Regional genetic differentiation among populations of Cladocora caespitosa in the Western Mediterranean

Cladocora caespitosa is the only reef-forming zooxanthellate scleractinian in the Mediterranean Sea. This endemic coral has suffered severe mortality events at different Mediterranean sites owing to anomalous summer heat waves related to global climate change. In this study, we assessed genetic structure and gene flow among four populations of this species in the Western Mediterranean Sea: Cape Palos (SE Spain), Cala Galdana (Balearic Islands), Columbretes Islands, and L’Ametlla (NE Spain). The results obtained from Bayesian approaches, F _ST statistics, and Bayesian analysis of migration rates suggest certain levels of genetic differentiation driven by high levels of self-recruitment, a fact that is supported by egg-retention mechanisms. Conversely, genetic connectivity among distant populations, even if generally low, seems to be related to sporadic dispersal events through regional surface currents linked to the spawning period that occurs at the end of summer-beginning of autumn. These features, together with a certain isolation of the Columbretes Islands, could explain the regional genetic differentiation found among populations. These results help to better understand population structure and connectivity of the species and will serve as an approach for further studies on different aspects of the biology and ecology of C. caespitosa.     

Environmental drivers of distribution and reef development of the Mediterranean coral Cladocora caespitosa

Coral Reefs - Cladocora caespitosa is the only Mediterranean scleractinian similar to tropical reef-building corals. While this species is part of the recent fossil history of the Mediterranean...     

Monthly variations in calix growth, polyp tissue, and density banding of the Mediterranean scleractinian Cladocora caespitosa (L.)

Monthly skeletal growth of the scleractinian, temperate coral Cladocora caespitosa (L.) from the Ligurian Sea (NW Mediterranean) was analysed for a period of 1 year and compared with seawater parameters. Measurements on corallite sections and on X-ray images showed that the formation of the high-density (HD) band and two dissepiments are favoured by fall–winter conditions, characterised by high quantities of rain, rough seas, and cold seawater. In summer, when the low-density (LD) band is formed, the corallites stretch upward and form one new dissepiment and one deep calix, where the polyps recede almost completely in August. These findings confirmed the adaptation of the temperate coral to winter environmental conditions, characterised by low irradiance and high availability of nutrients and food particles resuspended from bottom sediments. On the contrary, the high seawater temperature, irradiance, and ammonia contents stressed the coral in August and, when they persist in September, may cause the onset of mortality events.     

Relationship between symbiont density and photosynthetic carbon acquisition in the temperate coral Cladocora caespitosa

This study quantified variation in net photosynthetic carbon gain in response to natural fluctuations in symbiont density for the Mediterranean coral Cladocora caespitosa, and evaluated which density maximized photosynthetic carbon acquisition. To do this, carbon acquisition was modeled as an explicit function of symbiont density. The model was parameterized using measurements of rates of photosynthesis and respiration for small colonies with a broad range of zooxanthella concentrations. Results demonstrate that rates of net photosynthesis increase asymptotically with symbiont density, whereas rates of respiration increase linearly. In combination, these functional responses meant that colony energy acquisition decreased at both low and at very high zooxanthella densities. However, there was a wide range of symbiont densities for which net daily photosynthesis was approximately equivalent. Therefore, significant changes in symbiont density do not necessarily cause a change in autotrophic energy acquisition by the colony. Model estimates of the optimal range of cell densities corresponded well with independent observations of symbiont concentrations obtained from field and laboratory studies of healthy colonies. Overall, this study demonstrates that the seasonal fluctuations, in symbiont numbers observed in healthy colonies of the Mediterranean coral investigated, do not have a strong effect on photosynthetic energy acquisition.     

