Primary production of the kelp Lessonia corrugata varies with season and water motion: Implications for coastal carbon cycling.

Kelp forests provide vital ecosystem services such as carbon storage and cycling, and understanding primary production dynamics regarding seasonal and spatial variations is essential. We conducted surveys at three sites in southeast Tasmania, Australia, that had different levels of water motion, across four seasons to determine seasonal primary production and carbon storage as living biomass for kelp beds of Lessonia corrugata (Order Laminariales). We quantified blade growth, erosion rates, and the variation in population density and estimated both the net biomass accumulation (NBA) per square meter and the carbon standing stock. We observed a significant difference in blade growth and erosion rates between seasons and sites. Spring had the highest growth rate (0.02 g C · blade-1 · d-1 ) and NBA (1.62 g C · m-2 · d-1 ), while summer had the highest blade erosion (0.01 g C · blade-1 · d-1 ), with a negative NBA (-1.18 g C · m-2 · d-1 ). Sites exhibiting lower blade erosion rates demonstrated notably greater NBA than sites with elevated erosion rates. The sites with the highest water motion had the slowest erosion rates. Moreover, the most wave-exposed site had the densest populations, resulting in the highest NBA and a greater standing stock. Our results reveal a strong seasonal and water motion influence on carbon dynamics in L. corrugata populations. This knowledge is important for understanding the dynamics of the carbon cycle in coastal regions.

Thermal performance curves identify seasonal and site-specific variation in the development of Ecklonia radiata (Phaeophyceae) gametophytes and sporophytes.

Rapid ocean warming is affecting kelp forests globally. While the sporophyte life stage has been well studied for many species, the microscopic life stages of laminarian kelps have been understudied, particularly regarding spatial and temporal variations in thermal tolerance and their interaction. We investigated the thermal tolerance of growth, survival, development, and fertilization of Ecklonia radiata gametophytes, derived from zoospores sampled from two sites in Tasmania, Australia, throughout a year, over a temperature gradient (3-30°C). For growth we found a relatively stable thermal optimum at ~20.5°C and stable thermal maxima (25.3-27.7°C). The magnitude of growth was highly variable and depended on season and site, with no consistent spatial pattern for growth and gametophyte size. Survival also had a relatively stable thermal optimum of ~17°C, 3°C below the optimum for growth. Gametophytes grew to single cells between 5 and 25°C, but sporophytes were only observed between 10 and 20°C, indicating reproductive failure outside this range. The results reveal complex effects of source population and season of collection on gametophyte performance in E. radiata, with implications when comparing results from material collected at different localities and times. In Tasmania, gametophytes grow considerably below the estimated thermal maxima and thermal optima that are currently only reached during summer heatwaves, whereas optima for survival (~17°C) are frequently reached and surpassed during heatwaves, which may affect the persistence and recruitment of E. radiata in a warmer climate.

Density-dependence and seasonal variation in reproductive output and sporophyte production in the kelp, Ecklonia radiata.

The kelp, Ecklonia radiata, is an abundant subtidal ecosystem engineer in southern Australia. Density-dependent changes in the abiotic environment engineered by Ecklonia may feedback to affect reproduction and subsequent recruitment. Here, we examined: 1) how the reproductive capacity of Ecklonia individuals in the field (zoospores released · mm-2 reproductive tissue) varied with adult density and time, and 2) how the recruitment of microscopic gametophytes and sporophytes was influenced by zoospore density at two times. Zoospore production did not vary with adult density, with only one month out of ten sampled over a 2-y period showing a significant effect of density. However, zoospore production varied hugely over time, being generally highest in mid-autumn and lowest in mid-late summer. There were strong effects of initial zoospore density on gametophyte and sporophyte recruitment with both a minimum and an optimum zoospore density for sporophyte recruitment, but these varied in time. Almost no sporophytes developed when initial zoospore density was <6.5 · mm-2 in spring or <0.5 · mm-2 in winter with optimum densities of 90-355 · mm-2 in spring and 21-261 · mm-2 in winter, which resulted in relatively high recruitment of 4-7 sporophytes · mm-2 . Sporophyte recruitment declined at initial zoospore densities >335 · mm-2 in spring and >261 · mm-2 in winter and was zero at very high zoospore densities. These findings suggest that although adult Ecklonia density does not affect per-capita zoospore production, because there is a minimum zoospore density for sporophyte production, a decline in population-level output could feedback to impact recruitment.

