Responses of three tropical seagrass species to CO2 enrichment

Increased atmospheric carbon dioxide leads to ocean acidification and carbon dioxide (CO2) enrichment of seawater. Given the important ecological functions of seagrass meadows, understanding their responses to CO2 will be critical for the management of coastal ecosystems. This study examined the physiological responses of three tropical seagrasses to a range of seawater pCO2 levels in a laboratory. Cymodocea serrulata, Halodule uninervis and Thalassia hemprichii were exposed to four different pCO2 treatments (442-1204 µatm) for 2 weeks, approximating the range of end-of-century emission scenarios. Photosynthetic responses were quantified using optode-based oxygen flux measurements. Across all three species, net productivity and energetic surplus (PG:R) significantly increased with a rise in pCO2 (linear models, P < 0.05). Photosynthesis-irradiance curve-derived photosynthetic parameters-maximum photosynthetic rates (P max) and efficiency (alpha) also increased as pCO2 increased (linear models, P < 0.05). The response for productivity measures was similar across species, i.e. similar slopes in linear models. A decrease in compensation light requirement (Ec) with increasing pCO2 was evident in C. serrulata and H. uninervis, but not in T. hemprichii. Despite higher productivity with pCO2 enrichment, leaf growth rates in C. serrulata did not increase, while those in H. uninervis and T. hemprichii significantly increased with increasing pCO2 levels. While seagrasses can be carbon-limited and productivity can respond positively to CO2 enrichment, varying carbon allocation strategies amongst species suggest differential growth response between species. Thus, future increase in seawater CO2 concentration may lead to an overall increase in seagrass biomass and productivity, as well as community changes in seagrass meadows.

Relative abundance of diatom species in surface samples from the Southern Ocean, Tropical/Subtropical species

This paper gives a modern circumscription of Tropical/Subtropical diatoms regarding their relationship with sea-surface temperatures (SST) and sea ice cover. Diatoms from 228 core-top sediment samples collected from the Southern Ocean were studied to determine the geographic distribution of eight major diatom species/taxa preserved in surface sediments generally located north of the Subantarctic Front. The comparison of the relative contribution of diatom species with modern February SST and sea-ice cover reveals species-specific sedimentary distributions regulated both by water temperatures and sea ice conditions. Although selective preservation might have played some role, their presence in surface and downcore sediments from the Southern Ocean are reliable indicators of high SST and poleward transport of waters from the Tropical/Subtropical Atlantic. Our work supports the use of diatom remains to reconstruct past variations of these environmental parameters via qualitative and transfer function approaches.

Projected near-future CO2 levels increase activity and alter defensive behaviours in the tropical squid Idiosepius pygmaeus

Carbon dioxide (CO2) levels projected to occur in the oceans by the end of this century cause a range of behavioural effects in fish, but whether other highly active marine organisms, such as cephalopods, are similarly affected is unknown. We tested the effects of projected future CO2 levels (626 and 956 µatm) on the behaviour of male two-toned pygmy squid, Idiosepius pygmaeus. Exposure to elevated CO2 increased the number of active individuals by 19-25% and increased movement (number of line-crosses) by nearly 3 times compared to squid at present-day CO2. Squid vigilance and defensive behaviours were also altered by elevated CO2 with >80% of individuals choosing jet escape responses over defensive arm postures in response to a visual startle stimulus, compared with 50% choosing jet escape responses at control CO2. In addition, more escape responses were chosen over threat behaviours in body pattern displays at elevated CO2 and individuals were more than twice as likely to use ink as a defence strategy at 956 µatm CO2, compared with controls. Increased activity could lead to adverse effects on energy budgets as well as increasing visibility to predators. A tendency to respond to a stimulus with escape behaviours could increase survival, but may also be energetically costly and could potentially lead to more chases by predators compared with individuals that use defensive postures. These results demonstrate that projected future ocean acidification affects the behaviours of a tropical squid species.