Development of microsatellite markers as a molecular tool for conservation studies of the Mediterranean reef builder coral Cladocora caespitosa (Anthozoa, Scleractinia)

Cladocora caespitosa is a reef-building zooxanthellate scleractinian coral in the Mediterranean Sea. Mortality events have recurrently affected this species during the last decade. Thus, knowledge of its genetic structure, population diversity, and connectivity is needed to accomplish suitable conservation plans. In order to obtain a better understanding of the population genetics of this species, 13 highly variable microsatellites markers were developed from a naturally bleached colony. The developed primers failed to amplify zooxanthella DNA, isolated from C. caespitosa, verifying that these markers were of the coral and not algal symbiont origin. The degree of polymorphism of these loci was tested on tissue samples from 28 colonies. The allele number for each loci ranged from 2 to 13 (mean N(a) = 5.4), with an average observed heterozygosity of 0.42 (H(e) = 0.43) and all loci were in Hardy-Weinberg equilibrium. These new markers should be useful in future conservation genetic studies and will help to improve the resolution of the individual identification within this coral species. Primers were also tested in Oculina patagonica, with successful amplifications of several loci.     

Controlling Effects of Irradiance and Heterotrophy on Carbon Translocation in the Temperate Coral Cladocora caespitosa

Temperate symbiotic corals, such as the Mediterranean species Cladocora caespitosa, live in seasonally changing environments, where irradiance can be ten times higher in summer than winter. These corals shift from autotrophy in summer to heterotrophy in winter in response to light limitation of the symbiont's photosynthesis. In this study, we determined the autotrophic carbon budget under different conditions of irradiance (20 and 120 μmol photons m(-2) s(-1)) and feeding (fed three times a week with Artemia salina nauplii, and unfed). Corals were incubated in H(13)CO(3) (-)-enriched seawater, and the fate of (13)C was followed in the symbionts and the host tissue. The total amount of carbon fixed by photosynthesis and translocated was significantly higher at high than low irradiance (ca. 13 versus 2.5-4.5 μg cm(-2) h(-1)), because the rates of photosynthesis and carbon fixation were also higher. However, the percent of carbon translocation was similar under the two irradiances, and reached more than 70% of the total fixed carbon. Host feeding induced a decrease in the percentage of carbon translocated under low irradiance (from 70 to 53%), and also a decrease in the rates of carbon translocation per symbiont cell under both irradiances. The fate of autotrophic and heterotrophic carbon differed according to irradiance. At low irradiance, autotrophic carbon was mostly respired by the host and the symbionts, and heterotrophic feeding led to an increase in host biomass. Under high irradiance, autotrophic carbon was both respired and released as particulate and dissolved organic carbon, and heterotrophic feeding led to an increase in host biomass and symbiont concentration. Overall, the maintenance of high symbiont concentration and high percentage of carbon translocation under low irradiance allow this coral species to optimize its autotrophic carbon acquisition, when irradiance conditions are not favourable to photosynthesis.     

Recruitment and mortality of the temperate coral Cladocora caespitosa: implications for the recovery of endangered populations

Long-lived species are characterized by low recruitment and mortality. In these species, longevity buffers low recruitment, but when catastrophic disturbances alter mortality, recruitment becomes critical for population recovery. In this study, we assessed basic biological traits—recruitment, post-settlement growth, and the mortality of juvenile corals—and related these factors to the adult mortality of one of the most important populations of the Mediterranean reef-building coral Cladocora caespitosa over a period of 6 yr. Adult mortality and recruitment rates were low (~1 % and 0.30 recruits m^?2 yr^?1, respectively), whereas the juvenile colony mortality was comparatively high (29 % in the smallest size-class, <5 polyps). The low recruitment rates will hardly balance the recurrent climate-related mortality that has affected this population. Conservation plans and inclusion in the protection lists are urgently needed, given the escalating threats and slow dynamics of this species.     

Invasive macrophytes in a marine reserve (Columbretes Islands,NW Mediterranean): spread dynamics and interactions with the endemic scleractinian coral Cladocora caespitosa