Plastic and natural inorganic microparticles do not differ in their effects on adult mussels (Mytilidae) from different geographic regions.

Microplastics are ubiquitous in the marine environment and studies on their effects on benthic filter feeders at least partly revealed a negative influence. However, it is still unclear whether the effects of microplastics differ from those of natural suspended microparticles, which constitute a common stressor in many coastal environments. We present a series of experiments that compared the effects of six-week exposures of marine mussels to two types of natural particles (red clay and diatom shells) to two types of plastic particles (Polymethyl Methacrylate and Polyvinyl Chloride). Mussels of the family Mytilidae from temperate regions (Japan, Chile, Tasmania) through subtropical (Israel) to tropical environments (Cabo Verde) were exposed to concentrations of 1.5 mg/L, 15 mg/L and 150 mg/L of the respective microparticles. At the end of this period, we found significant effects of suspended particles on respiration rate, byssus production and condition index of the animals. There was no significant effect on clearance rate and survival. Surprisingly, we observed only small differences between the effects of the different types of particles, which suggests that the mussels were generally equally robust towards exposure to variable concentrations of suspended solids regardless of whether they were natural or plastic. We conclude, that microplastics and suspended solids elicit similar effects on the tested response variables, and that both types of microparticles mainly cause acute responses rather than more persistent carry-over effects.

Interactive effects of canopy-driven changes in light, scour and water flow on microscopic recruits in kelp.

Ecosystem engineering kelp forms habitat and influences associated communities by altering abiotic conditions. These conditions can also affect the engineer's own demographic rates but the mechanisms underpinning these feedbacks are not well known. Here, we tested the interactive effects of three abiotic factors engineered by the Australasian kelp Ecklonia radiata (light, water flow and scour) on the early survivorship and growth of its outplanted microscopic recruits. After six weeks, recruit survivorship was high in the absence of scour and low light (2-3 times higher than when scour was present) and under low water flow-ambient light conditions. Growth of sporophytes was strongly related to light, with recruits under ambient light approximately four times larger after six weeks. Overall, reduced scour (for survivorship) and ambient light (for growth) appear crucial for maximising E. radiata recruitment suggesting a healthy forest can provide microenvironments to enhance survivorship while gaps in the canopy enhance growth.

Demography of the Intertidal Fucoid Hormosira banksii: Importance of Recruitment to Local Abundance.

Canopy-forming macroalgae form the basis of diverse coastal ecosystems globally. The fucoid Hormosira banksii is often the dominant canopy-forming macroalga in the temperate intertidal of southern Australia and New Zealand, where it is commonly associated with an understory of coralline turf. Hormosira banksii is susceptible to both natural and anthropogenic disturbance and despite its abundance, few studies have examined the demography of this important species. This study determined the demographic response of H. banksii to different gradients of disturbance to both its canopy and to the understory coralline turf. We established plots in which the density of H. banksii and/or understory coralline turf was manipulated in a pulse perturbation to simulate a disturbance event. The manipulated plots contained eight treatments ranging from 100% removal of H. banksii to 100% removal of the understory coralline turf. We then measured recruitment and followed individual recruits for up to 18 months to determine growth and survivorship. We found that H. banksii recruitment was seasonally variable throughout the experiment and highest over summer, survivorship of recruits was generally high, and the species was slow-growing and long-lived. Moreover, the level of disturbance did not seem to affect recruitment, growth, or survivorship and post-recruitment mortality was independent of H. banksii density. In this system, it appears that H. banksii is a relatively long-lived perennial species whose demography is density-independent which appears to allow recovery from disturbance.

A comparison with natural particles reveals a small specific effect of PVC microplastics on mussel performance.

Effects of microplastics on marine taxa have become a focal point in marine experimental biology. Almost all studies so far, however, assessed the influence of microplastics on animals only in relation to a zero-particle group. Documented microplastic impacts may thus be overestimated, since many marine species also experience natural suspended solids as a stressor. Here, we compared the effects of polyvinyl chloride (PVC) and red clay (mean for both particles: ~12-14 µm) on the Mediterranean mussel Mytilus galloprovincialis across three particle concentrations (1.5, 15, 150 mg l-1). Exposure to PVC for 35 days lowered mussel body condition index by 14% in relation to clay, but no difference in byssus production, respiration and survival rates emerged between the two particle types. This suggests that the effects of synthetic particles on filter feeders may emulate those of natural suspended solids, and highlights the importance of including natural particles in microplastic exposure studies.

