The effect of sedimentation on spore settlement and recruitment of the endemic Arctic kelp, Laminaria solidungula (Phaeophyceae).

Environmental changes associated with rapid climate change in the Arctic, such as the increased rates of sedimentation from climatic or anthropogenic sources, can enhance the impact of abiotic stressors on coastal ecosystems. High sedimentation rates can be detrimental to nearshore kelp abundance and distribution, possibly due to increased mortality at the spore settlement stage. Spore settlement and viability of the Arctic kelp Laminaria solidungula were examined through a series of lab-based sedimentation experiments. Spores were exposed to increasing sediment loads in three experimental designs simulating different sedimentation scenarios: sediment deposition above settled spores, settlement of spores on sediment-covered substrate, and simultaneous suspension of spores and sediments during settlement. Spore settlement was recorded upon completion of each experiment, and gametophyte abundance was assessed following a growth period with sediments removed to examine short-term spore viability via a gametophyte-to-settled-spore ratio. In all three types of sediment exposure, the addition of sediments caused a 30%-40% reduction in spore settlement relative to a no-sediment control. Spore settlement decreased significantly between the low and high sediment treatments when spores were settled onto sediment-covered substrates. In all experiments, increasing amounts of sediment had no significant effect on spore viability, indicating that spores that had settled under different short-term sediment conditions were viable. Our results indicate that depending on spore-sediment interaction type, higher rates of sedimentation resulting from increased sediment loading could affect L. solidungula spore settlement success with potential impacts on the long-term persistence of a diverse and productive benthic habitat.

Gersemiols A-C and Eunicellol A, Diterpenoids from the Arctic Soft Coral Gersemia fruticosa.

Three new diterpenes named gersemiols A-C (1-3) and a new eunicellane diterpene, eunicellol A (4), have been isolated together with the known sesquiterpene (+)-α-muurolene (5) from the Arctic soft coral Gersemia fruticosa. The name gersemiane was assigned to the rare and unnamed diterpene skeleton of compounds 1-3 corresponding to 4-isopropyl-1,5,8a-trimethyltetradecahydrophenanthrene. The chemical structures were elucidated on the basis of extensive spectroscopic analysis (HR-ESIMS, 1D and 2D NMR) as well as coupling constant calculations for the determination of the relative configurations. All compounds were tested for their antimicrobial activity against several bacteria and fungi and eunicellol A was found to exhibit moderate and selective antibacterial activity.

Biodiversity mediates top-down control in eelgrass ecosystems: a global comparative-experimental approach.

Nutrient pollution and reduced grazing each can stimulate algal blooms as shown by numerous experiments. But because experiments rarely incorporate natural variation in environmental factors and biodiversity, conditions determining the relative strength of bottom-up and top-down forcing remain unresolved. We factorially added nutrients and reduced grazing at 15 sites across the range of the marine foundation species eelgrass (Zostera marina) to quantify how top-down and bottom-up control interact with natural gradients in biodiversity and environmental forcing. Experiments confirmed modest top-down control of algae, whereas fertilisation had no general effect. Unexpectedly, grazer and algal biomass were better predicted by cross-site variation in grazer and eelgrass diversity than by global environmental gradients. Moreover, these large-scale patterns corresponded strikingly with prior small-scale experiments. Our results link global and local evidence that biodiversity and top-down control strongly influence functioning of threatened seagrass ecosystems, and suggest that biodiversity is comparably important to global change stressors.

Fatty acid and stable isotope characteristics of sea ice and pelagic particulate organic matter in the Bering Sea: tools for estimating sea ice algal contribution to Arctic food web production.

We determined fatty acid (FA) profiles and carbon stable isotopic composition of individual FAs (δ(13)CFA values) from sea ice particulate organic matter (i-POM) and pelagic POM (p-POM) in the Bering Sea during maximum ice extent, ice melt, and ice-free conditions in 2010. Based on FA biomarkers, differences in relative composition of diatoms, dinoflagellates, and bacteria were inferred for i-POM versus p-POM and for seasonal succession stages in p-POM. Proportions of diatom markers were higher in i-POM (16:4n-1, 6.6-8.7%; 20:5n-3, 19.6-25.9%) than in p-POM (16:4n-1, 1.2-4.0%; 20:5n-3, 5.5-14.0%). The dinoflagellate marker 22:6n-3/20:5n-3 was highest in p-POM. Bacterial FA concentration was higher in the bottom 1 cm of sea ice (14-245 µg L(-1)) than in the water column (0.6-1.7 µg L(-1)). Many i-POM δ(13)C(FA) values were higher (up to ~10‰) than those of p-POM, and i-POM δ(13)C(FA) values increased with day length. The higher i-POM δ(13)C(FA) values are most likely related to the reduced dissolved inorganic carbon (DIC) availability within the semi-closed sea ice brine channel system. Based on a modified Rayleigh equation, the fraction of sea ice DIC fixed in i-POM ranged from 12 to 73%, implying that carbon was not limiting for primary productivity in the sympagic habitat. These differences in FA composition and δ(13)C(FA) values between i-POM and p-POM will aid efforts to track the proportional contribution of sea ice algal carbon to higher trophic levels in the Bering Sea and likely other Arctic seas.

