MARINE ECOLOGY PROGRESS SERIES

Carbon limitation in response to nutrient loading in an eelgrass mesocosm: influence of water residence time
Kaldy JE, Brown CA and Pacella SR
Altered primary productivity associated with eutrophication impacts not only ecosystem structure but also the biogeochemical cycling of oxygen and carbon. We conducted laboratory experiments to empirically determine how residence time (1, 3, 10 d) influences eutrophication responses in a simplified Pacific Northwest -green macroalgal community. We expected long-residence time (RT) systems to exhibit eutrophication impairments. Instead, we observed an accumulation of nutrients at all RTs and a shift in the dissolved inorganic carbon speciation away from CO with unexpected consequences for eel grass plant condition, including shoot mortality. Most metrics responded more strongly to temperature treatments than to RT treatments. No dramatic shifts in the relative abundance of and green macro algae were detected. shoot density proliferated in cool temperatures (12°C) with a modest decline at 20°C. Eelgrass loss was associated with high total scale pH (pH) and CO concentrations of <10 μmol kg CO, but not with high nutrients. δC values support the hypo thesis that carbon availability was greater at short RT. Further, very low leaf sugar concentrations are consistent with extreme photosynthetic CO limitation. We suggest that the effects of extremely low environ mental car bon concentrations (CO) and increased respiration at warm temperatures (20°C) and other physiological processes can lead to internal carbon limitation and shoot mortality. Eutrophication responses to nutrient loading are more nuanced than just light limitation of eelgrass and require additional research on the interaction of the biogeochemical environment and plant physiology to better understand estuarine ecosystem disruption.
Oyster Calcifying Fluid Harbors Persistent and Dynamic Autochthonous Bacterial Populations That May Aid in Shell Formation
Sakowski EG, Wommack KE and Polson SW
The eastern oyster () is a keystone species in estuarine environments but faces threats to shell formation associated with warming temperatures and acidification. Extrapallial fluid (EF), which is responsible for shell formation, harbors diverse and abundant microbial communities. Commensal microbial communities are vital to host health and fitness, yet long-term studies investigating temporal responses of the EF microbiome and its function in oyster fitness are lacking. In this study, bacterial communities of oyster EF and the water column were characterized monthly from October 2010 to September 2011. We investigated the selection, composition, and dynamics of resident and transient community members, evaluated the impact of temperature on EF microbial communities, and examined the functional role of the EF microbiome. Oyster EF communities were significantly different from the water column and were enriched for several taxa, including the Deltaproteobacteria, Epsilonproteobacteria, and Gammaproteobacteria. Overall, 94 resident members were identified in oyster EF. These members were persistent and abundant, comprising on average 33% of EF communities. Resident EF communities formed high-temperature and low-temperature groups and were more abundant overall at colder temperatures. Oyster EF resident communities were predicted to be enriched for dissimilatory nitrate reduction, nitrogen fixation, nitrification, and sulfite reductase genes. Sulfate and nitrate reduction may have a synergistic effect on calcium carbonate precipitation and indirectly aid in shell formation. Therefore, the potential role of the oyster EF microbiome in shell formation warrants further investigation as oysters and other shellfish face the future impacts of ocean warming and acidification.
Tracking the marine migration routes of South Pacific silver eels
Chang YK, Dall'Olmo G and Schabetsberger R
Three catadromous Pacific eels (2 , 1 ) from the Archipelago of Vanuatu were tagged with pop-up satellite archival transmitters and their migration tracks towards their presumed spawning area approximately 870 km northeast of the point of release were reconstructed in order to evaluate their movements in relation to oceanographic conditions. We used the timing of diel vertical migrations to derive the eels' positions. Two exhibited steep-angled turns resulting in a zig-zag migration path along the east-west axis, while one took a relatively straight course towards the presumed spawning area. They migrated with a speed over ground of 21-23 km day. In this region, the eastward flow of the South Equatorial Counter Current (SECC, ∼ 5-10°S) separates the westward flowing South Equatorial Current (SEC; ∼0-5°S and 10-18°S) into two branches. During shallower nighttime migration depths around 150 m eels crossed a variable flow field through the southern branch of the westward SEC with westward propagating mesoscale eddies and the eastward SECC, but stayed south of the stronger northern branch of SEC possibly increasing retention time of larvae within this area. The eels headed towards a tongue of high-salinity Subtropical Underwater (STUW) that may have provided cues for orientation. The eels did not move beyond a salinity front of 35.9-36.0 at a depth of 100-200 m, which may have provided cues for orientation towards the spawning area. These 3 tracks may represent the movements of mature silver eels all the way to where they spawn.
