Effects of allelopathic compounds produced by native and invasive marine macroalgae on the associated communities

In this study, the production of bioactive secondary metabolites called "allelochemicals" by algae has been investigated, specifically focusing on polyunsaturated aldehydes (PUAs). PUAs are known to have adverse effects on planktonic grazers and on phytoplankton; however, their effect on benthic communities has been poorly studied. Macroalgae are ecosystem engineers that play an important role in the structure of the habitat and associated communities, presenting a great variability in their morphology and structural complexity, which is a primary factor in the structuring of associated communities. In recent decades, it has been seen how the introduction of invasive species can modify the benthic habitat structure, causing cascading effects on the trophic chain. The thesis includes several field and laboratory studies. Field studies examined aldehyde production by native and invasive macroalgal species (Sargassum muticum, in the Adriatic Sea, and Rugulopterix okamurae in the Strait of Gibraltar), their structural complexity, together with their associated phyto and meiobenthos. Two laboratory studies were conducted. The first one, based on microcosms experiments, evaluated the effect of PUA (produced by the diatom Skeletonema marinoi, or as decadienal analytical standard) on meiofauna. The second one evaluated the inhibitory effect of dilkamural, an allelopathic compound isolated from R. okamurae, on unicellular phototrophs. Our results showed that PUAs produced by macroalgae were species-specific and had a significant impact on the benthic community. The morphology of macroalgae was an important factor in shaping associated communities, particularly for microphytobenthos. Invasive species, such as S. muticum and R. okamurae, could reduce the biodiversity of native benthic communities and simplify the habitat. Dilkamural was hypothesized to be an allelochemical defense, and laboratory toxicity tests confirmed this hypothesis. Overall, this thesis sheds light on the importance of allelochemicals and macroalgal structural complexity in the benthic environment and highlights the potential impact of invasive species.

Effects of concentrated viral communities on photosynthesis and community composition of co-occurring benthic microalgae and phytoplankton

Marine viruses have been shown to affect phytoplankton productivity; however, there are no reports on the effect of viruses on benthic microalgae (microphytobenthos). Hence, this study investigated the effects of elevated concentrations of virus-like particles on the photosynthetic physiology and community composition of benthic microalgae and phytoplankton. Virus populations were collected near the sediment surface and concentrated by tangential flow ultrafiltration, and the concentrate was added to benthic and water column samples that were obtained along a eutrophication gradient in the Brisbane River/Moreton Bay estuary, Australia. Photosynthetic and community responses of benthic microalgae, phytoplankton and bacteria were monitored over 7 d in aquaria and in situ. Benthic microalgal communities responded to viral enrichment in both eutrophic and oligotrophic sediments. In eutrophic sediments, Euglenophytes (Euglena sp.) and bacteria decreased in abundance by 20 to 60 and 26 to 66%, respectively, from seawater controls. In oligotrophic sediments, bacteria decreased in abundance by 30 to 42% from seawater controls but the dinoflagellate Gymnodinium sp. increased in abundance by 270 to 3600% from seawater controls, The increased abundance of Gymnodinium sp. may be related to increased availability of dissolved organic matter released from lysed bacteria. Increased (140 to 190% from seawater controls) initial chlorophyll a fluorescence measured with a pulse-amplitude modulated fluorometer was observed in eutrophic benthic microalgal incubations following virus enrichment, consistent with photosystem II damage. Virus enrichment in oligotrophic water significantly stimulated carbon fixation rates, perhaps due to increased nutrient availability by bacterial lysis. The interpretation of data from virus amendment experiments is difficult due to potential interaction with unidentified bioactive compounds within seawater concentrates. However, these results show that viruses are capable of influencing microbial dynamics in sediments.

Freshwater sponges (Porifera, Demospongiae) in a benthic filter feeding community at the Guanacaste Dry Forest, Costa Rica

The freshwater sponges Trochospongilla variabilis Bonetto & Ezcurra de Drago (1973), Radiospongilla crateriformis (Potts, 1882), Spongilla cenota Penney & Racek (1968) and Corvoheteromeyenia heterosclera (Ezcurra de Drago, 1974) compose with the sphaerid bivalve Eupera cubensis (Prime, 1865) and several Phylactolaemata bryozoans a benthic filter feeding community living in seasonal lentic and lotic habitats with high Particulate Organic Carbon (POC), low conductivity and acid pH within the Costa Rica Dry Forest biome. The sponge specimens gathered led to the re-description of the four species.