Reduced efficiency of pelagic-benthic coupling in the Arctic deep sea during lower ice cover.

Pelagic-benthic coupling describes the connection between surface-water production and seafloor habitats via energy, nutrient and mass exchange. Massive ice loss and warming in the poorly studied Arctic Chukchi Borderland are hypothesized to affect this coupling. The strength of pelagic-benthic coupling was compared between 2 years varying in climate settings, 2005 and 2016, based on δ13C and δ15N stable isotopes of food-web end-members and pelagic and deep-sea benthic consumers. Considerably higher isotopic niche overlap and generally shorter isotopic distance were found between pelagic and benthic food web components in 2005 than in 2016, suggesting weaker coupling in the latter, low-ice year. δ15N values indicated more refractory food consumed by benthos in 2016 and fresher food reaching the seafloor in 2005. Higher δ13C values of zooplankton indirectly suggested a higher contribution of ice algae in 2005 than 2016. The difference in pelagic-benthic coupling between these years is consistent with higher energy retention within the pelagic system, perhaps due to strong stratification in the Amerasian Basin in the recent decade. Weaker coupling to the benthos can be expected to continue with ice loss in the study area, perhaps reducing benthic biomass and remineralization capacity; monitoring of the area is needed to confirm this prediction.

Ecosystem response persists after a prolonged marine heatwave.

Some of the longest and most comprehensive marine ecosystem monitoring programs were established in the Gulf of Alaska following the environmental disaster of the Exxon Valdez oil spill over 30 years ago. These monitoring programs have been successful in assessing recovery from oil spill impacts, and their continuation decades later has now provided an unparalleled assessment of ecosystem responses to another newly emerging global threat, marine heatwaves. The 2014-2016 northeast Pacific marine heatwave (PMH) in the Gulf of Alaska was the longest lasting heatwave globally over the past decade, with some cooling, but also continued warm conditions through 2019. Our analysis of 187 time series from primary production to commercial fisheries and nearshore intertidal to offshore oceanic domains demonstrate abrupt changes across trophic levels, with many responses persisting up to at least 5 years after the onset of the heatwave. Furthermore, our suite of metrics showed novel community-level groupings relative to at least a decade prior to the heatwave. Given anticipated increases in marine heatwaves under current climate projections, it remains uncertain when or if the Gulf of Alaska ecosystem will return to a pre-PMH state.

Palmadorins A--C, diterpene glycerides from the antarctic nudibranch Austrodoris kerguelenensis.

The nudibranch Austrodoris kerguelenensis is distributed widely around the Antarctic coast and continental shelves. Earlier collections from McMurdo Sound and the Weddell Sea shelf have afforded a suite of diterpene glyceride esters, a compound class implicated as a chemical defense in nudibranchs. The present chemical investigation of A. kerguelenensis collected near Palmer Station on the Western Antarctic Peninsula has revealed additional examples, palmadorins A-C (1-3), as the first three members of a new series of clerodane diterpenes. In this paper we describe their isolation, structure elucidation, and stereochemical analysis using a combination of one- and two-dimensional NMR spectroscopy and wet chemical methods.

Sources of primary production to Arctic bivalves identified using amino acid stable carbon isotope fingerprinting.

