globalchange  > 气候变化事实与影响
DOI: doi:10.1038/nclimate2758
论文题名:
Shift from coral to macroalgae dominance on a volcanically acidified reef
作者: I. C. Enochs
刊名: Nature Climate Change
ISSN: 1758-801X
EISSN: 1758-6921
出版年: 2015-08-10
卷: Volume:5, 页码:Pages:1083;1088 (2015)
语种: 英语
英文关键词: Climate-change ecology ; Community ecology ; Marine biology
英文摘要:

Rising anthropogenic CO2 in the atmosphere is accompanied by an increase in oceanic CO2 and a concomitant decline in seawater pH (ref. 1). This phenomenon, known as ocean acidification (OA), has been experimentally shown to impact the biology and ecology of numerous animals and plants2, most notably those that precipitate calcium carbonate skeletons, such as reef-building corals3. Volcanically acidified water at Maug, Commonwealth of the Northern Mariana Islands (CNMI) is equivalent to near-future predictions for what coral reef ecosystems will experience worldwide due to OA. We provide the first chemical and ecological assessment of this unique site and show that acidification-related stress significantly influences the abundance and diversity of coral reef taxa, leading to the often-predicted shift from a coral to an algae-dominated state4, 5. This study provides field evidence that acidification can lead to macroalgae dominance on reefs.

Coral reefs contain the highest concentration of biodiversity in the marine realm, with abundant flora and fauna that form the backbone of complex and dynamic ecosystems6. From an anthropocentric standpoint, coral reefs provide valuable goods and services, supporting fisheries and tourism, and protect shorelines from storms7. Recently, widespread coral mortality has led to the flattening of reef frameworks and the loss of essential habitat4. This trend will be accelerated by ocean acidification (OA), as calcification is impaired, and dissolution is accelerated8, 9. Furthermore, experimental evidence suggests that OA could enhance the growth10 and competitive ability of fleshy macroalgae11. This OA-induced shift in the competitive balance between corals and algae could exacerbate direct effects of OA on calcifying reef species12 and lead to ecosystem shifts favouring non-reef-forming algae over coral4, 5. Understanding the individual responses of taxa to OA, as well as alteration of multi-species assemblages, is therefore critical to predicting ecosystem persistence and managing reef health in an era of global change.

At present, much of what is known concerning the impacts of OA on coral reef biota has been laboratory-based experimental work focused on the responses of select taxa2. This has been expanded to mesocosm-based studies, allowing manipulation of groups of organisms and investigation of community responses13. Although these multi-species experimental studies are vital, they cannot recreate the variability (physical, chemical, biological) of real-world reef systems14. In an effort to overcome the limitations of laboratory studies, real-world low-saturation-state (Ω) sites have been investigated. In the eastern Pacific, nutrient and CO2-enriched upwelled waters impact coral calcification and the precipitation of carbonate cements, influencing the distribution of reefs15. In Mexico, freshwater springs depress Ω, influencing coral calcification and species distributions16. In Palau, restricted circulation and biological activity contribute to elevated pCO2, with little impact on reef communities17. These locations provide insight into community-scale responses to OA; however, variation in other environmental parameters can complicate conclusions.

Volcanic enrichment of CO2 from submarine vents has been shown to impact the structure of temperate and sub-tropical ecosystems, including seagrasses18, rocky-shore and rocky-reef communities19, 20, soft sediments21 and vermetid reefs22. The occurrence of CO2 vents near coral reef ecosystems is rare and, at present, only two regions have been studied: Papua New Guinea (PNG; ref. 23) and a sub-tropical system at Iwotorishima Island, Japan24. High-pCO2 vent communities in Japan, comparable to conditions projected for the end of the century (pH ≈ 7.8) are dominated by soft corals, whereas nearby control sites (pH ≈ 8.1) are dominated by hard scleractinian corals24. In PNG, coral cover is not significantly different at a pH of 7.8, although species composition changes and diversity is reduced23. Given the different ecosystem responses observed at the previously described sites and the paucity of temporally and spatially explicit data sets, further work is necessary to examine the multifarious influences of OA on coral reefs.

Here we identify and characterize a CO2 vent impacting a tropical coral reef ecosystem in the Commonwealth of the Northern Mariana Islands (CNMI; Fig. 1). We use high-accuracy instrumentation to characterize the carbonate chemistry of this system and investigate its relationship with reef community composition (Methods).