Elevated CO and Hypoxia Alter the Acid-Base Homeostasis of Developing Sheepshead Minnows,
Enzor LA, Hankins C, Hamilton-Frazier M, Moso E, Raimondo S and Barron MG
Low dissolved oxygen and increased acidification are two environmental variables that concomitantly change in an estuarine environment, both of which are exacerbated by nutrient pollution and subsequent eutrophication. To better understand how estuarine residents compensate for daily fluctuations in these environmental variables, the interactive effects of acidification and hypoxia were assessed in developing sheepshead minnows () using a 2 by 2 factorial design over a 42-day exposure. Embryos were exposed to either acidic (partial pressure of CO, CO, ~2000 μatm), hypoxic (reduced dissolved oxygen, ~2 mg l), or combined acidic and hypoxic conditions and monitored for development, hatch rate, and survival. Changes in oxygen consumption, anaerobic metabolism, oxidative stress, and acid-base balance were evaluated at three life stages (embryo, larval, and juvenile fish) to discern if and how fish compensate for these stressors during development. The combination of acidification and hypoxia delayed hatching in embryos and significantly decreased oxygen consumption (p<0.001) in all three life-stages. Neither acidification, hypoxia, nor the combination of the stressors impacted the anaerobic metabolism or oxidative stress of juvenile fish, but acid-base equilibrium was disrupted by all three treatments in larval fish. Elevated carbonic anhydrase activity was observed in the multi-stress treatment in embryos and larval fish, but not in juvenile fish. These results show that developing sheepshead minnows can re-establish cellular homeostasis in compensating to acidified and hypoxic waters.
Effects of coastal acidification on North Atlantic bivalves: interpreting laboratory responses in the context of populations
Grear JS, O'Leary CA, Nye JA, Tettelbach ST and Gobler CJ
Experimental exposure of early life stage bivalves has documented negative effects of elevated CO on survival and growth, but the population consequences of these effects are unknown. Following standard practices from population viability analysis and wildlife risk assessment, we substituted laboratory-derived stress-response relationships into baseline population models of and . The models were constructed using inverse demographic analyses with time series of size-structured field data in NY, USA, whereas the stress-response relationships were developed using data from a series of previously published laboratory studies. We used stochastic projection methods and diffusion approximations of extinction probability to estimate cumulative risk of 50% population decline during ten-year population projections at 1, 1.5 and 2 times ambient CO levels. Although the population exhibited higher growth in the field data (12% per year) than the declining population (-8% per year), cumulative risk was high for in the first ten years due to high variance in the stochastic growth rate estimate (log λ = -0.02, σ = 0.24). This ten-year cumulative risk increased from 69% to 94% and >99% at 1.5 and 2 times ambient scenarios. For (log λ = -0.09, σ = 0.01), ten-year risk was 81%, 96% and >99% at 1, 1.5 and 2 times ambient CO, respectively. These estimates of risk could be improved with detailed consideration of harvest effects, disease, restocking, compensatory responses, other ecological complexities, and the nature of interactions between these and other effects that are beyond the scope of available data. However, results clearly indicate that early life stage responses to plausible levels of CO enrichment have the potential to cause significant increases in risk to these marine bivalve populations.
Effect of Green Macroalgal Blooms on the Behavior, Growth, and Survival of Cockles () in Pacific NW Estuaries
Lewis NS and DeWitt TH
Nutrient over-enrichment can produce adverse ecological effects within coastal ecosystems and negatively impact the production of ecosystem goods and services. In small estuaries of the U.S. Pacific Northwest, seasonal blooms of green macroalgae (GMA; Family Ulvaceae) are primarily associated with natural nutrient input, rather than anthropogenic sources. This provided us a unique opportunity to investigate the effects of naturally-stimulated macroalgal blooms on intertidal bivalves. (heart cockles) are an important species for shellfisheries in the region. In summer population surveys, we found that cockles emerged from the sediment with greater frequency as GMA biomass increased. Experimental manipulation of GMA biomass in the field showed that GMA elicited emergence, evoked above-ground lateral movement, inhibited shell growth, and increased mortality (by 34.0 ± 15.2%) in cockles. Laboratory experiments revealed that the interaction of a weighted barrier at the sediment surface and GMA presence elicited rapid emergence among cockles. Risk assessment of the emergence response in cockles showed that the emergent population experienced 11.0 ± 8.0% mortality due to gull predation, while laboratory exposure to elevated temperatures (≥34 °C) slowed valve-closure, inhibited reburial, and increased mortality, which could have translated to 7.1 ± 1.5% mortality. We found that cockles avoided mortality due to burial below GMA mats by emerging from the sediment, but that behavior consequently put them at risk of mortality due to heat stress or gull predation. Regardless of nutrient source, our research showed that GMA blooms pose a threat to the survival of intertidal bivalves.