Benthic invertebrates are a crucial trophic link in Arctic marine food webs. However, estimates of the contribution of different primary production sources sustaining these organisms are not well characterised. We measured the stable carbon isotope values (δ13C) of essential amino acids (EAAs) in muscle tissue from two common bivalve genera (Macoma spp. and Astarte spp.) collected in Hanna Shoal in the northeastern Chukchi Sea. Mixing models comparing the δ13CEAA fingerprints of the bivalves to a suite of primary production endmembers revealed relatively high contributions of EAAs from phytoplankton and bacteria in both species. We also examined whether δ13CEAA fingerprints could be produced from the EAAs preserved in bivalve shells, which could allow primary production sources to be estimated from ancient bivalve shells. The δ13CEAA fingerprints from a suite of paired modern bivalve shells and muscle from Macoma calcarea from across the Chukchi Sea revealed a correspondence between the estimates of the dominant primary production source of EAAs derived from analyses of these two tissue types. Our findings indicate that δ13CEAA fingerprinting of marine bivalves can be used to examine dominant organic matter sources in the Arctic marine benthos in recent years as well as in deeper time.

Sea ice meiofauna distribution on local to pan-Arctic scales.

Arctic sea ice provides microhabitats for biota that inhabit the liquid-filled network of brine channels and the ice-water interface. We used meta-analysis of 23 published and unpublished datasets comprising 721 ice cores to synthesize the variability in composition and abundance of sea ice meiofauna at spatial scales ranging from within a single ice core to pan-Arctic and seasonal scales. Two-thirds of meiofauna individuals occurred in the bottom 10 cm of the ice. Locally, replicate cores taken within meters of each other were broadly similar in meiofauna composition and abundance, while those a few km apart varied more; 75% of variation was explained by station. At the regional scale (Bering Sea first-year ice), meiofauna abundance varied over two orders of magnitude. At the pan-Arctic scale, the same phyla were found across the region, with taxa that have resting stages or tolerance to extreme conditions (e.g., nematodes and rotifers) dominating abundances. Meroplankton, however, was restricted to nearshore locations and landfast sea ice. Light availability, ice thickness, and distance from land were significant predictor variables for community composition on different scales. On a seasonal scale, abundances varied broadly for all taxa and in relation to the annual ice algal bloom cycle in both landfast and pack ice. Documentation of ice biota composition, abundance, and natural variability is critical for evaluating responses to decline in Arctic sea ice. Consistent methodology and protocols must be established for comparability of meiofauna monitoring across the Arctic. We recommend to (1) increase taxonomic resolution of sea ice meiofauna, (2) focus sampling on times of peak abundance when seasonal sampling is impossible, (3) include the bottom 30 cm of ice cores rather than only bottom 10 cm, (4) preserve specimens for molecular analysis to improve taxonomic resolution, and (5) formulate a trait-based framework that relates to ecosystem functioning.

Environmental drivers of the Canadian Arctic megabenthic communities.

Environmental gradients and their influence on benthic community structure vary over different spatial scales; yet, few studies in the Arctic have attempted to study the influence of environmental gradients of differing spatial scales on megabenthic communities across continental-scales. The current project studied for the first time how megabenthic community structure is related to several environmental factors over 2000 km of the Canadian Arctic, from the Beaufort Sea to northern Baffin Bay. Faunal trawl samples were collected between 2007 and 2011 at 78 stations from 30 to 1000 m depth and patterns in biomass, density, richness, diversity, and taxonomic composition were examined in relation to indirect/spatial gradients (e.g., depth), direct gradients (e.g., bottom oceanographic variables), and resource gradients (e.g., food supply proxies). Six benthic community types were defined based on their biomass-based taxonomic composition. Their distribution was significantly, but moderately, associated with large-scale (100-1000 km) environmental gradients defined by depth, physical water properties (e.g., bottom salinity), and meso-scale (10-100 km) environmental gradients defined by substrate type (hard vs. soft) and sediment organic carbon content. We did not observe a strong decline of bulk biomass, density and richness with depth or a strong increase of those community characteristics with food supply proxies, contrary to our hypothesis. We discuss how local- to meso-scale environmental conditions, such as bottom current regimes and polynyas, sustain biomass-rich communities at specific locations in oligotrophic and in deep regions of the Canadian Arctic. This study demonstrates the value of considering the scales of variability of environmental gradients when interpreting their relevance in structuring of communities.

Large-scale spatial distribution patterns of gastropod assemblages in rocky shores.