Dataset for concurrent echosounder and ADCP measurements at a tidal energy candidate site in Australia.

Interaction uncertainties between tidal energy devices and marine animals have the potential to impede the tidal energy industry as it moves closer towards commercial-scale array installations. Developing standardised environmental impact assessment (EIA) practices would allow for potential impact concerns to the marine environment to be identified and mitigated early during project development. In an effort to help formulate a standardised EIA framework that addresses knowledge gaps in fish-current interactions at tidal energy candidate sites, Scherelis et al. [1] presented a case study for investigating changes in fish aggregations in response to changing environmental conditions including tidal currents at a tidal energy candidate site in Australia prior to turbine installation. Here, we present the dataset utilised for this study titled "Investigating biophysical linkages at tidal energy candidate sites: a case study for combining environmental assessment and resource characterisation" [1]. The dataset includes tidal current information from an Acoustic Doppler Current Profiler (ADCP), volume backscattering measurements from a four-frequency biological echosounder (Acoustic Zooplankton and Fish Profiler - AZFP) as an indicator for fish biomass, and fish aggregation metrics calculated from volume backscatter in post-processing. ADCP and AZFP were installed on a bottom-mounted mooring and engaged in a concurrent sampling plan for ∼2.5 months from December 2018 to February 2019. The mooring was deployed in the Banks Strait, a tidal energy candidate site located in the northeast of Tasmania, Australia, at a location favourable for tidal turbine installations considering current speed, depth, substrate, sediment type and proximity to shore. The ADCP dataset includes current velocity and direction measurements at 1 m vertical and 1-min time intervals. The raw AZFP dataset includes volume backscattering strength collected in 4-s time intervals with a vertical resolution of 0.072 m in raw, and 0.1 m in pre-processed form. Several post-processing steps were implemented to mitigate changes in background noise due to current speed and wind stress, and to isolate acoustic fish returns from remaining scattering sources. Once isolated, volume backscatter containing fish targets underwent post-processing to determine fish aggregation metrics including density, abundance, centre of mass, dispersion,% water column occupied, evenness, and index for aggregation. Each aggregation metric was then binned by minute matched with corresponding environmental conditions for current speed, shear, temperature, diel stage, and tidal stage. Raw and processed datasets for the AZFP and ADCP are provided. Post-processed data includes the derived fish aggregation metrics along with corresponding environmental conditions. The described datasets are freely available on the Australian Ocean Data Network (AODN).

Habitat provided by native species facilitates higher abundances of an invader in its introduced compared to native range.

The impacts invasive species have on biodiversity and ecosystem function globally have been linked to the higher abundances they often obtain in their introduced compared to native ranges. Higher abundances of invaders in the introduced range are often explained by a reduction in negative species interactions in that range, although results are equivocal. The role of positive interactions in explaining differences in  the abundance of invaders between native and invasive ranges has not been tested. Using biogeographic surveys, we showed that the rocky shore porcelain crab, Petrolisthes elongatus, was ~4 times more abundant in its introduced (Tasmania, Australia) compared to its native (New Zealand) range. The habitat of these crabs in the invaded range (underside of intertidal boulders) was extensively covered with the habitat-forming tubeworm Galeolaria caespitosa. We tested whether the habitat provided by the tubeworm facilitates a higher abundance of the invasive crab by creating mimics of boulders with and without the tubeworm physical structure and measured crab colonisation into these habitats at three sites in both Tasmania and New Zealand. Adding the tubeworm structure increased crab abundance by an average of 85% across all sites in both ranges. Our intercontinental biogeographic survey and experiment demonstrate that native species can facilitate invader abundance and that positive interactions can be important drivers of invasion success.

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.

Resilience and stability of kelp forests: The importance of patch dynamics and environment-engineer feedbacks.

