findings in relation to the history of each invasion, the complex life history of D. vexillum, and available evidence of the relative invasiveness of these populations. Our results show numerous interesting patterns which highlight further research avenues to elucidate the complex factors underlying the global spread of this successful invader.

Marine Natural Products from Tunicates and Their Associated Microbes.

Marine tunicates are identified as a potential source of marine natural products (MNPs), demonstrating a wide range of biological properties, like antimicrobial and anticancer activities. The symbiotic relationship between tunicates and specific microbial groups has revealed the acquisition of microbial compounds by tunicates for defensive purpose. For instance, yellow pigmented compounds, “tambjamines”, produced by the tunicate, Sigillina signifera (Sluiter, 1909), primarily originated from their bacterial symbionts, which are involved in their chemical defense function, indicating the ecological role of symbiotic microbial association with tunicates. This review has garnered comprehensive literature on MNPs produced by tunicates and their symbiotic microbionts. Various sections covered in this review include tunicates’ ecological functions, biological activities, such as antimicrobial, antitumor, and anticancer activities, metabolic origins, utilization of invasive tunicates, and research gaps. Apart from the literature content, 20 different chemical databases were explored to identify tunicates-derived MNPs. In addition, the management and exploitation of tunicate resources in the global oceans are detailed for their ecological and biotechnological implications.

Genome-Wide Identification, Comparison, and Expression Analysis of Transcription Factors in Ascidian Styela clava.

Tunicates include diverse species, as they are model animals for evolutionary developmental biology study. The embryonic development of tunicates is known to be extensively regulated by transcription factors (TFs). Styela clava, the globally distributed invasive tunicate, exhibits a strong capacity for environmental adaptation. However, the TFs were not systematically identified and analyzed. In this study, we reported 553 TFs categorized into 60 families from S. clava, based on the whole genome data. Comparison of TFs analysis among the tunicate species revealed that the gene number in the zinc finger superfamily displayed the most significant discrepancy, indicating this family was under the highly evolutionary selection and might be related to species differentiation and environmental adaptation. The greatest number of TFs was discovered in the Cys2His2-type zinc finger protein (zf-C2H2) family in S. clava. From the point of temporal view, more than half the TFs were expressed at the early embryonic stage. The expression correlation analysis revealed the existence of a transition for TFs expression from early embryogenesis to the later larval development in S. clava. Eight Hox genes were identified to be located on one chromosome, exhibiting different arrangement and expression patterns, compared to Ciona robusta (C. intestinalis type A). In addition, a total of 23 forkhead box (fox) genes were identified in S. clava, and their expression profiles referred to their potential roles in neurodevelopment and sensory organ development. Our data, thus, provides crucial clues to the potential functions of TFs in development and environmental adaptation in the leathery sea squirt.

Testing ecological theories in the Anthropocene: alteration of succession by an invasive marine species

AbstractEcologists employ a diverse body of theory to help explain patterns and processes in ecological systems, with the consistency of ecological theories tested against an increasingly altered world. The global redistribution of species is a prominent impact of climate change and human‐mediated biological invasions and often results in negative impacts to ecosystems. Recently, a species of colonial tunicate not previously present, Clavelina oblonga, has become prominent within the marine fouling community of Beaufort, North Carolina, USA. Fifty years ago, researchers tested theories of ecological succession developed in terrestrial systems and found that this marine community was characterized by a heterogeneous mixture of species that varied inter‐annually (i.e., multiple community states) and increased in diversity over time. A recent follow‐up study found that the fouling community is dominated by C. oblonga with patterns of community development and structure that led to the loss of alternative community states, domination by C. oblonga, and reduced species diversity. The present study addressed the question: Will patterns of community development identified over 50 yr ago still operate under reduced cover of invasive C. oblonga? This study also quantified the impacts of two large‐scale environmental disturbances (extremely cold winter and hurricane) on the abundance of C. oblonga and the resulting fouling community. Clavelina oblonga remains a dominant component of the fouling community due to its rapid growth and strong seasonal recruitment. Under conditions of reduced percent cover of C. oblonga, the local fouling community displayed unique community states that became even more distinct over time, consistent with the pattern of multiple community states identified by researchers over 50 yr ago, and that were dependent upon date of disturbance. Natural disturbances in this study caused by a harsh winter and a hurricane greatly reduced the presence of C. oblonga. This experiment advanced our understanding of marine community ecology by testing whether the concept of multiple community states identified over 50 yr ago is still operable in the absence of invasive C. oblonga, and highlights how natural environment disturbances can potentially moderate the spread of this invasive tunicate.