Gastropod assemblages from nearshore rocky habitats were studied over large spatial scales to (1) describe broad-scale patterns in assemblage composition, including patterns by feeding modes, (2) identify latitudinal pattern of biodiversity, i.e., richness and abundance of gastropods and/or regional hotspots, and (3) identify potential environmental and anthropogenic drivers of these assemblages. Gastropods were sampled from 45 sites distributed within 12 Large Marine Ecosystem regions (LME) following the NaGISA (Natural Geography in Shore Areas) standard protocol (www.nagisa.coml.org). A total of 393 gastropod taxa from 87 families were collected. Eight of these families (9.2%) appeared in four or more different LMEs. Among these, the Littorinidae was the most widely distributed (8 LMEs) followed by the Trochidae and the Columbellidae (6 LMEs). In all regions, assemblages were dominated by few species, the most diverse and abundant of which were herbivores. No latitudinal gradients were evident in relation to species richness or densities among sampling sites. Highest diversity was found in the Mediterranean and in the Gulf of Alaska, while highest densities were found at different latitudes and represented by few species within one genus (e.g. Afrolittorina in the Agulhas Current, Littorina in the Scotian Shelf, and Lacuna in the Gulf of Alaska). No significant correlation was found between species composition and environmental variables (r≤0.355, p>0.05). Contributing variables to this low correlation included invasive species, inorganic pollution, SST anomalies, and chlorophyll-a anomalies. Despite data limitations in this study which restrict conclusions in a global context, this work represents the first effort to sample gastropod biodiversity on rocky shores using a standardized protocol across a wide scale. Our results will generate more work to build global databases allowing for large-scale diversity comparisons of rocky intertidal assemblages.

Aspects of benthic decapod diversity and distribution from rocky nearshore habitat at geographically widely dispersed sites.

Relationships of diversity, distribution and abundance of benthic decapods in intertidal and shallow subtidal waters to 10 m depth are explored based on data obtained using a standardized protocol of globally-distributed samples. Results indicate that decapod species richness overall is low within the nearshore, typically ranging from one to six taxa per site (mean = 4.5). Regionally the Gulf of Alaska decapod crustacean community structure was distinguishable by depth, multivariate analysis indicating increasing change with depth, where assemblages of the high and mid tide, low tide and 1 m, and 5 and 10 m strata formed three distinct groups. Univariate analysis showed species richness increasing from the high intertidal zone to 1 m subtidally, with distinct depth preferences among the 23 species. A similar depth trend but with peak richness at 5 m was observed when all global data were combined. Analysis of latitudinal trends, confined by data limitations, was equivocal on a global scale. While significant latitudinal differences existed in community structure among ecoregions, a semi-linear trend in changing community structure from the Arctic to lower latitudes did not hold when including tropical results. Among boreal regions the Canadian Atlantic was relatively species poor compared to the Gulf of Alaska, whereas the Caribbean and Sea of Japan appeared to be species hot spots. While species poor, samples from the Canadian Atlantic were the most diverse at the higher infraordinal level. Linking 11 environmental variables available for all sites to the best fit family-based biotic pattern showed a significant relationship, with the single best explanatory variable being the level of organic pollution and the best combination overall being organic pollution and primary productivity. While data limitations restrict conclusions in a global context, results are seen as a first-cut contribution useful in generating discussion and more in-depth work in the still poorly understood field of biodiversity distribution.

Spatial relationships between polychaete assemblages and environmental variables over broad geographical scales.

This study examined spatial relationships between rocky shore polychaete assemblages and environmental variables over broad geographical scales, using a database compiled within the Census of Marine Life NaGISA (Natural Geography In Shore Areas) research program. The database consisted of abundance measures of polychaetes classified at the genus and family levels for 74 and 93 sites, respectively, from nine geographic regions. We tested the general hypothesis that the set of environmental variables emerging as potentially important drivers of variation in polychaete assemblages depend on the spatial scale considered. Through Moran's eigenvector maps we indentified three submodels reflecting spatial relationships among sampling sites at intercontinental (>10,000 km), continental (1000-5000 km) and regional (20-500 km) scales. Using redundancy analysis we found that most environmental variables contributed to explain a large and significant proportion of variation of the intercontinental submodel both for genera and families (54% and 53%, respectively). A subset of these variables, organic pollution, inorganic pollution, primary productivity and nutrient contamination was also significantly related to spatial variation at the continental scale, explaining 25% and 32% of the variance at the genus and family levels, respectively. These variables should therefore be preferably considered when forecasting large-scale spatial patterns of polychaete assemblages in relation to ongoing or predicted changes in environmental conditions. None of the variables considered in this study were significantly related to the regional submodel.

Patterns of spatial variation of assemblages associated with intertidal rocky shores: a global perspective.