Relationship between space distribution of the benthic macroinvertebrates community and trophic state in a Neotropical reservoir (Itupararanga, Brazil)

The purpose of this work was to verify the benthic macroinvertebrates community responses through environmental factors along a headwater tropical reservoir. Samplings were taken with a Van-Veen grab along the reservoir in littoral and profundal regions and in the headwater, next to the dam and the middle of the reservoir. Samples were taken during both wet and dry seasons. Dissolved oxygen concentrations, electric conductivity, temperature and pH near the sediment have been performed in situ, at every sampling station by using a multiprobe and Secchi disc. Total water phosphorus and chlorophyll a concentrations were analyzed to determine the trophic state index. Sediment's organic matter, total phosphorus, nitrogen concentrations and granulometric composition were measured. In order to verify which environmental variables would have more influence over the benthic macroinvertebrates community, a canonical correspondence analysis (CCA) was performed. The total number of recorded taxa was 28. Among them, the family Chironomidae (Diptera) was the richest group (19 taxa). It can be proposed that the benthic macroinvertebrates community may be influenced by environmental conditions such as nutrient and organic matter availability, as well as dissolved oxygen concentration. Macroinvertebrates are adequate bioindicators of water quality due to their sensibility to environmental changes mentioned before. Chironomus sp, Limnodrilus hoffmeisteri and Branchiura sowerbyi comprises a group that can be considered bio-indicators of eutrophic conditions. A second group can be considered as indicator of mesotrophic conditions. The presence of two or more members from that group which comprises Tanytarsini spp, Fissimentum sp, Pelomus sp and Goeldichironomus sp, like predominant taxa, may indicates mesotrophic conditions.

A STUDY OF DIVERSITY AND COMMUNITY COMPARISON INDICES BY BIOASSAY OF COPPER USING COLONIZED BENTHIC MACROINVERTEBRATES

A community bioassay of copper was performed using benthic macroinvertebrates colonized on multiplate substrate samplers. Five copper concentrations ranging from 0.080-2.20 mg/l total copper were administered to five artificial streams by a Mount and Brungs proportional dilutor. Free copper ion as Cu('++) ranged from .002-.053 mg/l. A sixth stream received no copper and served as a control. Substrates were sampled at days 0, 14, and 28, and the results were used to compare 13 indices used or proposed to assess aquatic environmental impact. Sensitivity of the indices to changes in communities with respect to concentration and time was the basis for the comparison.^ Results indicated that all of the 8 diversity or richness indices tested gave approximately the same result (with the exception of number of species); they increased over the first 2-3 concentrations, then declined. Included among these was the Shannon index which gave false positive results, i.e., it increased, indicating enrichment, when in fact perturbation had occurred. This result was due to the disproportionate effect on the most abundant taxa, which caused a more even distribution of individuals among species. Number of species and individuals declined with increased concentration and time, with only one exception in the case of species, indicating perturbation.^ Results of five community comparison indices were varied at day 14 but by day 28 the results indicated a clear, nearly monotonic, trend due to copper impact. It was assumed that day 28 observations, though probably still changing, were nearer stability than at day 14 and therefore more representative of natural conditions. The changes in community comparison indices showed good agreement at 28 days and reflected the general decline in species and individuals. No single community comparison index could be set apart as superior to the others. ^

Eocene benthic foraminiferal community DSDP of Holes 95-612, 95-613 and 11-108

Benthic foraminiferal biofacies may vary independently of water depth and water mass; however, calibration of biofacies and stratigraphic ranges with independent paleodepth estimates allows reconstruction of age-depth patterns applicable throughout the deep Atlantic (Tjalsma and Lohmann, 1983). We have attempted to test these faunal calibrations in a continental margin setting, reconstructing Eocene benthic foraminiferal distributions along a dip section afforded by the New Jersey Transect (DSDP Sites 612, 108, 613). The following independent estimates of Eocene depths for the transect were obtained by "backtracking," "backstripping," and by assuming increasing depth downdip ("paleoslope"): Site 612, near the middle/lower bathyal boundary (about 1000 m); Site 108, in the middle bathyal zone (about 1600 m); and Site 613, near the lower bathyal/upper abyssal boundary (about 2000 m). Within uncertainties of backtracking (hundreds of meters), these estimates agree with estimates of paleodepth based on comparison of the New Jersey margin biofacies with other backtracked faunas. The stratigraphic ranges of many benthic taxa correspond to those found at other Atlantic DSDP sites. The major biofacies patterns show: (1) a depth dichotomy between an early to middle Eocene Nuttallides truempyidominated biofacies (greater than 2000 m) and a Lenticulina-Osangularia-Alabamina cf. dissonata biofacies (1000- 2000 m); and (2) a difference between a middle and a late Eocene biofacies at Site 612. The faunal boundary at about 2000 m, between bathyal and abyssal zones, occurs not only on the margin, but also throughout the deep Atlantic. The faunal change between the middle and late Eocene at Site 612 was due to a decrease of Lenticulina spp., the local disappearance of N. truempyi, and establishment of a Bulimina alazanensis-Gyroidinoides spp. biofacies. Although this change could be attributed to local paleoceanographic or water-depth changes, we argue that it is the bathyal expression of a global deep-sea benthic foraminiferal change which occurred across the middle/late Eocene boundary.