Planktonic foraminifera in the latest Maastrichtian of Walvis Ridge, South Atlantic

An abrupt global warming of 3-4°C occurred near the end of the Maastrichtian at 65.45-65.10 Ma. The environmental effects of this warm event are here documented based on stable isotopes and quantitative analysis of planktonic foraminifera at the South Atlantic DSDP Site 525A. Stable isotopes of individual species mark a rapid increase in temperature and a reduction in the vertical water mass stratification that is accompanied by a decrease in niche habitats, reduced species diversity and/or abundance, smaller species morphologies or dwarfing, and reduced photosymbiotic activity. During the warm event, the relative abundance of a large number of species decreased, including tropical-subtropical affiliated species, whereas typical mid-latitude species retained high abundances. This indicates that climate warming did not create favorable conditions for all tropical-subtropical species at mid-latitudes and did not cause a massive retreat in the local mid-latitude population. A noticeable exception is the ecological generalist Heterohelix dentata Stenestad that dominated during the cool intervals, but significantly decreased during the warm event. However, dwarfing is the most striking response to the abrupt warming and occurred in various species of different morphologies and lineages (e.g. biserial, trochospiral, keeled globotruncanids). Dwarfing is a typical reaction to environmental stress conditions and was likely the result of increased reproduction rates. Similarly, photosymbiotic activity appears to have been reduced significantly during the maximum warming, as indicated by decreased delta13C values. The foraminiferal response to climate change is thus multifaceted resulting in decreased species diversity, decreased species populations, increased competition due to reduced niche habitats, dwarfing and reduced photosymbiotic activity.

Neogene planktonic foraminifera of ODP Site 130-806

The Neogene biostratigraphy presented here is based on the study of 230 samples through 737 m of pelagic sediment in Hole 806B. Sediment accumulation is interrupted only once in the uppermost lower Miocene (Zone N6), apparently coincident with a widespread deep-sea hiatus. Preservation of planktonic foraminifers through the section ranges from good to moderately poor. One hundred and ten species of planktonic foraminifers were identified; taxonomic notes on most species are included. All of the standard low-latitude Neogene foraminiferal zones are delineated, with the exceptions of Zones N8 and N9 because of a high first occurrence of Orbulina, and Zones N18 and N19 because of a high first occurrence of Sphaeroidinella dehiscens. Good agreement exists between the published account of the variation in planktonic foraminiferal species richness and the rates of diversification and turnover, and measurements of these evolutionary indexes in the record of Hole 806B. The global pattern of change in tropical/transitional species richness is paralleled in Hole 806B, with departures caused by either ecological conditions peculiar to the western equatorial Pacific or by inexactness in the estimation of million-year intervals in Hole 806B. Temporal changes in the relative abundance of taxa in the sediment assemblages, considered in light of their depth habitats, reveal a detailed picture of historical change in the structure of the upper water column over the Ontong Java Plateau. The dominance of surface dwellers (Paragloborotalia kugleri, P. mayeri, Dentoglobigerina altispira, Globigerinita glutinata, and Globigerinoides spp.) throughout the lower and middle Miocene is replaced by a more equitable distribution of surface (D. altispira and Globigerinoides spp.), intermediate (Globorotalia menardii plexus), and deep (Streptochilus spp.) dwellers in the late Miocene, following the closing of the Indo-Pacific Seaway and the initiation of large-scale glaciation in the Antarctic. The shoaling of the thermocline along the equator engendered by these climatic and tectonic events persisted through the Pliocene, when initial increases in the abundance of a new set of shallow, intermediate, and deep dwelling species of planktonic foraminifers coincide with the closing of the Panamanian Seaway.

Surface sediment analyses, bivalve and gastropod taxa of marine sediments from the Banc d'Arguin, Mauritania

High-nutrient tropical carbonate systems are known to produce sediments that, in terms of skeletal composition, are reminiscent of their extra-tropical counterparts. Such carbonate systems and associated carbonate grain assemblages in the tropics are rare in the present-day world. Nonetheless, it is crucial to gain a better understanding of those ecosystems, including their drivers and players because such settings potentially represent models for ancient depositional systems as well as for predicted future environmental conditions. One of the modern occurrences of eutrophic tropical carbonate systems is the northern Mauritanian Shelf. The marine environment is characterized by an eastern boundary upwelling system that pushes cool and nutrient-rich intermediate waters onto a wide epicontinental platform (Golfe d'Arguin) where the waters warm up to tropical temperatures. The resulting facies is mixed carbonate-siliciclastic with a dominant foramol association grading into bimol and barnamol grain assemblages in the shallowest areas forming the Banc d'Arguin. Besides this cool water-related heterozoan association, the carbonate sediment is characterized by tropical molluskan species, while chlorozoan biota (e.g., corals and algal symbiont-bearing foraminifers) are entirely absent. We here present a first comprehensive facies analysis of this model example of eutrophic tropical carbonates. Furthermore, we reconstruct the loci of carbonate production and provide a conclusive depositional model of the Banc d'Arguin that received little attention to date due to its poorly accessible nature.