Assemblages associated with intertidal rocky shores were examined for large scale distribution patterns with specific emphasis on identifying latitudinal trends of species richness and taxonomic distinctiveness. Seventy-two sites distributed around the globe were evaluated following the standardized sampling protocol of the Census of Marine Life NaGISA project (www.nagisa.coml.org). There were no clear patterns of standardized estimators of species richness along latitudinal gradients or among Large Marine Ecosystems (LMEs); however, a strong latitudinal gradient in taxonomic composition (i.e., proportion of different taxonomic groups in a given sample) was observed. Environmental variables related to natural influences were strongly related to the distribution patterns of the assemblages on the LME scale, particularly photoperiod, sea surface temperature (SST) and rainfall. In contrast, no environmental variables directly associated with human influences (with the exception of the inorganic pollution index) were related to assemblage patterns among LMEs. Correlations of the natural assemblages with either latitudinal gradients or environmental variables were equally strong suggesting that neither neutral models nor models based solely on environmental variables sufficiently explain spatial variation of these assemblages at a global scale. Despite the data shortcomings in this study (e.g., unbalanced sample distribution), we show the importance of generating biological global databases for the use in large-scale diversity comparisons of rocky intertidal assemblages to stimulate continued sampling and analyses.

Large-scale spatial distribution patterns of echinoderms in nearshore rocky habitats.

This study examined echinoderm assemblages from nearshore rocky habitats for large-scale distribution patterns with specific emphasis on identifying latitudinal trends and large regional hotspots. Echinoderms were sampled from 76 globally-distributed sites within 12 ecoregions, following the standardized sampling protocol of the Census of Marine Life NaGISA project (www.nagisa.coml.org). Sample-based species richness was overall low (<1-5 species per site), with a total of 32 asteroid, 18 echinoid, 21 ophiuroid, and 15 holothuroid species. Abundance and species richness in intertidal assemblages sampled with visual methods (organisms >2 cm in 1 m(2) quadrats) was highest in the Caribbean ecoregions and echinoids dominated these assemblages with an average of 5 ind m(-2). In contrast, intertidal echinoderm assemblages collected from clearings of 0.0625 m(2) quadrats had the highest abundance and richness in the Northeast Pacific ecoregions where asteroids and holothurians dominated with an average of 14 ind 0.0625 m(-2). Distinct latitudinal trends existed for abundance and richness in intertidal assemblages with declines from peaks at high northern latitudes. No latitudinal trends were found for subtidal echinoderm assemblages with either sampling technique. Latitudinal gradients appear to be superseded by regional diversity hotspots. In these hotspots echinoderm assemblages may be driven by local and regional processes, such as overall productivity and evolutionary history. We also tested a set of 14 environmental variables (six natural and eight anthropogenic) as potential drivers of echinoderm assemblages by ecoregions. The natural variables of salinity, sea-surface temperature, chlorophyll a, and primary productivity were strongly correlated with echinoderm assemblages; the anthropogenic variables of inorganic pollution and nutrient contamination also contributed to correlations. Our results indicate that nearshore echinoderm assemblages appear to be shaped by a network of environmental and ecological processes, and by the differing responses of various echinoderm taxa, making generalizations about the patterns of nearshore rocky habitat echinoderm assemblages difficult.

Current patterns of macroalgal diversity and biomass in northern hemisphere rocky shores.

Latitudinal gradients in species abundance and diversity have been postulated for nearshore taxa but few analyses have been done over sufficiently broad geographic scales incorporating various nearshore depth strata to empirically test these gradients. Typically, gradients are based on literature reviews and species lists and have focused on alpha diversity across the entire nearshore zone. No studies have used a standardized protocol in the field to examine species density among sites across a large spatial scale while also focusing on particular depth strata. The present research used field collected samples in the northern hemisphere to explore the relationships between macroalgal species density and biomass along intertidal heights and subtidal depths and latitude. Results indicated no overall correlations between either estimates of species density or biomass with latitude, although the highest numbers of both were found at mid-latitudes. However, when strata were examined separately, significant positive correlations were found for both species numbers and biomass at particular strata, namely the intertidal ones. While the data presented in this paper have some limitations, we show that latitudinal macroalgal trends in species density and biomass do exist for some strata in the northern hemisphere with more taxa and biomass at higher latitudes.

Global patterns and predictions of seafloor biomass using random forests.