The hydrographic features of anguillid spawning areas: potential signposts for migrating eels
Schabetsberger R, Miller MJ, Dall'Olmo G, Kaiser R, Økland F, Watanabe S, Aarestrup K and Tsukamoto K
Catadromous anguillid eels (Genus ) migrate from their freshwater or estuarine habitats to marine spawning areas. Evidence from satellite tagging studies indicates that tropical and temperate eel species exhibit pronounced diel vertical migrations between 150 to 300 m nighttime depths to 600 to 800 m during the day. Collections of eggs and larvae of Japanese eels ( ) show they may spawn at these upper nighttime migration depths. How anguillid eels navigate through the ocean and find their spawning areas remains unknown, so this study describes the salinity, temperature and geostrophic currents between 0 and 800 m depths within two confirmed and three hypothetical anguillid spawning areas during likely spawning seasons. Within all four ocean gyres many eels would encounter subducted 'Subtropical Underwater' during their nighttime ascents that could provide odor plumes as signposts. Four spawning areas are located near the western margins of where subducted water masses form cores of elevated salinities (~35.0 to 36.8) around 150 m depths, while one is found near the center of subduction. Low salinity surface waters and fronts are present in some of the areas above the high-salinity cores. Spawning may occur at temperatures between 16 to 24°C where the thermocline locally deepens. At spawning depths, weak westward currents (~0 to 0.1 m s) prevail, and eastward surface countercurrents are present. Anguillid eels possess acute sensory capabilities to detect these hydrographic features as potential signposts guiding them to where they spawn.
Effect of marine protected areas (MPAs) on consumer diet: MPA fish feed higher in the food chain
Dell C, Montoya J and Hay M
Marine Protected Areas (MPAs) are often established to mitigate the effects of overfishing and other human disturbances. In Fiji these are locally managed and, where enforced, have significantly higher coral cover, higher fish biomass, and lower seaweed cover than in the adjacent, unprotected reefs (non-MPAs). We investigated how the isotopic signatures of a common, mid-level consumer, , differed among three small (0.5- 0.8km) MPAs versus adjacent, unprotected reefs. Isotopic ratios suggested that the fish in the MPAs fed higher in the food chain than those in the adjacent non-MPAs, despite being slightly smaller in size. Calculations using a brown alga as representative of the basal level of the food chain estimate this difference to be about half a trophic level. Thus, the isotopic ratio of a mid-level consumer can be noticeably altered over scales of only a few hundred meters. This may result from more complete food webs and hence greater prey choice and availability in the MPAs and implies that MPAs affect not only species' abundance and diversity, but also diet composition and trophic biology of member individuals. Our findings suggest exhibits considerable site fidelity in its feeding biology and thus provides a localized isotopic signal of its reef of residence. If the isotopic signal of this mid-level carnivore is reflective of the composition of the food web beneath it, the signal might provide an easily obtained indication of reef conditions in that area.
