Coral Condition Research Articles

Emergence of a multi host biofilm forming opportunistic pathogen Staphylococcus sciuri D26 in coral Favites abdita

Corals are hotspots of ocean microbial diversity and imbalance in the composition of coral associated microbes has been mostly correlated with the emergence of climate change driven diseases which affect the overall stability of the reef ecosystem. Coral sampling was performed by SCUBA diving at Palk Bay (latitude 9.271580, longitude 79.132203) south Indian coast. Among the 54 bacterial isolates, an isolate MGL-D26 showed comparatively high biofilm formation and was identified as Staphylococcus sciuri based on phylogenetic analysis. The production of exopolysaccharide (EPS) confirmed the formation of a slimy EPS matrix associated with the biofilm. The biofilm formation in S. sciuri D26 was induced significantly by UV exposure followed by other stress factors including pollution, agitation, and salinity. The strain inhibited innate immune factors of corals such as melanin synthesis and phenoloxidase. Challenge experiments in a model organism Aiptasia sp. showed pathogenicity of S. sciuri. Histopathological analysis revealed tissue invasion by S. sciuri which was a predisposing factor leading to mortality in challenged Aiptasia sp. However, specific disease condition of corals infected by S. sciuri requires continuous field monitoring and further investigation. Based on the findings, S. sciuri was a first reported multi-host opportunistic pathogen which has emerged in corals under environmental stress.

Effects of suspended drill cuttings on the coral Lophelia pertusa using pulsed and continuous exposure scenarios

ABSTRACTThis study aimed at providing confidence in the predictability of the impacts of drill cuttings (DC) discharge on the cold-water coral Lophelia pertusa. L. pertusa was exposed to DC from offshore exploration in the lab with the goal to assess precautionary thresholds of effects. Two exposure scenarios with DC were tested: a long-term (LT) pulsed exposure (12 weeks, peak concentrations: 2–50 mg/L, mean concentrations: 1–25 mg/L) and a short-term (ST) continuous exposure (2.5 weeks, mean concentrations: 4–42 mg/L). After exposure, a recovery period of 16 and 4 weeks was maintained in LT and ST, respectively. While there was an assumption that DC might result in an increase in respiration, decrease in growth, enhanced mucus production, reduced fatty acid content, only a significant rise was noted in skeleton growth at DC 4 mg/L and a significant increase of mucus particulate organic carbon at 25 mg/L at end of the exposure. DC did not markedly reduce prey capture rate consecutive to DC exposure. However, the effect of DC produced an increase of coral polyp activity during exposure and a return to pre-exposure conditions after cessation of DC, and coenosarc was smothered from DC even after a long recovery period (4 weeks). Overall, a DC concentration of 10 mg/L seems to represent a threshold above which changes in coral conditions were observed however with no apparent physiological consequences for the coral within the experimental time scale.

Diversity of Coral Fish At Saebus Island, East Java, Indonesia

Coral reef ecosystem is known as an important place to live various types of fish, where coral conditions will affect the diversity and abundance of the fish. In healthy coral reef ecosystems generally can be found many types of fish with high density. This research aims to investigate the diversity and abundance of coral fishes at Saebus Island, East Java. The observation conducted at 4 stations, according to cardinal point by UVS (underwater visual census) methods with belt transect with the visibility of 2,5 m horizontally, and 5 m vertically. The length of transect was 100 m parallel with coastline, with the area of observation is 500 m2. The censuses were conducted at 2 different depths (3 and 10 m). This study found 70 kinds of coral fish originated form 20 family at all stations. These fishes were from 3 different fish categories i.e. 7 target fishes, 13 indicator fishes, and 50 major fishes. Three different fishes that dominated target fish, indicator fish and major fish wereEpinephelus fasciatus, Chaetodon baronessaandAulostomus chinensis,respectively. There was similar value of fish diversity index at two different depths which were 3.635 and 3,623. While uniformity index at the depth of 3m was 0.153 and at 10m was 0.217, and domination index at the depth of 3m was 0.11 and at 10m was 0.167. These values suggest that diversity of coral fish at Saebus island can be categorized as high diversity.