Paleogene and early Neogene planktonic foraminifera of ODP sites 119-738 and 119-744

A virtually complete composite history of Cenozoic pelagic sedimentation was recovered from ODP Sites 738 (62°43' S) and 744 (61°35' S), drilled during Leg 119 on the Kerguelen Plateau. An excellent magnetobiochronologic record was obtained from upper Eocene through Holocene sediments at Site 744, and an expanded lower Paleocene through lower Oligocene sequence was cored at Hole 738. Analysis of the stratigraphic distribution of over 125 planktonic foraminifer taxa from these sites reveals changes in species composition that were strongly influenced by the climatic evolution of Antarctic water masses. Early Paleocene planktonic foraminifer assemblages are nearly identical in species composition to coeval assemblages from low and middle latitude sites, showing the same patterns of post-extinction recovery and taxonomic radiation. Biogeographic isolation, revealed by the absence of tropical keeled species, became apparent by late early Paleocene time. Diversity increased near the Paleocene/Eocene boundary when keeled morozovellids immigrated to the Kerguelen Plateau. Greatest diversity (23 species) was achieved by early Eocene time, corresponding to a Cenozoic warming maximum that has been recognized in lower Eocene deep sea and terrestrial sediments worldwide. A gradual decline in diversity from the late early through middle Eocene, primarily due to the disappearance of acarininids, parallels the record of cooling paleotemperatures in Southern Ocean surface waters. Chiloguembelina-dominated assemblages appeared in the late middle Eocene and persisted through the early Oligocene as Antarctic surface waters became thermally isolated. Late Eocene and early Oligocene assemblages exhibit considerably lower diversity than the older Eocene faunas, and were dominated by chiloguembelinids, subbotinids, and catapsydracids during a time of pronounced climatic cooling and development of continental glaciation on East Antarctica. The small foraminifer Globigerinit? juvenilis replaced chiloguembelinids as the dominant taxon during the late Oligocene. Diversity increased slightly toward the end of the late Oligocene with new appearances of several tenuitellid, globoturborotalitid, and globigerinid species. The trend toward diminishing planktonic foraminifer diversity was renewed during the late early Miocene as siliceous productivity increased in the Antarctic surface waters, culminating with the reduction to nearly monospecific assemblages of Neogloboqu?drin? p?chyderm? that occur in Pliocene-Holocene biosiliceous sediments. An Antarctic Paleogene zonal scheme previously devised for ODP Sites 689 and 690 in the Weddell Sea is used to biostratigraphically subdivide the Kerguelen Plateau sequence. The definition of one Antarctic Paleogene biozone is modified in the present study to facilitate correlation within the southern high latitudes. The ages of 13 late Eoceneearly Miocene datum events are calibrated based on a magnetobiochronologic age model developed for Site 744.

Seawater carbonate chemistry, thickness and carbonate elemental composition of the test of juvenile sea urchins in a laboratory experiment

Ocean surface CO2 levels are increasing in line with rising atmospheric CO2 and could exceed 900 µatm by year 2100, with extremes above 2000 µatm in some coastal habitats. The imminent increase in ocean pCO2 is predicted to have negative consequences for marine fishes, including reduced aerobic performance, but variability among species could be expected. Understanding interspecific responses to ocean acidification is important for predicting the consequences of ocean acidification on communities and ecosystems. In the present study, the effects of exposure to near-future seawater CO2 (860 µatm) on resting (M O2rest) and maximum (M O2max) oxygen consumption rates were determined for three tropical coral reef fish species interlinked through predator-prey relationships: juvenile Pomacentrus moluccensis and Pomacentrus amboinensis, and one of their predators: adult Pseudochromis fuscus. Contrary to predictions, one of the prey species, P. amboinensis, displayed a 28-39% increase in M O2max after both an acute and four-day exposure to near-future CO2 seawater, while maintaining M O2rest. By contrast, the same treatment had no significant effects on M O2rest or M O2max of the other two species. However, acute exposure of P. amboinensis to 1400 and 2400 µatm CO2 resulted in M O2max returning to control values. Overall, the findings suggest that: (1) the metabolic costs of living in a near-future CO2 seawater environment were insignificant for the species examined at rest; (2) the M O2max response of tropical reef species to near-future CO2 seawater can be dependent on the severity of external hypercapnia; and (3) near-future ocean pCO2 may not be detrimental to aerobic scope of all fish species and it may even augment aerobic scope of some species. The present results also highlight that close phylogenetic relatedness and living in the same environment, does not necessarily imply similar physiological responses to near-future CO2.