Size matters at deep-sea hydrothermal vents: different diversity and habitat fidelity patterns of meio- and macrofauna
Gollner S, Govenar B, Fisher CR and Bright M
Species with markedly different sizes interact when sharing the same habitat. Unravelling mechanisms that control diversity thus requires consideration of a range of size classes. We compared patterns of diversity and community structure for meio- and macrofaunal communities sampled along a gradient of environmental stress at deep-sea hydrothermal vents on the East Pacific Rise (9° 50' N) and neighboring basalt habitats. Both meio- and macrofaunal species richnesses were lowest in the high-stress vent habitat, but macrofaunal richness was highest among intermediate-stress vent habitats. Meiofaunal species richness was negatively correlated with stress, and highest on the basalt. In these deep-sea basalt habitats surrounding hydrothermal vents, meiofaunal species richness was consistently higher than that of macrofauna. Consideration of the physiological capabilities and life history traits of different-sized animals suggests that different patterns of diversity may be caused by different capabilities to deal with environmental stress in the 2 size classes. In contrast to meiofauna, adaptations of macrofauna may have evolved to allow them to maintain their physiological homeostasis in a variety of hydrothermal vent habitats and exploit this food-rich deep-sea environment in high abundances. The habitat fidelity patterns also differed: macrofaunal species occurred primarily at vents and were generally restricted to this habitat, but meiofaunal species were distributed more evenly across proximate and distant basalt habitats and were thus not restricted to vent habitats. Over evolutionary time scales these contrasting patterns are likely driven by distinct reproduction strategies and food demands inherent to fauna of different sizes.
Population characteristics of a recovering US Virgin Islands red hind spawning aggregation following protection
Nemeth RS
Many species of groupers form spawning aggregations, dramatic events where 100s to 1000s of individuals gather annually at specific locations for reproduction. Spawning aggregations are often targeted by local fishermen, making them extremely vulnerable to over fishing. The Red Hind Bank Marine Conservation District located in St. Thomas, United States Virgin Islands, was closed seasonally in 1990 and closed permanently in 1999 to protect an important red hind Epinephelus guttatus spawning site. This study provides some of the first information on the population response of a spawning aggregation located within a marine protected area. Tag-and-release fishing and fish transects were used to evaluate population characteristics and habitat utilization patterns of a red hind spawning aggregation between 1999 and 2004. Compared with studies conducted before the permanent closure, the average size of red hind increased mostly during the seasonal closure period (10 cm over 12 yr), but the maximum total length of male red hind increased by nearly 7 cm following permanent closure. Average density and biomass of spawning red hind increased by over 60% following permanent closure whereas maximum spawning density more than doubled. Information from tag returns indicated that red hind departed the protected area following spawning and migrated 6 to 33 km to a ca. 500 km(2) area. Protection of the spawning aggregation site may have also contributed to an overall increase in the size of red hind caught in the commercial fishery, thus increasing the value of the grouper fishery for local fishermen.
Distribution of (Dinophyceae) cysts in Greenland and Iceland, with an emphasis on viability and growth in the Arctic
Richlen ML, Zielinski O, Holinde L, Tillmann U, Cembella A, Lyu Y and Anderson DM
The bloom-forming dinoflagellate has been extensively studied due its toxin-producing capabilities and consequent impacts to human health and economies. This study investigated the prevalence of resting cysts of in western Greenland and Iceland to assess the historical presence and magnitude of bloom populations in the region, and to characterize environmental conditions during summer, when bloom development may occur. Analysis of sediments collected from these locations showed that cysts were present at low to moderate densities in most areas surveyed, with highest densities observed in western Iceland. Additionally, laboratory experiments were conducted on clonal cultures established from isolated cysts or vegetative cells from Greenland, Iceland, and the Chukchi Sea (near Alaska) to examine the effects of photoperiod interval and irradiance levels on growth. Growth rates in response to the experimental treatments varied among isolates, but were generally highest under conditions that included both the shortest photoperiod interval (16h:8h light:dark) and higher irradiance levels (~146-366 μmol photons m s), followed by growth under an extended photoperiod interval and low irradiance level (~37 μmol photons m s). Based on field and laboratory data, we hypothesize that blooms in Greenland are primarily derived from advected populations, as low bottom temperatures and limited light availability would likely preclude in situ bloom development. In contrast, the bays and fjords in Iceland may provide more favorable habitat for germling cell survival and growth, and therefore may support indigenous, self-seeding blooms.