The invasive algae Lophocladia lallemandii and Caulerpa racemosa are becoming an important threat to benthic assemblages in the Mediterranean Sea. Both species were first detected in Illa Grossa Bay (Columbretes Islands Marine Reserve, NW Mediterranean) in 2006, and their invasion was monitored until 2012. L. lallemandii showed a rapid outburst, spreading around the entire bay in just 2 years and showing the highest abundances between 5 and 10 m of depth (82.07 ± 3.53 % (±SE) in 2011). Caulerpa racemosa showed a slower but steady spread and remained in deeper areas during the first years; however, drastic changes in the depth distribution, with algae invading toward shallower areas, were noted beginning in 2010 and reached abundances of 57.76 ± 1.07 % (±SE) between 10 and 20 m of depth in 2011. Illa Grossa Bay hosts one of the most important populations of the endemic coral Cladocora caespitosa. This study is the first to quantitatively assess interactions between the coral and invasive algae. Although both invasive species L. lallemandii and C. racemosa had overlapping distributions with C. caespitosa, we did not find any lethal or sublethal effects of either invasive algal species. On the other hand, C. caespitosa exhibited toxic activity, which could explain the low overgrowth of living colony parts by C. racemosa.     

Fossil analogues of present-day Cladocora caespitosa coral banks: Sedimentary setting, dwelling community, and taphonomy (Late Pliocene, W Mediterranean)

 Cladocora caespitosa is a common zooxanthellate, ahermatypic, constructional scleractinian coral in shallow waters of the present-day Mediterranean. Extensive coral banks in Upper Pliocene shallow marine deposits of the Almería-Níjar Basin (SE Spain) contain the same species. These banks occur on debris-flow conglomerates deposited in a fan delta, or on bioclastic accumulations interpreted as storm deposits. Direct relationships of coral beds with coastal facies indicate that C. caespitosa colonized shallow settings near the paleocoast, probably not deeper than 20–30 m. Low turbulence allowed corals to colonize substrates, which remained stable for long periods. Activity of organisms in the coral community, storms, and detritic discharges from the fan delta were the most significant mechanisms disturbing the coral development. The hard substrata provided by coral banks promoted colonization by cemented and epibyssate organisms. Coral banks marked maximum flooding surfaces at the end of transgressive systems tracks. They were suddenly buried by sediment input into the basin. Taphonomic signatures measured on components of the coral bank communities indicate a low turbulence environment, probably a bay. The low hydraulic energy further inhibited post-burial reworking, thus promoting the in situ preservation of a great part of the organisms inhabiting the bioconstructions. Accepted: 2 December 1997     

Effects of temperature, light and heterotrophy on the growth rate and budding of the temperate coral Cladocora caespitosa

Recent investigations have shown the temperate scleractinian coral Cladocora caespitosa to be a new potential climate archive for the Mediterranean Sea. Whilst earlier studies have demonstrated a seasonal variation in growth rates, they were unable to distinguish which environmental parameter (light, temperature, or food) was influencing growth. In this study, the effect of these three factors on the coral physiology and calcification rate was characterized to aid the correct interpretation of skeletal trace element variations. Two temperatures (13 and 23°C), irradiances (50 and 120 μmol m⁻² s⁻¹), and feeding regimes (unfed and fed with nauplii of Artemia salina) were tested under controlled laboratory conditions on the growth, zooxanthellae density, chlorophyll (chl) content, and asexual reproduction (budding) of C. caespitosa during a 7-week factorial experiment. Unlike irradiance, which had no effect, high temperature and food supply increased the growth rates of C. caespitosa. The effect of feeding was however higher for corals maintained at low temperature, suggesting that heterotrophy is especially important during the cold season, and that temperature is the predominant factor affecting the coral’s growth. At low temperature, fed samples had higher zooxanthellae density and chl content, possibly for maximizing photosynthetic efficiency. Sexual reproduction investment of C. caespitosa was higher during favourable conditions characterised by high temperatures and zooplankton availability.     

The ecology of the Mediterranean stony coral Cladocora caespitosa (Linnaeus, 1767) in the Gulf of Trieste (northern Adriatic Sea): a 30-year long story