Porites coral response to an oceanographic and human impact gradient in the Line Islands

AbstractCoral bleaching caused by heat stress (warm water) will arguably be the greatest driver of coral reef loss in the coming decades. Understanding how corals have adapted to distinct oceanographic regimes on multiple scales can provide insight into future tolerance and persistence, information critical to directed intervention or targeted protections. The northern Line Islands span a gradient in oceanographic regimes across latitudes, with seawater becoming warmer, fresher, more oligotrophic, and more saturated with aragonite away from the equator. The combination of this regional gradient and island‐scale (local) conditions was used as a natural experiment against which to test how massive Porites corals respond to these background conditions and episodic heat stress. We found that coral condition, represented by a metric combining tissue thickness, lipids, and calcification, was similar at almost all islands, though there were differences in how corals allocate resources among these biological parameters. Corals at Teraina, the most densely inhabited island, showed evidence for reduced calcification, potentially associated with human impacts and/or outflow of freshwater from the island. In contrast, Porites corals at Palmyra, a wildlife reserve, exhibited unexpectedly high tissue‐condition metrics for its latitude, suggesting an additional food source, possibly plankton‐rich lagoonal outflow. We did not find a strong relationship between human habitation and the response of Porites corals to recent heat stress. However, differences in coral calcification rates and energy stores between observed values and those expected based solely on the regional oceanographic gradient, suggests local effects have indirect impacts on these corals.

Macroalgae reduces survival of nursery‐rearedAcroporacorals in the Florida reef tract

Recent declines in coral populations along the Florida reef tract have prompted the establishment of coral restoration programs which raise coral species, such as the threatenedAcropora cervicornis, in nurseries ready for outplanting. Large numbers of nursery‐reared coral colonies have been outplanted along the Florida reef tract in recent years, yet few studies have characterized benthic habitats that are considered optimal for colony survival. In 2016, we surveyed 23A. cervicornisrestoration sites, located at six different reefs in the upper Florida Keys. We examined the condition of the outplanted corals and quantified the benthic assemblages adjacent to the outplanted coral colonies. We found that whereA. cervicornissurvived for more than 1 year, the substrate significantly supported less brown macroalgae of the genusDictyotathan at sites whereA. cervicornishad died. Coral survival was highest at sites with less than 15%Dictyotacover. These results suggest that the habitat conditions that supportedDictyotaspp. were not conducive toA. cervicornisgrowth and survival. Restoration practitioners should avoid attaching nursery‐raised corals to substrate withDictyotaspp. cover greater than 15%.

A comparison of two common sample preparation techniques for lipid and fatty acid analysis in three different coral morphotypes reveals quantitative and qualitative differences.

Lipids are involved in a host of biochemical and physiological processes in corals. Therefore, changes in lipid composition reflect changes in the ecology, nutrition, and health of corals. As such, accurate lipid extraction, quantification, and identification is critical to obtain comprehensive insight into a coral’s condition. However, discrepancies exist in sample preparation methodology globally, and it is currently unknown whether these techniques generate analogous results. This study compared the two most common sample preparation techniques for lipid analysis in corals: (1) tissue isolation by air-spraying and (2) crushing the coral in toto. Samples derived from each preparation technique were subsequently analysed to quantify lipids and their constituent classes and fatty acids in four common, scleractinian coral species representing three distinct morphotypes (Acropora millepora, Montipora crassotuberculata, Porites cylindrica, and Pocillopora damicornis). Results revealed substantial amounts of organic material, including lipids, retained in the skeletons of all species following air-spraying, causing a marked underestimation of total lipid concentration using this method. Moreover, lipid class and fatty acid compositions between the denuded skeleton and sprayed tissue were substantially different. In particular, the majority of the total triacylglycerol and total fatty acid concentrations were retained in the skeleton (55–69% and 56–64%, respectively). As such, the isolated, sprayed tissue cannot serve as a reliable proxy for lipid quantification or identification in the coral holobiont. The in toto crushing method is therefore recommended for coral sample preparation prior to lipid analysis to capture the lipid profile of the entire holobiont, permitting accurate diagnoses of coral condition.