Extensive geographic and ontogenetic variation characterizes the trophic ecology of a temperate reef fish on southern California (USA) rocky reefs
Hamilton SL, Caselle JE, Lantz CA, Egloff TL, Kondo E, Newsome SD, Loke-Smith K, Pondella DJ, Young KA and Lowe CG
Interactions between predator and prey act to shape the structure of ecological communities, and these interactions can differ across space. California sheephead are common predators of benthic invertebrates in kelp beds and rocky reefs in southern California, USA. Through gut content and stable isotope (δC and †N) analyses, we investigated geographic and ontogenetic variation in trophic ecology across 9 populations located at island and mainland sites throughout southern California. We found extensive geographic variation in California sheephead diet composition over small spatial scales. Populations differed in the proportion of sessile filter/suspension feeders or mobile invertebrates in the diet. Spatial variation in diet was highly correlated with other life history and demographic traits (e.g. growth, survivorship, reproductive condition, and energy storage), in addition to proxies of prey availability from community surveys. Multivariate descriptions of the diet from gut contents roughly agreed with the spatial groupings of sites based on stable isotope analysis of both California sheephead and their prey. Ontogenetic changes in diet occurred consistently across populations, despite spatial differences in size structure. As California sheephead increase in size, diets shift from small filter feeders, like bivalves, to larger mobile invertebrates, such as sea urchins. Our results indicate that locations with large California sheephead present, such as many marine reserves, may experience increased predation pressure on sea urchins, which could ultimately affect kelp persistence.
Escaping paradise: Larval export from Hawaii in an Indo-Pacific reef fish, the Yellow Tang ()
Eble JA, Toonen RJ, Sorenson L, Basch LV, Papastamatiou YP and Bowen BW
The depauperate marine ecosystems of the Hawaiian Archipelago share a high proportion of species with the southern and western Pacific, indicating historical and/or ongoing connections across the large oceanic expanse separating Hawaii from its nearest neighbors. The rate and direction of these interactions are, however, unknown. While previous biogeographic studies have consistently described Hawaii as a diversity sink, prevailing currents likely offer opportunities for larval export. To assess interactions between the remote reefs of the Hawaiian Archipelago and the species rich communities of the Central and West Pacific, we surveyed 14 nuclear microsatellite loci (nDNA; = 857) and a 614 bp segment of mitochondrial cytochrome (mtDNA; = 654) in the Yellow Tang (). Concordant frequency shifts in both nDNA and mtDNA reveal significant population differentiation among three West Pacific sites and Hawaii (nDNA ' = 0.116, mtDNA ϕ = 0.098, < 0.001). SAMOVA analyses of microsatellite data additionally indicate fine scale differentiation within the 2600 km Hawaiian Archipelago (' = 0.026; < 0.001), with implications for management of this heavily-exploited aquarium fish. Mismatch analyses indicate the oldest contemporary populations are in the Hawaiian Archipelago (circa 318,000 y), with younger populations in the West Pacific (91,000 - 175,000 y). Estimates of Yellow Tang historical demography contradict expectations of Hawaii as a population sink, and instead indicate asymmetrical gene flow, with Hawaii exporting rather than importing Yellow Tang larvae.
Lack of adult novel northern lineages of invasive green crab along much of the northern US Atlantic coast
Williams LM, Nivison CL, Ambrose WG, Dobbin R and Locke WL
Introduced over 200 yr ago to the east coast of North America, now ranges from New York to Newfoundland. In the 1980s, a secondary invasion of European lineages, termed northern haplotypes, occurred in Nova Scotia. Young-of-the-year sampled in 2007 revealed that northern haplotypes were present in low frequencies at several northwestern Atlantic sites as far south as New York; a model predicted an increase in their range and frequency over time. We collected samples in 2013 and 2014 to determine the haplotypes of adult crabs from New York to Nova Scotia. Six haplotypes, encompassing previously identified northern and southern haplotypes, 1 novel southern haplotype, and 1 Scandinavian haplotype, were identified in 275 crabs sampled at 11 sites. Northern haplotypes were only found in Nova Scotia, Beals Island (Maine), and Mount Desert Island (Maine) at a frequency of 60, 8, and 24%, respectively; remaining sites were predominantly composed of a previously identified southern haplotype. Northern haplotypes are limited in adult crabs to Mount Desert Island and north, indicating that the southern haplotype is selectively favored at some point during their life history, recruitment of northern larvae is limited south of Mount Desert Island, or entire year-classes post-2007 were lost. Our results do not support the predictions of an increase in the range and frequency of northern haplotypes, at least among adults, and indicate that a more complete knowledge of factors affecting life stages is necessary to understand the current distribution of haplotypes.