Cladocora caespitosa is an endemic coral of the Mediterranean Sea and an important carbonate bioconstructor that adds 3D complexity to the habitat, thus increasing marine biodiversity. Despite its important role in the ecosystem, the real status of the population along most of the Mediterranean coastline is still poorly investigated and very little is known about the resilience of the species. Using non-destructive visual surveys, colonies of C. caespitosa were investigated by SCUBA diving in 2013 and 2015 at seven sites of the northern Adriatic Sea (southern part of the Gulf of Trieste). Data about colony size, index of sphericity and corallite diameter were collected. Almost all biometrical parameters differed significantly among sampling sites, showing low occurrence of the larger size classes compared to the abundance of small-sized colonies. This pattern of distribution is typical of long-lived organisms. The positively skewed colony size distribution could be due to both a high mortality rate of small colonies unable to reach larger size classes, and to a high fragmentation rate of colonies, related to a strong hydrodynamic forces. The northern Adriatic population of C. caespitosa has previously been investigated by Schiller, who reported size and abundance data of colonies from one site, at a depth range of 2–5?m. We compared these data with our findings from the same sampling site, adding new information about the ecology of C. caespitosa. After a 30-year period, the comparison shows a change in the size distribution of colonies, with a decrease of the small class and an increase of the medium class of colonies. In view of these conclusions, further assessments are required in order to evaluate the trend of the northernmost C. caespitosa population in the Mediterranean Sea.     

Long-Term Responses of Mediterranean Mountain Forests to Climate Change,Fire and Human Activities in the Northern Apennines (Italy)

Ecosystems - Fagus sylvatica (beech) dominates the montane forests of the Apennines and builds old-growth high-conservation value stands. However, recent severe drought-induced diebacks raise...     

Divergent Responses of Soil Fungi Functional Groups to Short-term Warming

Soil fungi fill pivotal ecological roles in biogeochemical processes, particularly dominating decomposition of lignin. Little is known, however, about the responses of different fungal groups to climate warming with respect to bacteria. In this study, using barcode pyrosequencing, we showed that short-term (15 months) of field exposure of an alpine meadow to warming (elevated 1 and 2 °C) did not markedly alter the overall soil fungal community structures and α-diversity on Tibetan Plateau, but the average β-diversity dramatically decreased in response to warming. However, soil respiration rates were stimulated in the growing season, which significantly (P?Ascomycota and rare taxa (relative abundance?Basidiomycota-affiliated members significantly increased, while Ascomycota showed a range of responses to warming. Collectively, we conclude that the fungal communities are resistant to short-term warming, though variations are observed in certain species and rare taxa. This report indicates that changes in a relatively small subset of the soil fungal community are sufficient to produce substantial changes in function, such as CO₂ efflux rates.     

The impact of seawater temperature on coral growth parameters of the colonial coral Cladocora caespitosa (Anthozoa, Scleractinia) in the eastern Adriatic Sea

The scleractinian coral Cladocora caespitosa deserves a special place among the major carbonate bioconstructors of the Mediterranean Sea. Annual coral skeleton growth, coral calcification, and skeleton density of the colonial coral C. caespitosa taken from 25 locations in the eastern Adriatic Sea were analyzed and compared with annual sea surface temperatures (SST). The growth rates of the coral C. caespitosa from the 25 stations in the Adriatic Sea ranged from 1.92 to 4.19?mm per year, with higher growth rates of the investigated corallites in the southern part of the Adriatic Sea. These growth rates are similar to those measured in other areas of the Mediterranean Sea. The correlation between coral growth and sea temperatures in the Adriatic Sea is seen as follows: An X-radiograph analysis of coral growth in C. caespitosa colonies that are over 60?years old showed that higher growth rates of this coral coincided with a warmer period in the Mediterranean Sea. A positive significant correlation exists between corallite growth rates and SST and coral calcification and SST. A negative correlation exists between coral density and SST. Coral growth rates also showed a correlation with higher eutrophication caused by nearby fish farms, along with a greater depth of the investigated colonies and high bottom currents.     

Vulnerability of Mediterranean Ecosystems to Long-Term Changes along the Coast of Israel