The O2, pH and Ca2+ Microenvironment of Benthic Foraminifera in a High CO2 World

Ocean acidification (OA) can have adverse effects on marine calcifiers. Yet, phototrophic marine calcifiers elevate their external oxygen and pH microenvironment in daylight, through the uptake of dissolved inorganic carbon (DIC) by photosynthesis. We studied to which extent pH elevation within their microenvironments in daylight can counteract ambient seawater pH reductions, i.e. OA conditions. We measured the O2 and pH microenvironment of four photosymbiotic and two symbiont-free benthic tropical foraminiferal species at three different OA treatments (~432, 1141 and 2151 µatm pCO2). The O2 concentration difference between the seawater and the test surface (delta O2) was taken as a measure for the photosynthetic rate. Our results showed that O2 and pH levels were significantly higher on photosymbiotic foraminiferal surfaces in light than in dark conditions, and than on surfaces of symbiont-free foraminifera. Rates of photosynthesis at saturated light conditions did not change significantly between OA treatments (except in individuals that exhibited symbiont loss, i.e. bleaching, at elevated pCO2). The pH at the cell surface decreased during incubations at elevated pCO2, also during light incubations. Photosynthesis increased the surface pH but this increase was insufficient to compensate for ambient seawater pH decreases. We thus conclude that photosynthesis does only partly protect symbiont bearing foraminifera against OA.

Photosynthate translocation increases in response to low seawater pH in a coral-dinoflagellate symbiosis

This study has examined the effect of low seawater pH values (induced by an increased CO2 partial pressure) on the rates of photosynthesis, as well as on the carbon budget and carbon translocation in the scleractinian coral species Stylophora pistillata, using a new model based on 13C labelling of the photosynthetic products. Symbiont photosynthesis contributes to a large part of the carbon acquisition in tropical coral species, and it is thus important to know how environmental changes affect this carbon acquisition and allocation. For this purpose, nubbins of S. pistillata were maintained for six months at two pHTs (8.1 and 7.2, by bubbling seawater with CO2). The lowest pH value was used to tackle how seawater pH impacts the carbon budget of a scleractinian coral. Rates of photosynthesis and respiration of the symbiotic association and of isolated symbionts were assessed at each pH. The fate of 13C photosynthates was then followed in the symbionts and the coral host for 48 h. Nubbins maintained at pHT 7.2 presented a lower areal symbiont concentration, and lower areal rates of gross photosynthesis and carbon incorporation compared to nubbins maintained at pHT 8.1. The total carbon acquisition was thus lower under low pH. However, the total percentage of carbon translocated to the host as well as the amount of carbon translocated per symbiont cell were significantly higher under pHT 7.2 than under pHT 8.1 (70% at pHT 7.2 vs. 60% at pHT 8.1), such that the total amount of photosynthetic carbon received by the coral host was equivalent under both pHs (5.5 to 6.1 µg C/cm**2/h). Although the carbon budget of the host was unchanged, symbionts acquired less carbon for their own needs (0.6 compared to 1.8 µg C/cm**2/h), explaining the overall decrease in symbiont concentration at low pH. In the long term, such decrease in symbiont concentration might severely affect the carbon budget of the symbiotic association.