Molecular characterization of community structure and species ecology in a hydrographically complex estuarine system (Puget Sound, Washington, USA)
Hubbard KA, Olson CH and Armbrust EV
Species within the toxic marine diatom genus coexist in coastal and estuarine waters globally and are difficult to distinguish by microscopy. Here, we describe a sensitive, high throughput PCR-based Automated Ribosomal Intergenic Spacer Analysis (ARISA) approach to determine the relative abundance of species within natural communities over space and time. The method was quantitatively validated using simplified mixtures of DNA or ITS1 standards from isolates of and . Relative abundance calculations based on ARISA profiles from these mixtures reflected input ratios, with minor deviations resulting from intraspecific variability. ARISA was used to identify and quantify at least eight species within Puget Sound and the eastern Strait of Juan de Fuca, Washington, USA: and genotypes corresponding to var. and var. were identified by environmental sequencing. The different species were significantly correlated with physical (temperature, salinity), biological (chlorophyll fluorescence, oxygen), and/or chemical (ammonium, nutrient ratios) factors. The ability to determine shifts in the relative abundance of species over spatial and temporal scales relevant to dispersion and selection facilitates dissection of the varied mechanisms driving vertical and horizontal species distribution patterns in hydrographically complex systems.
Trophodynamics and mercury bioaccumulation in reef and open-ocean fishes from The Bahamas with a focus on two teleost predators
Shipley ON, Lee CS, Fisher NS, Burruss G, Frisk MG, Brooks EJ, Zuckerman ZC, Herrmann AD and Madigan DJ
Identifying prey resource pools supporting fish biomass can elucidate trophic pathways of pollutant bioaccumulation. We used multiple chemical tracers (carbon [δC] and nitrogen [δN] stable isotopes and total mercury [THg]) to identify trophic pathways and measure contaminant loading in upper trophic level fishes residing at a reef and open-ocean interface near Eleuthera in the Exuma Sound, The Bahamas. We focused predominantly on the trophic pathways of mercury bioaccumulation in dolphinfish and wahoo , 2 commonly consumed pelagic sportfish in the region. Despite residing within close proximity to productive and extensive coral reefs, both dolphinfish and wahoo relied almost exclusively on open-ocean prey over both short and long temporal durations. A larger isotopic niche of dolphinfish suggested a broader diet and some potential prey differentiation between the 2 species. THg concentrations in dolphinfish (0.2 ± 0.1 ppm) and wahoo (0.3 ± 0.3 ppm) were mostly below recommended guidelines for humans (US Environmental Protection Agency (EPA) = 0.3 ppm, US Food and Drug Administration (FDA)= 1.0 ppm) and were within ranges previously reported for these species. However, high THg concentrations were observed in muscle and liver tissue of commonly consumed reef-associated fishes, identifying a previously unrecognized route of potentially toxic Hg exposure for human consumers on Eleuthera and neighboring islands.
Induced defence to grazing by vertebrate herbivores: uncommon or under-investigated?
Dell C and Hay ME
There are many examples of macroalgae inducing defence in response to small invertebrate herbivores like amphipods, isopods, and gastropods but few cases of induction in response to vertebrate macrograzers like herbivorous fishes. This may be because larger grazers rapidly consume large quantities of seaweed before induction can occur, thus selecting for constitutive rather than induced defences. Alternatively, the pattern could occur because induction due to feeding by macrograzers is less commonly investigated. In Fiji, field assays with the brown macroalga demonstrated that thalli growing in marine protected areas (MPAs) with abundant herbivorous fishes were significantly less palatable than those growing in adjacent fished areas (non-MPAs) with few herbivorous fishes. This significant preference occurred in 11 of 13 trials over 5 time periods and across 3 pairs of MPAs and spatially associated non-MPAs. This preference was not positively associated with algal nitrogen content or with the toughness of algal fronds. When ramets were taken from the non-MPA and half were partially grazed by fishes while the other half were protected from grazing, new growth from the controls was strongly preferred to new growth from the previously grazed ramets although these fronds originated from the same holdfast. This suggests that upregulates defences (probably chemical) in response to grazing by herbivorous fishes. This is one of the few published examples of induction of macroalgal defence in response to feeding by large, mobile grazers. It is unclear whether induced defences against fishes are rare or just under-investigated.