Habitat forming 'ecosystem engineers' such as kelp species create complex habitats that support biodiverse and productive communities. Studies of the resilience and stability of ecosystem engineers have typically focussed on the role of external factors such as disturbance. However, their population dynamics are also likely to be influenced by internal processes, such that the environmental modifications caused by engineer species feedback to affect their own demography (e.g. recruitment, survivorship). In numerous regions globally, kelp forests are declining and experiencing reductions in patch size and kelp density. To explore how resilience and stability of kelp habitats is influenced by this habitat degradation, we created an array of patch reefs of various sizes and supporting adult Ecklonia radiata kelp transplanted at different densities. This enabled testing of how sub-canopy abiotic conditions change with reductions in patch size and adult kelp density, and how this influenced demographic processes of microscopic and macroscopic juvenile kelp. We found that ecosystem engineering by adult E. radiata modified the environment to reduce sub-canopy water flow, sedimentation, and irradiance. However, the capacity of adult kelp canopy to engineer abiotic change was dependent on patch size, and to a lesser extent, kelp density. Reductions in patch size and kelp density also impaired the recruitment, growth and survivorship of microscopic and macroscopic juvenile E. radiata, and even after the provisioning of established juveniles, demographic processes were impaired in the absence of sufficient adult kelp. These results are consistent with the hypothesis that ecosystem engineering by adult E. radiata facilitates development of juvenile conspecifics. Habitat degradation seems to impair the ability of E. radiata to engineer abiotic change, causing breakdown of positive intraspecific feedback and collapse of demographic functions, and overall, leading to reductions in ecosystem stability and resilience well before local extirpation.

Family-level variation in early life-cycle traits of kelp.

Temperate kelp forests (Laminarians) are threatened by temperature stress due to ocean warming and photoinhibition due to increased light associated with canopy loss. However, the potential for evolutionary adaptation in kelp to rapid climate change is not well known. This study examined family-level variation in physiological and photosynthetic traits in the early life-cycle stages of the ecologically important Australasian kelp Ecklonia radiata and the response of E. radiata families to different temperature and light environments using a family × environment design. There was strong family-level variation in traits relating to morphology (surface area measures, branch length, branch count) and photosynthetic performance (Fv /Fm ) in both haploid (gametophyte) and diploid (sporophyte) stages of the life-cycle. Additionally, the presence of family × environment interactions showed that offspring from different families respond differently to temperature and light in the branch length of male gametophytes and oogonia surface area of female gametophytes. Negative responses to high temperatures were stronger for females vs. males. Our findings suggest E. radiata may be able to respond adaptively to climate change but studies partitioning the narrow vs. broad sense components of heritable variation are needed to establish the evolutionary potential of E. radiata to adapt under climate change.

Stronger positive association between an invasive crab and a native intertidal ecosystem engineer with increasing wave exposure.

Ecosystem engineers are predicted to have stronger facilitative effects when environmental stress is higher. Here we examined whether facilitation of the invasive porcelain crab Petrolisthes elongatus by the ecosystem engineering serpulid tube worm Galeolaria caespitosa increased with wave exposure. Petrolisthes occurs beneath intertidal boulders which often have a high cover of Galeolaria on their underside. Surveys across nine sites demonstrated Petrolisthes abundance beneath boulders increased with wave exposure and Galeolaria cover, although only when the habitat matrix beneath boulders was rock or mixed rock and sand. Moreover, as wave exposure increased, the strength of relationship between Petrolisthes abundance and the surface area of Galeolaria also increased. Experimentally, the presence of Galeolaria on the underside of boulders increased Petrolisthes abundance by 50% compared to boulders lacking Galeolaria. Our findings suggest the facilitative role of Galeolaria is stronger at more wave-exposed sites, which appears to contribute to a higher abundance of invasive Petrolisthes.

Local densities and habitat preference of the critically endangered spotted handfish (Brachionichthys hirsutus): Large scale field trial of GPS parameterised underwater visual census and diver attached camera.

The critically endangered spotted handfish (Brachionichthys hirsutus) is restricted to a limited number of locations in south-eastern Tasmania, Australia. As is often the case for rare species, conducting statistically adequate surveys for B. hirsutus can be costly and time consuming due to the low probability of encountering individuals. For the first time we used a highly efficient and rigorous Global Positioning System (GPS) parameterised underwater visual census (GUVC) to survey B. hirsutus abundance within all nine known local populations in the Derwent Estuary within one season. In addition, a benthic microhabitat assessment was conducted simultaneously using a GoPro® camera attached to diver to determine B. hirsutus microhabitat preferences. B. hirsutus local populations varied between sites, with densities ranging from 1.58 to 43.0 fishes per hectare. B. hirsutus demonstrates a strong preference for complex microhabitat features, such as depressions and ripple formations filled with biogenic substrates (e.g. shells) but avoids simple, low relief microhabitats (e.g. sand flats) and areas dominated by ephemeral, filamentous algae. Complex microhabitats may enable B. hirsutus to avoid predators, increase forage opportunities or provide higher quality spawning sites. This first wide-scale application of GUVC for B. hirsutus allowed us to survey a larger number of sites than previously possible to provide a robust reference point for future long-term monitoring.