Towards the scalable isolation of cellulose nanocrystals from tunicates

In order for sustainable nanomaterials such as cellulose nanocrystals (CNCs) to be utilized in industrial applications, a large-scale production capacity for CNCs must exist. Currently the only CNCs available commercially in kilogram scale are obtained from wood pulp (W-CNCs). Scaling the production capacity of W-CNCs isolation has led to their use in broader applications and captured the interest of researchers, industries and governments alike. Another source of CNCs with potential for commercial scale production are tunicates, a species of marine animal. Tunicate derived CNCs (T-CNCs) are a high aspect ratio CNC, which can complement commercially available W-CNCs in the growing global CNC market. Herein we report the isolation and characterization of T-CNCs from the tunicate Styela clava, an invasive species currently causing significant harm to local aquaculture communities. The reported procedure utilizes scalable CNC processing techniques and is based on our experiences from laboratory scale T-CNC isolation and pilot scale W-CNC isolation. To our best knowledge, this study represents the largest scale where T-CNCs have been isolated from any tunicate species, under any reaction conditions. Demonstrating a significant step towards commercial scale isolation of T-CNCs, and offering a potential solution to the numerous challenges which invasive tunicates pose to global aquaculture communities.

The role of invasive tunicates as reservoirs of molluscan pathogens

Ascidian tunicates frequently display rapid expansion when introduced beyond their native range and are considered successful invaders. This invasive potential may be exacerbated by a warming climate, allowing for the occupation of environmental niches previously held by native species. Research into tunicate invasion ecology is prevalent, but less is known about their role in pathogen maintenance. This study investigated the impact of invasive tunicates on the maintenance of pathogens that affect commercial bivalves, including the cultured species Ostrea edulis (European flat oyster) and Crassostrea gigas (Pacific oyster), and the fished species Cerastoderma edule (Common cockle). Focal pathogens included ostreid herpesvirus OsHV-1 μVar, Vibrio aestuarianus, Bonamia ostreae and Minchinia spp. The range of pathogens in their molluscan hosts was determined and the tunicates Botrylloides violaceus, Didemnum vexillum and Styela clava were then screened for these same pathogens, using both field samples from oyster culture sites and marinas and a series of laboratory cohabitation trials. Sample sites reflected areas close to and further away from known pathogen sources. PCR, Sanger sequencing and histology confirmed the presence of B. ostreae and Minchinia mercenariae-like in S. clava, and V. aestuarianus was confirmed by qPCR in B. violaceus and D. vexillum. Furthermore, histology confirmed Minchinia mercenariae-like sporonts in S. clava suggesting that the tunicate can facilitate replication of this species. S. clava also maintained B. ostreae in tanks with no oysters present. The results indicate that tunicates can act as reservoirs of infection in areas where disease occurs and potentially transport diseases to uninfected sites.

Stress Memory of Recurrent Environmental Challenges in Marine Invasive Species: Ciona robusta as a Case Study.

Fluctuating environmental changes impose tremendous stresses on sessile organisms in marine ecosystems, in turn, organisms develop complex response mechanisms to keep adaptive homeostasis for survival. Physiological plasticity is one of the primary lines of defense against environmental challenges, and such defense often relies on the antioxidant defense system (ADS). Hence, it is imperative to understand response mechanisms of ADS to fluctuating environments. Invasive species provide excellent models to study how species cope with environmental stresses, as invasive species encounter sudden, and often recurrent, extensive environmental challenges during the whole invasion process. Here, we studied the roles of ADS on rapid response to recurrent cold challenges in a highly invasive tunicate (Ciona robusta) by simulating cold stresses during its invasion process. We assessed antioxidative indicators, including malondialdehyde (MDA), total antioxidant capacity (T-AOC), superoxide dismutase (SOD), catalase (CAT), and glutathione (GSH), as well as transcriptional changes of ADS-related genes to reveal the physiological plasticity under recurring cold stresses. Our results demonstrated that physiological homeostasis relied on the resilience of ADS, which further accordingly tuned antioxidant activity and gene expression to changing environments. The initial cold stress remodeled baselines of ADS to promote the development of stress memory, and subsequent stress memory largely decreased the physiological response to recurrent environmental challenges. All results here suggest that C. robusta could develop stress memory to maintain physiological homeostasis in changing or harsh environments. The results obtained in this study provide new insights into the mechanism of rapid physiological adaption during biological invasions.