Intergenerational effects of macroalgae on a reef coral: major declines in larval survival but subtle changes in microbiomes
Beatty DS, Clements CS, Stewart FJ and Hay ME
Tropical reefs are shifting from coral to macroalgal dominance, with macroalgae suppressing coral recovery, potentially via effects on coral microbiomes. Understanding how macroalgae affect corals and their microbiomes requires comparing algae- versus coral-dominated reefs without confounding aspects of time and geography. We compared survival, settlement, and post-settlement survival of larvae, as well as the microbiomes of larvae and adults, of the Pacific coral between an Marine Protected Area (MPA) dominated by corals versus an adjacent fished area dominated by macroalgae. Microbiome composition in adult coral, larval coral, and seawater did not differ between the MPA and fished area. However, microbiomes of adult coral were more variable in the fished area and Vibrionaceae bacteria, including strains most closely related to the pathogen , were significantly enriched, but rare, in adult and larval coral from the fished area. Larvae from the macroalgae-dominated area exhibited higher pre-settlement mortality and reduced settlement compared to those from the coral-dominated area. Juveniles planted into a coral-dominated area survived better than those placed into a fished area dominated by macroalgae. Differential survival depended on whether macroalgae were immediately adjacent to juvenile coral rather than on traits of the areas . Contrary to our expectations, coral microbiomes were relatively uniform at the community level despite dramatic differences in macroalgal cover between the MPA (~2% cover) and fished (~90%) area. Reducing macroalgae may elicit declines in rare but potentially harmful microbes in coral and their larvae, as well as positive intergenerational effects on offspring survival.
Spatial and temporal limits of coral-macroalgal competition: the negative impacts of macroalgal density, proximity, and history of contact
Clements CS, Rasher DB, Hoey AS, Bonito VE and Hay ME
Tropical reefs are commonly transitioning from coral- to macroalgal-dominance, producing abrupt, and often lasting, shifts in community composition and ecosystem function. Although negative effects of macroalgae on corals are well documented, whether such effects vary with spatial scale or the density of macroalgae remains inadequately understood, as does the legacy of their impact on coral growth. Using closely adjacent coral- versus macroalgal-dominated areas, we tested effects of macroalgal competition on the Indo-Pacific corals and . When corals were transplanted to areas of: ) macroalgal-dominance, ) macroalgal-dominance but with nearby macroalgae removed, or ) coral-dominance lacking macroalgae, coral growth was equivalently high in plots without macroalgae and low (62-90% less) in plots with macroalgae, regardless of location. In a separate experiment, we raised corals above the benthos in each area and exposed them to differing densities of the dominant macroalga . Coral survivorship was high (≥ 93% after 3 months) and did not differ among treatments, whereas the growth of both coral species decreased as a function of density. When was removed after two months, there was no legacy effect of macroalgal density on coral growth over the next seven months; however, there was no compensation for previously depressed growth. In sum, macroalgal impacts were density dependent, occurred only if macroalgae were in close contact, and coral growth was resilient to prior macroalgal contact. The temporal and spatial constraints of these interactions suggest that corals may be surprisingly resilient to periodic macroalgal competition, which could have important implications for ecosystem trajectories that lead to reef decline or recovery.
Impact of nitrogen chemical form on the isotope signature and toxicity of a marine dinoflagellate
Armstrong CT, Erdner DL, McClelland JW, Sanderson MP, Anderson DM, Gobler CJ and Smith JL
Despite a global interest in the relationship between harmful algal blooms (HABs) and eutrophication, the impact of natural versus anthropogenic nutrient sources on species composition or toxicity of HABs remains unclear. Stable isotopes are used to identify and track nitrogen (N) sources to water bodies, and thus can be used to ascertain the N source(s) used by the phytoplankton in those systems. To focus this tool for a particular species, the fundamental patterns of N isotope fractionation by that organism must first be understood. While literature is available describing N isotope fractionation by diatoms and coccolithophores, data are lacking regarding dinoflagellates. Here we investigated the effects of N chemical form on isotope fractionation (Δ) and toxin content using isolates of the autotrophic dinoflagellate, in single-N and mixed-N experiments. Growth of exclusively on nitrate (NO ), ammonium (NH ), or urea, resulted in Δ of 2.7±1.4‰, 29±9.3‰, or 0.3±0.1‰, respectively, with the lowest cellular toxicity reported during urea utilization. Cells initially utilized NH and urea when exposed to mixed-N medium, and only utilized NO after NH decreased below 2-4 μM. This pattern of N preference was similar across all N treatments, suggesting that there is no effect of preconditioning on N chemical preference by . In NO and urea-rich environments, the δN of would resemble the source(s) of N utilized, supporting this tool's utility as a tracer of N source(s) facilitating bloom formation, however, caution is advisable in NH rich environments where the large Δ value could lead to misinterpretation of the signal.