Figure 1: Map showing the location of the study site at Maug.
Map showing the location of the study site at Maug.

a, Location of the Commonwealth of the Northern Mariana Islands (CNMI). b, Location of Maug. c, The three main islands of Maug, with 100m isobaths and the location of both the vent and control sites. d, Detail of the vent with the high-pCO2 and mid-pCO2 study sites, together with 2m isobaths.

Study site.

This study was conducted at Maug Island (20° 1′ N, 145° 13 E), in the northernmost region of the CNMI (Fig. 1). Initial investigation of the pH/CO2 gradient was conducted with a pH probe (ROSS Ultra pH, Orion) and non-dispersive infrared CO2 analyser (LI-820, LI-COR Biosciences) paired with a Global Positioning System (GPS). These data were used to inform subsequent chemical, environmental and biological sampling. For the purposes of this study, three sites were established along a gradient of vent influence. A high-pCO2 site was located along the vent field/reef margin. An intermediate, mid-pCO2 site was located roughly 50m south of the vent, in an area dominated by reef framework and coral. Finally, an unaffected control site was located on the southern end of the island, roughly 1km south of the research site. All were located at approximately nine metres depth, to control the influence of extraneous sources of variance during comparison.

Environmental data.

To characterize the extent of carbonate chemistry alteration, 33 discrete water samples were collected in a grid pattern over the area influenced by the vent, covering both the high-pCO2 and mid-pCO2 sites. Water was collected from 20cm below the water’s surface using borosilicate glass bottles, which were immediately fixed with HgCl2 and sealed. Temperature and salinity were recorded at the same depth using a handheld meter (EC300A, YSI) and sites were marked with a handheld GPS (GPSMAP 78S, Garmin). Water samples were collected in the same manner at the control site.

Samples were transported to NOAA’s Atlantic Oceanographic and Meteorological Laboratories (AOML), where they were analysed for dissolved inorganic carbon (DIC) and total alkalinity (TA) using autotitrators (AS-C3 and AS-ALK2 respectively, Apollo SciTech). The carbonic acid system was solved using CO2SYS (ref. 36) with the dissociation constants of ref. 37 as refitted by ref. 38 and ref. 39 for boric acid. Carbonate chemistry parameters were plotted over the extent of the vent using ArcGIS (ESRI). An interpolated raster map was created from these points using the Spatial Analyst Toolbox and the inverse distance weighted (IDW) technique.

SeaFET pH loggers were deployed and recorded data every half hour at each of the three sites (control, mid-pCO2, high-pCO2). Data were collected from 19 May to 10 August 2014. Shorter-term diel oscillation in carbonate chemistry was investigated using discrete water samples collected every 6h over a 48h period from 11 August to 13 August 2014. Water was collected at each of the three study sites immediately above the benthos using a Niskin bottle, and then immediately transferred to borosilicate bottles while minimizing bubble formation and gas exchange. Samples were analysed with the same methodology used for spatial characterization.

Temperature loggers (HOBO Water Temp Pro v2, Onset) were deployed over the same period as the SeaFETs and were attached to stable platform bases approximately 10cm above the benthos at the control, mid-pCO2 and high-pCO2 sites.

Light loggers (ECO-PAR, Wet Labs) were placed at each of the three sites and were programmed to record photosynthetically active radiation (PAR, 400–700nm) every 30min from 19 May to 9 August 2014. The instrument at the high-pCO2 site failed immediately on deployment. The mid-pCO2 and control site instruments were subsequently redeployed at the high-pCO2 and control sites, collecting every 10min from 10–13 August, to measure relative PAR levels. ECO-PAR instruments contain wipers that clean the sensor after each reading, and no drift was observed over the deployment period. We report daily PAR dose following ref. 40, where mean PAR over the period 10 am to 3 pm is multiplied by the total time of that period (5h).

Two acoustic Doppler current profilers (ADCPs, Nortek Aquadopp) were deployed at the high-pCO2 and control sites to measure current. The upward facing devices were turned over during a storm and stopped recording useable data on 4 July.

Vent gas was collected underwater using a conical collection cup connected to gas impermeable 1l Tedlar sampling bags. Sealed bags were transported to Miami and subsequently analysed using gas chromatography (Varian CP3800 and HP 5890).

Biological data.

