Mads S. Thomsen

Seaweed communities in retreat from ocean warming

In recent decades, global climate change [1] has caused profound biological changes across the planet [2-6]. However, there is a great disparity in the strength of evidence among different ecosystems and between hemispheres: changes on land have been well documented through long-term studies, but similar direct evidence for impacts of warming is virtually absent from the oceans [3, 7], where only a few studies on individual species of intertidal invertebrates, plankton, and commercially important fish in the North Atlantic and North Pacific exist. This disparity of evidence is precarious for biological conservation because of the critical role of the marine realm in regulating the Earths environmental and ecological functions, and the associated socioeconomic well-being of humans [8]. We interrogated a database of >20,000 herbarium records of macroalgae collected in Australia since the 1940s and documented changes in communities and geographical distribution limits in both the Indian and Pacific Oceans, consistent with rapid warming over the past five decades [9, 10]. We show that continued warming might drive potentially hundreds of species toward and beyond the edge of the Australian continent where sustained retreat is impossible. The potential for global extinctions is profound considering the many endemic seaweeds and seaweed-dependent marine organisms in temperate Australia.

Decreasing resilience of kelp beds along a latitudinal temperature gradient: potential implications for a warmer future

Successful mitigation of negative effects of global warming will depend on understanding the link between physiological and ecological responses of key species. We show that while metabolic adjustment may assist Australasian kelp beds to persist and maintain abundance in warmer waters, it also reduces the physiological responsiveness of kelps to perturbation, and suppresses canopy recovery from disturbances by reducing the ecological performance of kelp recruits. This provides a warning not to rely solely on inventories of distribution and abundance to evaluate ecosystem function. The erosion of resilience is mediated by a shift in adult-juvenile interactions from competitive under cool to facilitative under warm conditions, supporting the prediction that positive interactions may become increasingly important in a warmer future. Kelp beds may remain intact but with a lower threshold for where additional impacts (e.g., extreme storms or reduced water quality) will lead to persistent loss of habitat and ecological function.

Climate-driven regime shift of a temperate marine ecosystem

No turning back? Ecosystems over time have endured much disturbance, yet they tend to remain intact, a characteristic we call resilience. Though many systems have been lost and destroyed, for systems that remain physically intact, there is debate as to whether changing temperatures will result in shifts or collapses. Wernburg et al. show that extreme warming of a temperate kelp forest off Australia resulted not only in its collapse, but also in a shift in community composition that brought about an increase in herbivorous tropical fishes that prevent the reestablishment of kelp. Thus, many systems may not be resilient to the rapid climate change that we face. Science, this issue p. 169 Rapid warming tropicalizes a temperate kelp forest. Ecosystem reconfigurations arising from climate-driven changes in species distributions are expected to have profound ecological, social, and economic implications. Here we reveal a rapid climate-driven regime shift of Australian temperate reef communities, which lost their defining kelp forests and became dominated by persistent seaweed turfs. After decades of ocean warming, extreme marine heat waves forced a 100-kilometer range contraction of extensive kelp forests and saw temperate species replaced by seaweeds, invertebrates, corals, and fishes characteristic of subtropical and tropical waters. This community-wide tropicalization fundamentally altered key ecological processes, suppressing the recovery of kelp forests.

Habitat Cascades: The Conceptual Context and Global Relevance of Facilitation Cascades via Habitat Formation and Modification

The importance of positive interactions is increasingly acknowledged in contemporary ecology. Most research has focused on direct positive effects of one species on another. However, there is recent evidence that indirect positive effects in the form of facilitation cascades can also structure species abundances and biodiversity. Here we conceptualize a specific type of facilitation cascade-the habitat cascade. The habitat cascade is defined as indirect positive effects on focal organisms mediated by successive facilitation in the form of biogenic formation or modification of habitat. Based on a literature review, we demonstrate that habitat cascades are a general phenomenon that enhances species abundance and diversity in forests, salt marshes, seagrass meadows, and seaweed beds. Habitat cascades are characterized by a hierarchy of facilitative interactions in which a basal habitat former (typically a large primary producer, e.g., a tree) creates living space for an intermediate habitat former (e.g., an epiphyte) that in turn creates living space for the focal organisms (e.g., spiders, beetles, and mites). We then present new data on a habitat cascade common to soft-bottom estuaries in which a relatively small invertebrate provides basal habitat for larger intermediate seaweeds that, in turn, generate habitat for focal invertebrates and epiphytes. We propose that indirect positive effects on focal organisms will be strongest when the intermediate habitat former is larger and different in form and function from the basal habitat former. We also discuss how humans create, modify, and destroy habitat cascades via global habitat destruction, climatic change, over-harvesting, pollution, or transfer of invasive species. Finally, we outline future directions for research that will lead to a better understanding of habitat cascades.