Community structure and biodiversity in marine benthic communities during experiments, 2011

A mesocosm experiment was conducted to quantify the effects of reduced pH and elevated temperature on an intact marine invertebrate community. Standardised faunal communities, collected from the extreme low intertidal zone using artificial substrate units, were exposed to one of eight nominal treatments (four pH levels: 8.0, 7.7, 7.3 and 6.7, crossed with two temperature levels: 12 and 16°C). After 60 days exposure communities showed significant changes in structure and lower diversity in response to reduced pH. The response to temperature was more complex. At higher pH levels (8.0 and 7.7) elevated temperature treatments contained higher species abundances and diversity than the lower temperature treatments. In contrast, at lower pH levels (7.3 and 6.7), elevated temperature treatments had lower species abundances and diversity than lower temperature treatments. The species losses responsible for these changes in community structure and diversity were not randomly distributed across the different phyla examined. Molluscs showed the greatest reduction in abundance and diversity in response to low pH and elevated temperature, whilst annelid abundance and diversity was mostly unaffected by low pH and was higher at the elevated temperature. The arthropod response was between these two extremes with moderately reduced abundance and diversity at low pH and elevated temperature. Nematode abundance increased in response to low pH and elevated temperature, probably due to the reduction of ecological constraints, such as predation and competition, caused by a decrease in macrofaunal abundance. This community-based mesocosm study supports previous suggestions, based on observations of direct physiological impacts, that ocean acidification induced changes in marine biodiversity will be driven by differential vulnerability within and between different taxonomical groups. This study also illustrates the importance of considering indirect effects that occur within multispecies assemblages when attempting to predict the consequences of ocean acidification and global warming on marine communities.

Seawater carbonate chemistry and benthic marine community during experiments, 2011

Ocean acidification is predicted to impact all areas of the oceans and affect a diversity of marine organisms. However, the diversity of responses among species prevents clear predictions about the impact of acidification at the ecosystem level. Here, we used shallow water CO2 vents in the Mediterranean Sea as a model system to examine emergent ecosystem responses to ocean acidification in rocky reef communities. We assessed in situ benthic invertebrate communities in three distinct pH zones (ambient, low, and extreme low), which differed in both the mean and variability of seawater pH along a continuous gradient. We found fewer taxa, reduced taxonomic evenness, and lower biomass in the extreme low pH zones. However, the number of individuals did not differ among pH zones, suggesting that there is density compensation through population blooms of small acidification-tolerant taxa. Furthermore, the trophic structure of the invertebrate community shifted to fewer trophic groups and dominance by generalists in extreme low pH, suggesting that there may be a simplification of food webs with ocean acidification. Despite high variation in individual species' responses, our findings indicate that ocean acidification decreases the diversity, biomass, and trophic complexity of benthic marine communities. These results suggest that a loss of biodiversity and ecosystem function is expected under extreme acidification scenarios.

Neogene benthic foraminiferal community and organic carbon at DSDP Holes 95-612 and 95-613

Quantitative study of benthic foraminifers from the upper Miocene to lower Pliocene section at Site 612 (1404 m present water depth) and the Pliocene section at Site 613 (2323 m present water depth) shows no evidence of widespread downslope transport of shallow-water biofacies or reworking of older material in the greater than 150 µm size fraction. In contrast, upper Miocene sediments from Site 604 (2364 m present water depth) show extensive reworking and downslope transport. At Site 612, benthic foraminifers show a succession from an upper Miocene Bolivina alata-Nonionella sp. biofacies, to an uppermost Miocene Bulimina alazanensis biofacies, to a lower Pliocene Cassidulina reflexa biofacies, to an upper Pliocene Melonis barleeanum-Islandiella laevigata biofacies. Evidence suggests that the Pliocene biofacies are in situ, although they could have been transported downslope from the upper-middle bathyal zone. At Site 613, Uvigerina peregrina dominated the "middle" Pliocene, while Globocassidulina subglobosa was dominant in the early and late Pliocene. High abundances of U. peregrina at Site 613 are associated with high values of sedimentary organic carbon.