Native turncoats and indirect facilitation of species invasions.

At local scales, native species can resist invasion by feeding on and competing with would-be invasive species. However, this relationship tends to break down or reverse at larger scales. Here, we consider the role of native species as indirect facilitators of invasion and their potential role in this diversity-driven 'invasion paradox'. We coin the term 'native turncoats' to describe native facilitators of non-native species and identify eight ways they may indirectly facilitate species invasion. Some are commonly documented, while others, such as indirect interactions within competitive communities, are largely undocumented in an invasion context. Therefore, we use models to evaluate the likelihood that these competitive interactions influence invasions. We find that native turncoat effects increase with the number of resources and native species. Furthermore, our findings suggest the existence, abundance and effectiveness of native turncoats in a community could greatly influence invasion success at large scales.

Historical demography and colonization pathways of the widespread intertidal seaweed Hormosira banksii (Phaeophyceae) in southeastern Australia.

The palaeoceanography of southern Australia has been characterized by fluctuating sea levels during glacial periods, changing temperature regimes and modified boundary currents. Previous studies on genetic structuring of species in southeastern Australia have focused mainly on the differentiation of eastern and western populations while the potential role of Bass Strait as a region of overlap for three biogeographic provinces (Peronia, Maugea, and Flindersia) has been largely ignored. This study aimed to explore the likely roles of historic and contemporary factors in determining divergence patterns in the habitat-forming intertidal seaweed Hormosira banksii in southeastern Australia with a special focus on postglacial dispersal into Bass Strait. We examined the genetic diversity of 475 Hormosira specimens collected from 19 sites around southern Australia using DNA sequence analysis of cytochrome oxidase 1. Three major haplotype groups were identified (western, centre and eastern) corresponding with the three existing biogeographical provinces in this region. Historic break points appeared to be retained and reinforced by modern day dispersal barriers. Phylogeographic grouping of Hormosira reflected a combination of historic and contemporary oceanography. As western and eastern group haplotypes were largely absent within Bass Strait, re-colonization after the last glacial maximum appeared to have originated from refuges within or near present day Bass Strait. Patterns of genetic structure for Hormosira are consistent with other marine species in this region and highlight the importance of biogeographical barriers in contributing to modern genetic structure.

Disturbance-mediated facilitation by an intertidal ecosystem engineer.

Ecosystem engineers facilitate communities by providing a structural habitat that reduces abiotic stress or predation pressure for associated species. However, disturbance may damage or move the engineer to a more stressful environment, possibly increasing the importance of facilitation for associated communities. In this study, we determined how disturbance to intertidal boulders (i.e., flipping) and the subsequent movement of a structural ecosystem engineer, the tube-forming serpulid worm Galeolaria caespitosa, from the bottom (natural state, low abiotic stress) to the top (disturbed state, high abiotic stress) surface of boulders influenced the importance of facilitation for intertidal communities across two intertidal zones. Theory predicts stronger relative facilitation should occur in the harsher environments of the top of boulders and the high intertidal zone. To test this prediction, we experimentally positioned boulders with the serpulids either face up or face down for 12 months in low and high zones in an intertidal boulder field. There were very different communities associated with the different boulders and serpulids had the strongest facilitative effects on the more stressful top surface of boulders with approximately double the species richness compared to boulders lacking serpulids. Moreover, within the serpulid matrix itself there was also approximately double the species richness (both zones) and abundance (high zone only) of small invertebrates on the top of boulders compared to the bottom. The high relative facilitation on the top of boulders reflected a large reduction in temperature by the serpulid matrix on that surface (up to 10°C) highlighting a key role for modification of the abiotic environment in determining the community-wide facilitation. This study has demonstrated that disturbance and subsequent movement of an ecosystem engineer to a more stressful environment increased the importance of facilitation and allowed species to persist that would otherwise be unable to survive in that environment.

Relationships between an invasive crab, habitat availability and intertidal community structure at biogeographic scales.