Evidence for impacts of nonindigenous macroalgae: a meta-analysis of experimental field studies.

Invasions by nonindigenous macroalgal species (NIMS) potentially cause severe impacts on native species. We conducted a meta‐analysis of 18 field‐based manipulative experiments to quantify the direction and magnitude of impacts (Hedges effect size d, hereafter ES). We found significant small‐to‐medium negative effects on “macrophyte abundance” (cover, biomass of native taxa; EScumulative = −0.30) and medium‐to‐large negative effects on “macrophyte assemblages” (richness, diversity, total abundance; EScumulative = −0.70). In contrast, EScumulative were not significant for “macrophyte processes” (growth, mortality; EScumulative = −0.39), “animal abundance” (densities; EScumulative = −0.13), or “animal assemblages” (richness, diversity; EScumulative = 0.75). The nonsignificant effect sizes were characterized by low sample sizes and should be interpreted with caution. Three study‐specific effect sizes were particularly large (<−2.0), showing that, in specific cases, impacts can be highly negative. From a conservation perspective, focus could be on such worst‐case scenarios. Still, the reported EScumulative are likely biased toward larger effects because only the most conspicuous NIMS have been tested and because nonsignificant results are less likely to be published. To better understand the impacts of NIMS, more manipulative experiments are needed, testing more species and under contrasting environmental conditions. Future studies should include procedural control treatments and report the abundance of the NIMS to avoid ambiguous interpretations. In conclusion, current experimental evidence shows that NIMS have, on average, small‐to‐large negative impacts on native plant species and assemblages. It is possible that these effects can result in severe consequences when accumulated over long time periods and large spatial scales.

A meta-analysis of seaweed impacts on seagrasses: Generalities and knowledge gaps

Seagrasses are important habitat-formers and ecosystem engineers that are under threat from bloom-forming seaweeds. These seaweeds have been suggested to outcompete the seagrasses, particularly when facilitated by eutrophication, causing regime shifts where green meadows and clear waters are replaced with unstable sediments, turbid waters, hypoxia, and poor habitat conditions for fishes and invertebrates. Understanding the situations under which seaweeds impact seagrasses on local patch scales can help proactive management and prevent losses at greater scales. Here, we provide a quantitative review of available published manipulative experiments (all conducted at the patch-scale), to test which attributes of seaweeds and seagrasses (e.g., their abundances, sizes, morphology, taxonomy, attachment type, or origin) influence impacts. Weighted and unweighted meta-analyses (Hedges d metric) of 59 experiments showed generally high variability in attribute-impact relationships. Our main significant findings were that (a) abundant seaweeds had stronger negative impacts on seagrasses than sparse seaweeds, (b) unattached and epiphytic seaweeds had stronger impacts than ‘rooted’ seaweeds, and (c) small seagrass species were more susceptible than larger species. Findings (a) and (c) were rather intuitive. It was more surprising that ‘rooted’ seaweeds had comparatively small impacts, particularly given that this category included the infamous invasive Caulerpa species. This result may reflect that seaweed biomass and/or shading and metabolic by-products like anoxia and sulphides could be lower for rooted seaweeds. In conclusion, our results represent simple and robust first-order generalities about seaweed impacts on seagrasses. This review also documented a limited number of primary studies. We therefore identified major knowledge gaps that need to be addressed before general predictive models on seaweed-seagrass interactions can be build, in order to effectively protect seagrass habitats from detrimental competition from seaweeds.

Epibiota communities of the introduced and indigenous macroalgal relatives Sargassum muticum and Halidrys siliquosa in Limfjorden (Denmark)