Although human activity is considered to be a major driving force affecting the distribution and dynamics of Mediterranean ecosystems, the full consequences of projected climate variability and relative sea-level changes on fragile coastal ecosystems for the next century are still unknown. It is unclear how these waterfront ecosystems can be sustained, as well as the services they provide, when relative sea-level rise and global warming are expected to exert even greater pressures in the near future (drought, habitat degradation and accelerated shoreline retreat). Haifa Bay, northern Israel, has recorded a landward sea invasion, with a maximum sea penetration 4,000 years ago, during an important period of urban development and climate instability. Here, we examine the cumulative pressure of climate shifts and relative sea-level changes in order to investigate the patterns and mechanisms behind forest replacement by an open-steppe. We provide a first comprehensive and integrative study for the southern Levant that shows that (i) human impact, through urbanization, has been the main driver behind ecological erosion in the past 4,000 years; (ii) climate pressures have reinforced this impact; and (iii) local coastal changes have played a decisive role in eroding ecosystem resilience. These three parameters, which have closely interacted during the last 4,000 years in Haifa Bay, clearly indicate that for an efficient management of the coastal habitats, anthropogenic pressures linked to urban development must be reduced in order to mitigate the predicted effects of Global Change.     

森林凋落物动态及其对全球变暖的响应

综述了森林凋落物研究的进展,森林凋落物动态的研究随研究方法的改进而不断深化。制约凋落物分解速率的因素有内在因素即凋落物自身的化学物理性质和外在因素即凋落物分解过程发生的外部环境条件,如参与分解的异养微生物和土壤动物群落的种类、数量、活性（生物类因素）和气候、土壤、大气成分等（非生物类因素）。讨论了全球变暖可能引起的凋落物量和凋落物分解的变化。气温上升可能引发植被分布、物候特征和制约凋落物分解因素的改变,影响森林凋落物动态,最终影响森林生态系统物质循环的功能。     

Photoprotective Responses to Winter Stress in Evergreen Mediterranean Ecosystems

Abstract: The role of antioxidants and regulation of photosynthesis as photoprotective adaptations of Mediterranean vegetation to winter stress have been studied. These mechanisms are influenced by the degree of environmental stress (studied at different altitudes) and the taxonomic origin of each species. Thus, antioxidant composition was species-dependent, but not apparently affected by altitude; while the reductions in photochemical efficiency, associated with the presence of zeaxanthin and a lower chlorophyll content, were common among species with increasing altitude. Three different strategies to cope with winter stress were observed in Mediterranean evergreens. First, in Buxus sempervirens antioxidant concentration was very high and the photochemical efficiency was much reduced, with a large accumulation of zeaxanthin. Second, in other Mediterranean sclerophylls, reversible reductions of PSII efficiency related to activity of the xanthophyll cycle allowed day to day adaptation of photosynthetic rates. Third, the summer semideciduous Cistus species maintained a high photochemical efficiency and accumulated little antioxidant, and this strategy could be justified by the occurrence two leaf populations with different physiological tolerances to environmental stress.     

Responses of the Metabolism of the Larvae of Pocillopora damicornis to Ocean Acidification and Warming

Ocean acidification and warming are expected to threaten the persistence of tropical coral reef ecosystems. As coral reefs face multiple stressors, the distribution and abundance of corals will depend on the successful dispersal and settlement of coral larvae under changing environmental conditions. To explore this scenario, we used metabolic rate, at holobiont and molecular levels, as an index for assessing the physiological plasticity of Pocillopora damicornis larvae from this site to conditions of ocean acidity and warming. Larvae were incubated for 6 hours in seawater containing combinations of CO₂ concentration (450 and 950 µatm) and temperature (28 and 30°C). Rates of larval oxygen consumption were higher at elevated temperatures. In contrast, high CO₂ levels elicited depressed metabolic rates, especially for larvae released later in the spawning period. Rates of citrate synthase, a rate-limiting enzyme in aerobic metabolism, suggested a biochemical limit for increasing oxidative capacity in coral larvae in a warming, acidifying ocean. Biological responses were also compared between larvae released from adult colonies on the same day (cohorts). The metabolic physiology of Pocillopora damicornis larvae varied significantly by day of release. Additionally, we used environmental data collected on a reef in Moorea, French Polynesia to provide information about what adult corals and larvae may currently experience in the field. An autonomous pH sensor provided a continuous time series of pH on the natal fringing reef. In February/March, 2011, pH values averaged 8.075±0.023. Our results suggest that without adaptation or acclimatization, only a portion of naïve Pocillopora damicornis larvae may have suitable metabolic phenotypes for maintaining function and fitness in an end-of-the century ocean.