At local scales, habitat availability influences interactions between native and invasive species. Habitat availability may also predict patterns in native communities and invasive species at biogeographic scales when both native and invasive species have specific habitat requirements. The New Zealand porcelain crab, Petrolisthes elongatus, has invaded intertidal rocky shores around Tasmania, Australia, where it is found in high densities (>1800 m(2)) under rocks. A hierarchical sampling approach was used to investigate 1) the relationship between habitat availability (rock cover) and the biomass and abundance of P. elongatus, and 2) the relationship between P. elongatus biomass and native communities at local and regional scales. Invertebrate communities and habitat availability were sampled at multiple sites in the north and south regions of Tasmania. P. elongatus biomass and abundance were positively correlated with rock cover and patterns were consistent at the biogeographic scale (between regions). P. elongatus biomass was positively correlated with native species richness, biomass and abundance highlighting their co-dependence on rock cover. However, multivariate analyses indicated a different native community structure with increasing P. elongatus biomass. Flat, strongly adhering gastropods (chitons and limpets) were positively correlated with P. elongatus biomass, whereas mobile gastropods and crabs were negatively correlated with P. elongatus biomass. Despite local scale variation, there were clear consistent relationships between habitat-availability and the biomass of P. elongatus, and between native communities and the biomass of P. elongatus suggesting that the relationships between native and invasive species may be predictable at large spatial scales. Moreover, the strong relationships between P. elongatus biomass and changes in native community structure suggest a greater understanding of its impact is needed so that appropriate management plans can be developed.

Environmental correlates of phenotypic variation: do variable tidal regimes influence morphology in intertidal seaweeds?

Seaweed morphology is often shaped by the hydrodynamic environment. However, exposure to air at low tide represents an additional factor potentially affecting the morphology of intertidal species. Here, we examined the relationships between the morphology of Hormosira banksii, an important intertidal habitat-forming seaweed in southern Australia, and environmental factors across multiple spatial scales around the island of Tasmania, Australia. Tasmania is surrounded by a diverse coastline with differences in wave exposure, tidal parameters, and temperature. We sampled Hormosira from four regions (100s km apart), three sites (10s km apart) within each region, and two zones (meters apart; eulittoral and sublittoral) at each site, and measured multiple morphological variables to test for differences in morphology at those different spatial scales. Thirteen environmental variables reflecting wave exposure, tidal conditions, and temperature for each site were generated to assess the relationship between Hormosira morphology and environmental variation. Morphology varied at all spatial scales examined. Most notably, north coast individuals had a distinct morphology, generally having smaller vesicles and shorter fronds, compared to other regions. Tidal conditions were the main environmental factors separating north coast sites from other sites and tidal regime was identified as the best predictor of morphological differences between regions. In contrast to other studies, we found little evidence that wave exposure was associated with morphological variation. Overall, our study emphasizes the role of tidal conditions, associated with emersion stress during low tide, in affecting the morphology of intertidal seaweeds.

Phenotypic plasticity and biogeographic variation in physiology of habitat-forming seaweed: response to temperature and nitrate.

Southeastern Australian waters are warming at nearly four times the global average rate (~0.7°C · century(-1) ) driven by strengthening incursions of the warm oligotrophic East Australian Current. The growth rate hypothesis (GRH) predicts that nutrient depletion will impact more severely on seaweeds at high latitudes with compressed growth seasons. This study investigates the effects of temperature and nutrients on the ecophysiology of the habitat-forming seaweed Phyllospora comosa in a laboratory experiment using temperature (12°C, 17°C, 22°C) and nutrient (0.5, 1.0, 3.0 µM NO3 (-) ) scenarios representative of observed variation among geographic regions. Changes in growth, photosynthetic characteristics (via chlorophyll fluorescence), pigment content, tissue chemistry (δ(13) C, % C, % N, C:N) and nucleic acid characteristics (absolute RNA and DNA, RNA:DNA ratios) were determined in seaweeds derived from cool, high-latitude and warm, low-latitude portions of the species' range. Performance of P. comosa was unaffected by nitrate availability but was strongly temperature-dependent, with photosynthetic efficiency, growth, and survival significantly impaired at 22°C. While some physiological processes (photosynthesis, nucleic acid, and accessory pigment synthesis) responded rapidly to temperature, others (C/N dynamics, carbon concentrating processes) were largely invariant and biogeographic variation in these characteristics may only occur through genetic adaptation. No link was detected between nutrient availability, RNA synthesis and growth, and the GRH was not supported in this species. While P. comosa at high latitudes may be less susceptible to oligotrophy than predicted by the GRH, warming water temperatures will have deleterious effects on this species across its range unless rapid adaptation is possible.