Sargassum muticum (Phaeophyceae, Fucales) has recently been introduced to Limfjorden (Denmark) where its closest relative is the indigenous Halidrys siliquosa. Previous studies have demonstrated large quantitative (canopy biomass) and qualitative (canopy persistence) differences in the habitat available to epibiota within the canopies of these two macroalgae. We therefore hypothesised that these algae would support different epibiota communities and tested this by sampling the epibiota of S. muticum and H. siliquosa on seven occasions throughout 1997 by enclosing entire thalli in mesh bags. We found 53 epibiota taxa and, with only one exception, they were all recorded on both host species. Species richness and abundance of epibiota exhibited clear seasonal variation on both host species, although epibiota biomass was seasonally constant on H. siliquosa but not on S. muticum. These patterns were consistent with the different life histories of the host species. There was a weakly negative correlation between thallus size and epibiota biomass for both host species. When taking species-specific seasonal variation in thallus size into consideration, S. muticum and H. siliquosa were found to support significantly different epibiota biomasses. Multivariate analyses showed that epibiota community structure was different, although highly overlapping, between the two species, whereas there was an almost parallel temporal development in epibiota community structure. We conclude that it is unlikely that the introduction of S. muticum to Limfjorden has caused major changes in local epibiota community structure. However, the standing stock of epibiota is likely to have increased.

GRACILARIA VERMICULOPHYLLA (RHODOPHYTA, GRACILARIALES) IN HOG ISLAND BAY, VIRGINIA: A CRYPTIC ALIEN AND INVASIVE MACROALGA AND TAXONOMIC CORRECTION1

Gracilaria in Virginia, USA, is abundant and composed of thalli either having relatively flat or cylindrical branches. These two morphologies were referred to previously as G. foliifera (Forsskål) Bøgesen and G. verrucosa (Hudson) Papenfuss. However, G. verrucosa is regarded an invalid name, and the flat specimens are now referred to as G. tikvahiae McLachlan. This has created confusion about the nomenclature of Gracilaria from this region. Here we document that the cylindrical form that dominates Hog Island Bay, Virginia, is G. vermiculophylla (Ohmi) Papenfuss, an alien macroalga from the West Pacific. Most of the ecological studies performed at the Long Term Ecological Research (LTER) site in Hog Island Bay used this cylindrical species. The present study clarifies the taxonomical status of this species, and we identify attributes that make this alien successful in turbid coastal lagoons.

Stress tolerance of the invasive macroalgae Codium fragile and Gracilaria vermiculophylla in a soft-bottom turbid lagoon

Invasive species are often hypothesized to have superior performance traits. We compared stress tolerance (as change in biomass) of the invasive macroalgae Codium fragile ssp. tomentosoides and Gracilaria vermiculophylla to the native macroalgae Fucus vesiculosus, Agardhiella subulata, Hypnea musciformis and Ulva curvata in Hog Island Bay, a shallow lagoon in Virginia, USA. We hypothesized that the success of the two aliens is due to their high tolerances of turbidity, sedimentation, desiccation, grazing and nutrient enrichment. Like many lagoons, Hog Island Bay is characterized by extensive intertidal mudflats, high turbidity and sedimentation, and high densities of omnivorous mud snails. Nutrient enrichment may also become a problem as land use practices in adjacent watersheds change. Contrary to our hypothesis, C. fragile was less resistant to sedimentation, desiccation and grazing than other algae and had low growth at all light and nutrient levels. This suggests that any superior performance of this invasive species compared to native algae is probably limited to microhabitats where stress is minimal and where bivalve shells facilitate recruitment and long-term persistence. In contrast, G. vermiculophylla was resistant to desiccation, burial and grazing, and was not negatively influenced by either high or low light or nutrient levels. These traits reflect the current success of G. vermiculophylla in already invaded lagoons and estuaries, and indicates that it will likely continue its spread in European and North American turbid and tidal soft-sediment systems.

The effect of thallus size, life stage, aggregation, wave exposure and substratum conditions on the forces required to break or dislodge the small kelp Ecklonia radiata

Abstract Canopy removal by storms is a primary cause of mortality for the small kelp Ecklonia radiata in temperate Australasia. We simulated hydrodynamic drag from storms with in situ pull-tests to determine whether thallus size, life stage, aggregation, wave exposure and substratum affect the canopy removal process. A total of 466 individuals were pulled off 20 rocky reefs at 8–10 m depth in southwestern Australia. The majority (71%) of thalli dislodged at the rock implying that the canopy removal process in southwestern Australia is a substratum-controlled process. Dislodgment bared clean substratum where re-invasion by propagules or encroachment would be necessary to fill up the gaps. Maximum break forces (150–250 N) were found for large late stage kelps and kelp aggregates from wave exposed sandstone and granite reefs, and minimum values (25–100 N) for small juveniles and solitary kelps from protected limestone reefs. By applying size and break force data to the drag equation, water velocities required to break or dislodge E. radiata were calculated to 2–5 ms-1 for large kelps. These velocities are frequently encountered in wave-exposed shallow subtidal habitats, suggesting that thallus size is limited by the hydrodynamic environment.