Seasonal variation modulates coral sensibility to heat-stress and explains annual changes in coral productivity

The potential effects of seasonal acclimatization on coral sensitivity to heat-stress, has received limited attention despite differing bleaching thresholds for summer and winter. In this study, we examined the response of two contrasting phenotypes, termed winter and summer, of four Caribbean reef corals to similar light and heat-stress levels. The four species investigated were categorized into two groups: species with the ability to harbour large number of symbionts, Orbicella annularis and O. faveolata, and species with reduced symbiont density (Montastraea cavernosa and Pseudodiploria strigosa). The first group showed higher capacity to enhance photosynthetic rates per area (Pmax), while Pmax enhancement in the second group was more dependent on Symbiodinium performance (Psym). In summer all four species presented higher productivity, but also higher sensitivity to lose coral photosynthesis under heat-stress. In contrast, corals in winter exhibit symbionts with higher capacity to photoacclimate to the increased levels of light-stress elicited by heat-stress. Overall, our study supports the importance of the acclimatory coral condition in addition to the previous thermal history, to determine the severity of the impact of heat-stress on coral physiology, but also the dependence of this response on the particular structural and functional traits of the species.

Intraspecific differences in molecular stress responses and coral pathobiome contribute to mortality under bacterial challenge in Acropora millepora

Disease causes significant coral mortality worldwide; however, factors responsible for intraspecific variation in disease resistance remain unclear. We exposed fragments of eight Acropora millepora colonies (genotypes) to putatively pathogenic bacteria (Vibrio spp.). Genotypes varied from zero to >90% mortality, with bacterial challenge increasing average mortality rates 4–6 fold and shifting the microbiome in favor of stress-associated taxa. Constitutive immunity and subsequent immune and transcriptomic responses to the challenge were more prominent in high-mortality individuals, whereas low-mortality corals remained largely unaffected and maintained expression signatures of a healthier condition (i.e., did not launch a large stress response). Our results suggest that lesions appeared due to changes in the coral pathobiome (multiple bacterial species associated with disease) and general health deterioration after the biotic disturbance, rather than the direct activity of any specific pathogen. If diseases in nature arise because of weaknesses in holobiont physiology, instead of the virulence of any single etiological agent, environmental stressors compromising coral condition might play a larger role in disease outbreaks than is currently thought. To facilitate the diagnosis of compromised individuals, we developed and independently cross-validated a biomarker assay to predict mortality based on genes whose expression in asymptomatic individuals coincides with mortality rates.

Threat of blast fishing on coral diversity in Peucang Island, Ujung Kulon National Park, Indonesia

Gumbira RWW, Rizkia F, Pribadi TDK, Hidayat MS. 2017. Threat of blast fishing on coral diversity in Peucang Island National Park, Indonesia. Ocean Life 1: 26-31. As a maritime country with high biodiversity, the health of Indonesian coral reefs is of high concern. Coral reef ecosystems are friable, and mortality occurs relatively quickly after disturbance, not only by nature but also by human activities such as fish bombing. Fish bombing is a big issue that has always threatened coral reefs, including in Peucang Island, part of Ujung Kulon National Park, a place that is protected by law to preserve its natural diversity. A study on coral coverage was conducted to observe the condition of corals and related biota in a coral reef ecosystem. Data was collected using Point Intercept Transect method in 2m and 8m depths. The results showed that coral coverage in 8-meter depth is only 44.02 percent of the live coral in 2-meter depth. The reduction in the value of Diversity Indices, Dominance Indices as well as the Evenness Index in all study sites led to low diversity, with no dominance by a particular species at any of the study sites. The results also showed that corals are well distributed in both depths. Over the past three years, the damage to corals in Peucang Island had generally increased by 0.7 percent due to fish bombing activities.

Kondisi dan Keanekagaragaman Karang Batu di Perairan Sabang

<strong>Condition and Diversity of Stony Corals in Sabang Waters.</strong> Globally, coral reefs are facing a variety of threats, both natural and human activities. Coral reefs in the waters of Sabang in 2004 were hit by a tsunami and in 2010 an increase in sea surface temperatures caused a bleaching. These resulted in mass mortality of stony corals. This study aims to determine the current condition and the diversity of stony corals in Sabang waters. The study was conducted in May 2015 in ten stations spread over four islands, namely Pulau Weh, Pulau Rondo, Pulau Breueh, and Pulau Nasi Besar. The coral cover is calculated using Underwater Photo Transect (UPT) method. The images were analyzed using CPCe 4.1 program with 30 random points in each frame. The number of species and the number of coral colonies are calculated based on the number of species and coral colonies contained in the photo frames along the transect. Based on the results of the analysis, live coral cover ranged 22.45–58.60% with an average live coral cover of 41.99%. Throughout the transect, 148 species of stony corals from 37 genera and 15 families were discovered. In general, stony corals in Sabang waters are in a good condition, not different from the condition of coral reefs in 2005. Open waters have a positive impact on coral growth, especially for the genus Acropora.