Changes in benthic cover were investigated using photo quadrats and was conducted across two spatial scales: large-scale differences between the three instrumented high-pCO2, medium-pCO2 and control sites, and fine-scale community shifts occurring outside the zone of active bubbling, expressed as a function of proximity to the high-pCO2 site. For quantification of benthic cover among sites, high-resolution photomosaics were constructed following ref. 41. Mosaics were subsequently subsampled into 100 images per site and the benthic cover under 30 randomly located points were identified using the CPCe software package42. To examine changes in benthic cover with increasing distance from the area of active bubbling, East–West oriented transects, perpendicular to the CO2 gradient, were placed at increasing distance from the vent. Photos were taken every 2m along the 20-metre transect, and were subsequently analysed using CPCe (ref. 42), whereby 40 random points were overlaid over each image and identified.

Finer-level taxonomic identification of coral and algae can be difficult from photographs and community richness data were collected in situ using SCUBA. As with benthic cover, analysis was conducted across both large and small spatial scales. Immediately outside of the vent, six 15m transects were placed, starting at ~5m depth, and arranged perpendicular to the shore (East–West). Transects were spaced 10–20m apart, incrementally further away from the vent site (North–South). Five 0.25m2 quadrats were placed haphazardly along each transect. All algae species within each quadrat were identified to the lowest reliable taxonomic level. Analysis of algae richness was conducted on all identified algae taxa (excluding turf algae), as well as on calcifying algae species, as shown in Supplementary Table 8. Turf algae is defined as the low-lying (<2cm) community of small and juvenile algae species that are not taxonomically distinguishable in situ. Five additional 0.25m2 quadrats were used to quantify coral richness, as listed in Supplementary Table 7. Species-specific prevalences at each site were calculated as the proportion of richness quadrats containing each species, and 95% confidence intervals were calculated following ref. 43. For larger-scale site comparisons of community richness, near-vent data were grouped and compared against three additional transects placed at the control site.

Cores (5cm diam. × 10cm length) were taken from colonies of massive Porites sp. in close proximity to the instrumented mosaic sites using a pneumatic drill and SCUBA tank rig. Cores were slabbed parallel to the growth axis and scanned using microCT (Skyscan 1174, Bruker). Density was plotted versus distance and Coral XDS+ (ref. 44) was used to delineate yearly banding (peak–peak method), as well as to calculate extension, density and calcification rate.

Statistical analysis.

Light was analysed using a t-test (2-tailed). Current and pH data were analysed using nonparametric Mann–Whitney and Kruskal–Wallis tests, respectively. Percentage cover data were arcsine-transformed45 and analysed using general linear models (GLMs). Species richness data were loge-transformed and were analysed using t-tests (2-tailed). Transformation was unnecessary for coral core and calcification data, which were presented by year. Sample-specific averages over a five-year period (2009–2013) were compared between sites using GLMs. Post hoc pair-wise comparisons were conducted with Tukey’s tests.

To investigate the effects of vent proximity on benthic community composition at the vent site, linear (y = b0 + b1x), parabolic (y = b0 + b1x + b2x2), asymptotic (y = b0 + b1x−1) and Ricker models (b0xeb1x) were fitted to percentage coral cover data. Linear, parabolic and asymptotic models were fitted to coral, algae and calcifying algae community richness data. Goodness of fit was evaluated on statistical significance (p < 0.05), R2, and Akaike’s information criterion (AIC). Statistical analysis was conducted with the SPSS and GraphPad Prism software packages46, 47.

  1. Feely, R. A. et al. Impact of anthropogenic CO2 on the CaCO3 system in the oceans. Science 305, 362366 (2004).
  2. Fabry, V. J., Seibel, B. A., Feely, R. A. & James, O. C. Impacts of ocean acidification on marine fauna and ecosystem processes. ICES J. Mar. Sci. 65, 414432 (2008).
  3. Langdon, C. & Atkinson, M. Effect of elevated pCO2 on photosynthesis and calcification of corals and interactions with seasonal change in temperature/irradiance and nutrient enrichment. J. Geophys. Res. 110, C09S07 (2005). URL:
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资源类型: 期刊论文
标识符: http://119.78.100.158/handle/2HF3EXSE/4629
Appears in Collections:气候变化事实与影响
科学计划与规划
气候变化与战略

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I. C. Enochs. Shift from coral to macroalgae dominance on a volcanically acidified reef[J]. Nature Climate Change,2015-08-10,Volume:5:Pages:1083;1088 (2015).
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