A comprehensive seafloor biomass and abundance database has been constructed from 24 oceanographic institutions worldwide within the Census of Marine Life (CoML) field projects. The machine-learning algorithm, Random Forests, was employed to model and predict seafloor standing stocks from surface primary production, water-column integrated and export particulate organic matter (POM), seafloor relief, and bottom water properties. The predictive models explain 63% to 88% of stock variance among the major size groups. Individual and composite maps of predicted global seafloor biomass and abundance are generated for bacteria, meiofauna, macrofauna, and megafauna (invertebrates and fishes). Patterns of benthic standing stocks were positive functions of surface primary production and delivery of the particulate organic carbon (POC) flux to the seafloor. At a regional scale, the census maps illustrate that integrated biomass is highest at the poles, on continental margins associated with coastal upwelling and with broad zones associated with equatorial divergence. Lowest values are consistently encountered on the central abyssal plains of major ocean basins The shift of biomass dominance groups with depth is shown to be affected by the decrease in average body size rather than abundance, presumably due to decrease in quantity and quality of food supply. This biomass census and associated maps are vital components of mechanistic deep-sea food web models and global carbon cycling, and as such provide fundamental information that can be incorporated into evidence-based management.

Bioassay-guided fractionation of antifouling compounds using computer-assisted motion analysis of brown algal spore swimming.

Antifouling extracts from the sea stars Astropecten articulatus and Luidia clathrata and from the brittle star Astrocyclus caecilia were fractionated by solid phase extraction and high performance liquid chromatography. Bioactive fractions were identified with the use of computer-assisted motion analysis-based bioassays utilising previously described Hincksia irregularis spore swimming behaviour parameters. Quantified parameters of spore movement were rate of change of direction (RCD) and speed (SPEE). The methods used initially required only 10 microg equivalent amounts of total crude extract and each resultant resolving step (normalised to 1 mg ml(-1) of crude, unfractionated extract) required far less material. Statistical analyses of RCD and ratios of RCD:SPEE values in experiments comparing swimming in the presence of extract fractions to controls revealed that both parameters were useful individually and in combination for efficiently following compound bioactivity throughout the fractionation procedure. This technique was also able to detect synergistic or additive interactions between compounds.

Ainigmaptilones, sesquiterpenes from the Antarctic gorgonian coral Ainigmaptilon antarcticus.

Antarctic octocorals from the Weddell Sea of Western Antarctica appear to select either physical or chemical means of defense. Fractionation of the bioactive extract from the chemically defended Antarctic gorgonian coral Ainigmaptilon antarcticus yielded two sesquiterpenes, ainigmaptilones A (1) and B (2). Ainigmaptilone A has broad spectrum bioactivity toward sympatric predatory and fouling organisms, including antibiotic activity.

Individual and coupled effects of echinoderm extracts and surface hydrophobicity on spore settlement and germination in the brown alga Hincksia irregularis.

Marine substrata possess cues that influence the behavior of fouling organisms. Initial adhesion of fouling algal zoospores to surfaces is also theorized to depend primarily upon interactions between substrata and spore cell bodies and flagellar membranes. In an effort to identify cues and surface characteristics that influence spore settlement and early development, the effects of bioactive echinoderm extracts, surface charge, and surface hydrophobicity were examined individually and in tandem on zoospore settlement and germination in Hincksia irregularis. Experiments utilizing 96-well plastic culture plates confirmed that spore settlement and germination were significantly affected by surface charge and hydrophobicity as well as by echinoderm metabolites, both individually and in tandem. Spore settlement rates in the dark over 30 min were > 400% higher on hydrophobic surfaces than on positively and negatively charged surfaces. Spore germling numbers were > 300% higher on hydrophobic surfaces than on positively and negatively charged surfaces when spores were allowed to settle in the light for 30 min and the settled spores allowed to subsequently germinate for 24 h. Spore germling numbers were consistently > 25% higher on hydrophobic surfaces than on positively and negatively charged surfaces when equal numbers of spores were allowed to completely settle in the light and subsequently germinate for 24 h. H. irregularis germ tube lengths were also significantly longer on positively charged plates than on negatively charged plates. All echinoderm extracts tested had significant effects on germination and settlement at levels below those of estimated ecological concentrations. Short-term (30 min) exposure and subsequent germination experiments indicated that higher concentrations of extracts had rapid toxic effects on algal spores. Synchronous effects of echinoderm extracts and plate charge upon spore settlement varied considerably and did not show a strong dose response relationship. Long-term (24 h) exposure of spores to echinoderm extracts had dosage dependent effects on germination and spore survival. The results of this study indicate that H. irregularis spores possess the capacity for complex responses to their environment, utilizing combined cues of surface charge, surface energy and biochemistry to determine where to settle and germinate. These responses may aid spores in the detection of suitable substrata and conditions for settlement in the marine environment.