Facilitation in Caribbean coral reefs: high densities of staghorn coral foster greater coral condition and reef fish composition.

Recovery of the threatened staghorn coral (Acropora cervicornis) is posited to play a key role in Caribbean reef resilience. At four Caribbean locations (including one restored and three extant populations), we quantified characteristics of contemporary staghorn coral across increasing conspecific densities, and investigated a hypothesis of facilitation between staghorn coral and reef fishes. High staghorn densities in the Dry Tortugas exhibited significantly less partial mortality, higher branch growth, and supported greater fish abundances compared to lower densities within the same population. In contrast, partial mortality, branch growth, and fish community composition did not vary with staghorn density at the three other study locations where staghorn densities were lower overall. This suggests that density-dependent effects between the coral and fish community may only manifest at high staghorn densities. We then evaluated one facilitative mechanism for such density-dependence, whereby abundant fishes sheltering in dense staghorn aggregations deliver nutrients back to the coral, fueling faster coral growth, thereby creating more fish habitat. Indeed, dense staghorn aggregations within the Dry Tortugas exhibited significantly higher growth rates, tissue nitrogen, and zooxanthellae densities than sparse aggregations. Similarly, higher tissue nitrogen was induced in a macroalgae bioassay outplanted into the same dense and sparse aggregations, confirming greater bioavailability of nutrients at high staghorn densities. Our findings inform staghorn restoration efforts, suggesting that the most effective targets may be higher coral densities than previously thought. These coral-dense aggregations may reap the benefits of positive facilitation between the staghorn and fish community, favoring the growth and survivorship of this threatened species.

Current status of coral reefs in Tioman Island, Peninsular Malaysia

The present study aimed to determine the current status of corals using the coral video transect (CVT) method in the east coast, west coast, and isolated areas of Tioman Island, Peninsular Malaysia. A total of 65 genera from 21 families of corals were identified, from which the scleractinian corals of Acropora, Montipora, and Porites were dominant in the coral assemblages. Nine reef sites were categorised as having ‘good’ (51.4%–60.3%) coral condition and four reef sites as having ‘fair’ (37.6%–49.2%) coral condition. This study concluded that the reefs around Tioman Island were in ‘good’ average coral condition and have high generic diversity with mixed coral morphological structures. The east coast and isolated areas had better live coral cover compared to the west coast area due to less coastal development and human impacts.

Local Stressors, Resilience, and Shifting Baselines on Coral Reefs.

Understanding how and why coral reefs have changed over the last twenty to thirty years is crucial for sustaining coral-reef resilience. We used a historical baseline from Kosrae, a typical small island in Micronesia, to examine changes in fish and coral assemblages since 1986. We found that natural gradients in the spatial distribution of fish and coral assemblages have become amplified, as island geography is now a stronger determinant of species abundance patterns, and habitat forming Acropora corals and large-bodied fishes that were once common on the leeward side of the island have become scarce. A proxy for fishing access best predicted the relative change in fish assemblage condition over time, and in turn, declining fish condition was the only factor correlated with declining coral condition, suggesting overfishing may have reduced ecosystem resilience. Additionally, a proxy for watershed pollution predicted modern coral assemblage condition, suggesting pollution is also reducing resilience in densely populated areas. Altogether, it appears that unsustainable fishing reduced ecosystem resilience, as fish composition has shifted to smaller species in lower trophic levels, driven by losses of large predators and herbivores. While prior literature and anecdotal reports indicate that major disturbance events have been rare in Kosrae, small localized disturbances coupled with reduced resilience may have slowly degraded reef condition through time. Improving coral-reef resilience in the face of climate change will therefore require improved understanding and management of growing artisanal fishing pressure and watershed pollution.