Distribution and abundance of the invasive tunicate Ciona intestinalis (Linnaeus, 1767) in Icelandic harbours

Ciona intestinalis is within the group of taxa that are spreading globally and is one of the most thriving invasive marine species, known to depress both species richness and abundance at a local scale in distinct geographic areas. It is a dominant biofouling agent, associated with the decreasing economic incomes from aquaculture operations around the world. Ciona intestinalis was first observed in 2007 at Straumsvík in Southwest Iceland. The present study was designed to provide general information on its current distribution across Icelandic harbours. The species was found only on the SW coast of Iceland, in dense aggregations reaching up to 876 ind/m2 in all harbours from Grindavík to Akranes. The current work provides information that can be used to track the dispersal of C. intestinalis populations along the Icelandic coast and for the development of effective management decisions.

Study of plant and tunicate based nanocrystalline cellulose in hybrid polymeric nanocomposites

Nanocrystalline cellulose (NCC) is a sustainable and renewable nanomaterial which can be defined as the nanoscale crystalline region of cellulose fibrils. The present work describes polyvinyl alcohol (PVA) based nanocomposite materials containing reinforcing low aspect ratio NCC sourced from plants and high aspect ratio NCC from a local invasive tunicate species. The effect of size and surface chemistry of the NCC reinforcement on the mechanical, thermal and morphological properties of PVA based nanocomposites is studied. Hybrid nanocomposites which contain both plant and tunicate sourced NCC within the PVA matrix are also documented. To our best knowledge, this represents the first known study describing the effect of NCC with varying size and surface chemistry combined within PVA to form hybrid PVA–NCC nanocomposites. In all cases, increasing mechanical reinforcement is observed as NCC content is increased, which varies in magnitude with NCC aspect ratio and surface chemistry. A notable improvement in thermal stability is also observed in all nanocomposites when NCC content is increased. Precursor materials and the resulting nanocomposites are well characterized by transmission electron microscopy, dynamic mechanical analysis, differential scanning calorimetry, thermogravimetric analysis, elemental analysis and powder X-ray diffraction.

The Application of a Mathematical Model to Evaluate the Effectiveness of Control Strategies Against Ciona intestinalis in Mussel Production.

The Prince Edward Island (PEI) mussel industry has faced challenges associated with invasive tunicate species over the past two decades. Field experiments to find suitable mitigation strategies require considerable time and are resource intensive. This study demonstrates the application of a mathematical model to assess several control strategies against Ciona intestinalis populations under different temperature conditions in a mussel production area in PEI. A temperature dependent compartmental model was used to model the total abundance of C. intestinalis. Each mitigation strategy was defined in terms of a combination of timing and frequency of treatments. Various strategies were explored to obtain the combination that maximized the difference in predicted abundances between the control (untreated) and the different mitigation strategies. Treatment frequency was allowed to vary between one and four times over a given production year. The model was assessed under baseline conditions, which mimicked water temperatures from Georgetown Harbor, PEI, in 2008; as well as under scenarios that reflected prolonged summer or warm spring temperatures. Furthermore, the sensitivity of the model to variations in presumed treatment efficacy was evaluated. The use of all four available treatments, starting around the first week of July and correctly timed thereafter, provided the most effective strategy, assuming the baseline temperature scenario. However, the effectiveness of this mitigation strategy depended on temperature conditions. The mathematical model developed in this study allows decision makers to explore different strategies to control the abundance of C. intestinalis in mussel production areas under different environmental conditions. In addition, the modeling framework developed could be adapted to simulate comparable ectoparasitic infestation in aquatic environments.