Seawater carbonate chemistry and weight of two Atlantic corals Favia fragum and Porites astreoides during experiments, 2011

Rising concentrations of atmospheric CO2 are changing the carbonate chemistry of the oceans, a process known as ocean acidification (OA). Absorption of this CO2 by the surface oceans is increasing the amount of total dissolved inorganic carbon (DIC) and bicarbonate ion (HCO3) available for marine calcification yet is simultaneously lowering the seawater pH and carbonate ion concentration ([CO3]), and thus the saturation state of seawater with respect to aragonite. We investigated the relative importance of [HCO3] versus [CO3] for early calcification by new recruits (primary polyps settled from zooxanthellate larvae) of two tropical coral species, Favia fragum and Porites astreoides. The polyps were reared over a range of ?ar values, which were manipulated by both acid-addition at constant pCO2 (decreased total [HCO3] and [CO3]) and by pCO2 elevation at constant alkalinity (increased [HCO3], decreased [CO3]). Calcification after 2 weeks was quantified by weighing the complete skeleton (corallite) accreted by each polyp over the course of the experiment. Both species exhibited the same negative response to decreasing [CO3] whether ?ar was lowered by acid-addition or by pCO2 elevation-calcification did not follow total DIC or [HCO3]. Nevertheless, the calcification response to decreasing [CO3] was nonlinear. A statistically significant decrease in calcification was only detected between Omega aragonite = <2.5 and Omega aragonite = 1.1-1.5, where calcification of new recruits was reduced by 22-37% per 1.0 decrease in Omega aragonite. Our results differ from many previous studies that report a linear coral calcification response to OA, and from those showing that calcification increases with increasing [HCO3]. Clearly, the coral calcification response to OA is variable and complex. A deeper understanding of the biomineralization mechanisms and environmental conditions underlying these variable responses is needed to support informed predictions about future OA impacts on corals and coral reefs.

Avian species identity and predation success in tropical cacao agroforestry of the Napu Valley in Central Sulawesi, Indonesia

Avian ecosystem services such as the suppression of pests are considered being of high ecological and economic importance in a range of ecosystems, especially in tropical agroforestry. But how bird predation success is related to the diversity and composition of the bird community, as well as local and landscape factors, is poorly understood. The author quantified arthropod predation in relation to the identity and diversity of insectivorous birds, using experimental exposure of artificial, caterpillar-like prey on smallholder cacao agroforestry systems, differing in local shade management and distance to primary forest. The bird community was assessed using both mist netting (targeting on active understory insectivores) and point count (higher completeness of species inventories) sampling. The study was conducted in a land use dominated area in Central Sulawesi, Indonesia, adjacent to the Lore Lindu National Park. We selected 15 smallholder cacao plantations as sites for bird and bat exclosure experiments in March 2010. Until July 2011, we recorded several data in this study area, including the bird community data, cacao tree data and bird predation experiments that are presented here. We found that avian predation success can be driven by single and abundant insectivorous species, rather than by overall bird species richness. Forest proximity was important for enhancing the density of this key species, but did also promote bird species richness. The availability of local shade trees had no effects on the local bird community or avian predation success. Our findings are both of economical as well as ecological interest because the conservation of nearby forest remnants will likely benefit human needs and biodiversity conservation alike.

Species-specific consequences of ocean acidification for the calcareous tropical green algae Halimeda, 2011

Ocean acidification (OA), resulting from increasing dissolved carbon dioxide (CO2) in surface waters, is likely to affect many marine organisms, particularly those that calcify. Recent OA studies have demonstrated negative and/or differential effects of reduced pH on growth, development, calcification and physiology, but most of these have focused on taxa other than calcareous benthic macroalgae. Here we investigate the potential effects of OA on one of the most common coral reef macroalgal genera,Halimeda. Species of Halimeda produce a large proportion of the sand in the tropics and are a major contributor to framework development on reefs because of their rapid calcium carbonate production and high turnover rates. On Palmyra Atoll in the central Pacific, we conducted a manipulative bubbling experiment to investigate the potential effects of OA on growth, calcification and photophysiology of 2 species of Halimeda. Our results suggest that Halimeda is highly susceptible to reduced pH and aragonite saturation state but the magnitude of these effects is species specific. H. opuntiasuffered net dissolution and 15% reduction in photosynthetic capacity, while H. taenicola did not calcify but did not alter photophysiology in experimental treatments. The disparate responses of these species to elevated CO2 partial -pressure (pCO2) may be due to anatomical and physiological differences and could represent a shift in their relative dominance in the face of OA. The ability for a species to exert biological control over calcification and the species specific role of the carbonate skeleton may have important implications for the potential effects of OA on ecological function in the future.