THE ABUNDANCE OF ORNAMENTAL CORALS AFTER MASS DIE OFF IN 1997 ON THE PADANG SHELF REEF SYSTEM, WEST SUMATERA, INDONESIA

ABSTRACTThe Indonesian Ministry of Marine Affairs and Fisheries aims to establish export quota ornamental corals based on scientifically-founded data from the natural environment. The purpose of this study was to determine the abundance of ornamental coral after mass die off in 1997. Ornamental corals were classified into three groups based on the guideline of ornamental coral propagation for trading purposes. Classification categories are rapid (3-6 months), middle (8-12 months), and slow growth rate (>24 months), which relates to their harvesting time when transplanted for the trade purpose. The survey method used line intercept transects to get coral condition data and belt transect with 2 m width to get ornamental coral data. Both line and belt transect have 30 m length and with three replicates. The study showed number of colony and abundance of ornamental coral for fast growth rate group such as Acropora sp (1348 col & 7.49 col/m2), Galaxea sp, Hydnophora exesa, H. microconus, Pocillopora damicornis, Stylopora sp and Montipora sp with the highest abundant found on Acropora sp and Montipora sp (1348 col & 7.49 col/m2). Whereas the coral which from middle growth rate were Leptoseris sp, Pavona sp, Platygyra sp, Favia sp, Favites sp with the highest abundant found on Favia sp (101 col & 0.56 col/m2). Coral species typically used for ornamental coral trade within the slow growth rate category such massive coral were not found on this survey. Due to the massive coral die-off in 1997 and the paucity of ornamental corals, the utilization of ornamental coral should be supported by coral transplantation and the broodstock of slow growth of coral taken from other area. Keywords: ornamental coral, abundant, mass die off, Padang Waters

Acute and subacute toxicity of the polycyclic aromatic hydrocarbon 1-methylnaphthalene to the shallow-water coral Porites divaricata: Application of a novel exposure protocol.

Previous research evaluating hydrocarbon toxicity to corals and coral reefs has generally focused on community-level effects, and results often are not comparable between studies because of variability in hydrocarbon exposure characterization and evaluation of coral health and mortality during exposure. Toxicity of the polycyclic aromatic hydrocarbon 1-methylnaphthalene to the coral Porites divaricata was assessed in a constant exposure toxicity test utilizing a novel toxicity testing protocol uniquely applicable to shallow-water corals, which considered multiple assessment metrics and evaluated the potential for post-exposure mortality and/or recovery. Acute and subacute effects (gross morphological changes, photosynthetic efficiency, mortality, and histologic cellular changes) were evaluated during pre-exposure (4 wk), exposure (48 h), and post-exposure recovery (4 wk) periods. Coral condition scores were used to determine a 48-h median effective concentration of 7442 μg/L. Significant physical and histological changes resulted from exposure to 640 μg/L and 5427 μg/L 1-methylnaphthalene, with a 1-d to 3-d delay in photosynthetic efficiency effects (ΔF/Fm). Pigmented granular amoebocyte area was found to be a potentially useful sublethal endpoint for this species. Coral mortality was used to estimate a 48-h median lethal concentration of 12 123 μg/L. Environ Toxicol Chem 2017;36:212-219. © 2016 SETAC.

Corals in high diversity reefs resist human impact

Coral reefs are amongst the most diverse ecosystems in the biosphere. However, they also represent one of the most threatened marine systems. Apart from global change, especially fishing and tourism affect coral reefs either with mechanical damage or with increase of pollution and sedimentation. Recently, the increase of disturbances has induced extensive changes in community structure and composition of coral reefs. Well-balanced and rich communities can better resist disturbances and show a more rapid recovery, compared to less biodiverse systems.This study assesses the status of coral reefs subjected to several anthropogenic pressures, using a modified version of Coral Condition Index (CCI) that takes into account all Acropora and Pocillopora growth forms and considers a further category of coral damage: the presence of disease. The investigations were carried out at Bangka Island (North Sulawesi, Indonesia), where some of the most flourishing reefs in the country are present. The CCI takes into account the extent of different damages on coral colonies, particularly of the genera Acropora and Pocillopora, being among the most widespread bioconstructors of local coral reefs and very sensitive to anthropogenic disturbances. The aim of the present work is to test whether the CCI is a reliable index in different coral reefs, and to evaluate if highly biodiverse reefs show a better resistance to several human stressors.Data showed high values of CCI (0.9 on average) at all the investigated sites and at the two depths (3 and 9m) for each site, with the most abundant category represented by “healthy coral colonies”. These data indicate a reasonably good health status of the reef in the study area (CCI>0.8). The presence of different types of human pressure in the study area was evaluated through the use of metric proxies. Results do not seem to show any significant influence of such human activities on reef coral status, as shown by the low values of correlation between CCI values and the distances of the study sites from the three main sources of stress (Villages, Resorts and Other). Moreover, the present data seem to confirm that highly biodiverse and well-structured assemblages can resist disturbances more efficiently and that human pressure in the study area is sustainable. Compared to fishing activities, the impact of Scuba diving on coral reef is lower, resulting more sustainable and ecologically non-destructive.CCI summarizes many kinds of information and can be applicable in various areas with different pressures. It is a useful tool that might help to assist and guide management decisions towards alternative development models.