Invasive tunicate (Ascidiacea) metabolic and filtration rates in comparison to native tunicate and bivalve species

Invasive tunicate (Ascidiacea) metabolic and filtration rates in comparison to native tunicate and bivalve species

Association of the Mytilid Musculus viator with the Invasive Tunicate Ascidiella aspersa in San Matías Gulf, Argentine Patagonia

Ascidiella aspersa (Muller, 1776) is an exotic tunicate species widely distributed along the Sud-Occidental Atlantic coasts. In Patagonia waters, it forms dense aggregations at El Sotano (San Matias Gulf). We report an undescribed association between this invasive tunicate and a native mytilid species, Musculus viator (d'Orbigny, 1842). During a benthic study, a total of 543 individuals of A. aspersa were collected at El Sotano. A total of 689 individuals of M. viator were extracted from the tunics of 42 A. aspersa specimens. A maximum of 50 mytilid per tunicate was found. No relationship was found between the size of the tunicates and the size of M. viator. The prevalence of M. viator on A. aspersa individuals suggested that the tunicate invasion might have altered the community structure by improving local physical conditions -alternative substrate or refuge- for mytilid populations.

Compared stress tolerance to short-term exposure in native and invasive tunicates from the NE Atlantic: when the invader performs better

The combined impact of invasive species and climate change threatens natural systems worldwide, often facilitating the expansion of harmful invasive species. It is imperative to understand the mechanisms behind why species become invasive and widespread. Traditionally, it is thought that invasive species have greater tolerances to a wider array of environmental conditions than natives. We, therefore, tested the hypothesis that invasive species are more tolerant to the effects of short-term exposure to temperature and salinity stress. Using unifactorial experiments, we compared the tolerances of two common fouling NE Atlantic ascidians, the native Ciona intestinalis and the invasive Styela clava, to increased temperature and decreased salinity. We measured lethal and behavioural responses affecting 50% of populations to give an indication of the tolerance limits for temperature (LT50) and salinity (EC50), and respiration rate to give an indication of the change in metabolic response. The invasive S. clava was more tolerant to increased stress (LT50 = 29.5 °C, EC50 = 19.5) compared with C. intestinalis (LT50 = 27.0 °C, EC50 = 22.7), whereas both species displayed similar metabolic responses observed through increased respiration rates. This study is among the first to experimentally determine limits for temperature and hyposalinity stress for either species and supports the hypothesis that the invader performs better under extreme conditions. Future environmental changes caused by events such as heat waves and climate change could push species to the edge of their physiological limits, potentially facilitating competitive shifts between native and invasive species.

Invasive tunicate restructures invertebrate community on fishing grounds and a large protected area on Georges Bank

Marine invasive species can profoundly alter ecosystem processes by displacing native species and changing community structures. The invasive tunicate Didemnum vexillum was first found on the northern edge of Georges Bank in 1998. It can form encrusting colonies on gravel substrates that are also a preferred habitat for a number of other invertebrates. In this study we used data collected via HabCam, a vessel-towed underwater imaging system, to investigate the distribution of D. vexillum and its relationship to other epibenthic macroinvertebrates in a portion of Georges Bank that includes fishing grounds and an area protected from bottom-fishing. This novel technology provides imaging of epibenthic species distributions in areas of the benthic environment that were previously unobservable. We found that D. vexillum density is negatively correlated with the Atlantic sea scallop (Placopecten magellanicus), barnacles (genus Balanus), the tube anemone (genus Cerianthus), the green sea urchin (Strongylocentrotus droebachiensis), the globular sponge of the genus Polymastia, and Bryozoa. However D. vexillum is positively correlated with Cancer spp. Crabs, the tube forming polychaete, Filograna implexa, and Asterias spp. sea stars. The hypothesis that D. vexillum restructures the invertebrate community is supported by principal components analysis, revealing it as a primary driver of variation in the community when present. Additionally, there is an effect of the closed area as compared to fishing grounds on the structure of the invertebrate community and the abundance of certain species as consistent with previous studies, bottom-fishing affects invertebrate community structure. Principal components analysis revealed that bottom-fishing also appears to weaken clustering among species in the invertebrate community as compared to the community structure in the closed area. Biodiversity in high gravel sites of the epibenthic environment, as measured by the Shannon diversity index, also declined with increasing D. vexillum percent cover, while the open and closed areas were not significantly different in their level of biodiversity. D. vexillum appears to be the key driver of biodiversity decline in the epibenthos when present, rather than other processes such as direct disturbance and extraction from dredging. This research evaluates ecological responses to the presence of an invasive tunicate and suggests that this invasive species is a major force in shaping the ecological interactions in invaded areas.