Size Distribution of Massive Porites at Reef Flat in Kondang Merak, Malang, Indonesia

Kondang Merak was lied on south of Java sea that has monitored coral reef inside. Environment condition such as load of terrestrial’s nutrient into the sea, increase of surface water temperature and tourism activities were affected on coral reef health. The massive Porites that common coral found in this location need be monitored gradually to see coral condition over time. The research was conducted on May 2015 in reef flat area of Kondang Merak. The aims of the research were to know distribution of massive Porites and oceanography condition in Kondang Merak. The highest sea surface temperature was 26,46 o C in transect number 2 at station 1 (eastern part). Salinity of sea water was in normal range between 34, 42 o / oo to 35,75 o / oo , dissolved oxygen was on range 6 mg/l sampai 9,1 mg/l and pH was 7,23 sampai 8,68 respectively. Two species on massive Porites they were Porites lobata dan P. lutea . In station 1 the average diameter of P. lobata was 228.57 cm and P. lutea 63.89 cm, in while P. lobata in station 2 was 260.57 cm and P. lutea 153.56 cm.

Development of a Regional Coral Observation Method by a Fluorescence Imaging LIDAR Installed in a Towable Buoy

Coral bleaching and mortality is predicted to increase under global climate change. A new observation technique is required to monitor regional coral conditions. To this end, we developed a light detection and ranging (LIDAR) system installed in a towable buoy for boat observations, which acquires continuous fluorescent images of the seabed during day-time. Most corals have innate fluorescent proteins in their tissue, and they emit fluorescence by ultraviolet excitation. This fluorescence distinguishes living coral from dead coral skeleton, crustose coralline algae, and sea algae. This paper provides a proof of concept for using the LIDAR system and fluorescence to map coral distribution within 1 km scale and coral cover within 100 m scale for a single reef in Japan.

Fluctuations in coral health of four common inshore reef corals in response to seasonal and anthropogenic changes in water quality

Environmental drivers of coral condition (maximum quantum yield, symbiont density, chlorophyll a content and coral skeletal growth rates) were assessed in the equatorial inshore coastal waters of Singapore, where the amplitude of seasonal variation is low, but anthropogenic influence is relatively high. Water quality variables (sediments, nutrients, trace metals, temperature, light) explained between 52 and 83% of the variation in coral condition, with sediments and light availability as key drivers of foliose corals (Merulina ampliata, Pachyseris speciosa), and temperature exerting a greater influence on a branching coral (Pocillopora damicornis). Seasonal reductions in water quality led to high chlorophyll a concentrations and maximum quantum yields in corals, but low growth rates. These marginal coral communities are potentially vulnerable to climate change, hence, we propose water quality thresholds for coral growth with the aim of mitigating both local and global environmental impacts.

Codition of Coral Reef at Teluk Pandan Sub-District East Kutai District

Coral reef of TelukPandan Sub-district East Kutai District still low in both monitoring and management, thus also poorly in coral reef database. This research purpose was to obtain the data of coral reef condition and distribution description in this sub-district. Research was held on October 2013 at six observation stations using lifeform line intercept transect survey method. The coral condition estimation was based on live coral cover (LC) and hard coral mortality index (HCM). Coral reef distribution and wide were estimated by satellite image Landsat 7 ETM+ processing and direct field tracking using GPS. There estimated ± 618.63 ha of coral reef area consist of fringing and patch reef formation type. Live coral cover vary 2.1-67.8% or categorized “poor” to “good” reef condition and in average LC = 38.1% (“fair/moderate” condition). Hard coral morality (HCM) index ranged 0.14 – 0.8 and in average 0.31.