Latitudinal diversity patterns of continental benthic copepod species assemblages in the Americas

The diversity, considered as richness (numbers) of genera and species, of continental benthic Copepoda of the Americas was examined. Thirty-three local lists of planktonic and benthic Cyclopidae (Eucyclopinae and Cyclopinae), 21 local lists of epibenthic Harpacticoida (Canthocamptidae), and 8 local lists of interstitial Harpacticoida (Parastenocarididae) were compared. Available data do not allow the rejection of the null hypothesis of no significant difference in diversity of tropical and temperate copepod faunas. For Cyclopidae, 6–15 genera and 6–44 species were recorded from local areas; linear regression analysis showed no significant correlation between latitude and number of genera and species. For Canthocamptidae, 1–13 genera and 3–43 species were recorded locally, with no significant correlation between latitude and number of species; for genera the correlation between latitude and number of genera was strongly positive, but the effect of latitude is small. No relationship between latitude and diversity of the Parastenocarididae was evident from inspection. Endemism of Cyclopidae and Canthocamptidae is significantly greater in South America, as shown by comparisons of Sørensen's similarity coefficient between paired areas. Because of this high degree of endemism, conservation strategies for South American wetland invertebrates should emphasize preservation of many small sites.     

Hypoxia tolerance in the copepod Calanoides carinatus and the effect of an intermediate oxygen minimum layer on copepod vertical distribution in the northern Benguela Current upwelling system and the Angola-Benguela Front

There is a growing concern that hypoxic and anoxic areas in the sea spread in extent and intensity, posing a severe risk to marine ecosystems and fisheries. Hypoxia may affect fish stocks directly or via detrimental effects on important prey species, such as zooplankton. A unique feature of the northern Benguela Current upwelling region and Angola-Benguela frontal system is a pronounced intermediate oxygen minimum layer (IOML) at 60-500 m depth with oxygen concentrations ≤ 1.4 mg O₂l^− 1 (minimum < 0.7 mg O₂l^− 1). Field studies during February-March 2002 demonstrated that the abundance of calanoid copepods and the biomass of mesozooplankton in general were severely reduced within the IOML. The dominant copepod Calanoides carinatus showed a bimodal vertical distribution with parts of the population either comprising all developmental stages concentrated in the surface layer (0-60 m), or copepodids C5 diapausing below 400 m depth apparently avoiding the IOML. Accordingly, abundances of other calanoid copepods were higher at the surface and below 300 m than in the centre of the IOML. The scarcity of planktonic life within the IOML raises the question whether this layer represents an effective barrier for zooplankton vertical migrations. Especially in C. carinatus, ontogenetic vertical migration plays a key role in the retention of the population within the productive upwelling region and for the rapid re-colonisation of plumes of newly upwelled water. To address this issue, the hypoxia tolerance of C. carinatus was determined in a series of laboratory-based, closed-bottle experiments in January 2005. Copepods were kept in gas-tight bottles and the decreasing oxygen concentrations were monitored to establish their minimum oxygen demands. Although copepodids survived apparently unharmed at surprisingly low oxygen concentrations of ca. 1.5 mg O₂l^− 1, they could not tolerate oxygen levels < 1.1 mg O₂l^− 1, implying that the core of the IOML, where O₂ concentrations are below this threshold, is uninhabitable for C. carinatus. In contrast, the IOML may represent a refuge from competition and predation for other copepod species specifically adapted to hypoxic environments.     

Copepod communities related to water masses in the southwest East China Sea

The East China Sea is characterized by a complex hydrographic regime and high biological productivity and diversity. This environmental setting in particular challenged a case study on the use of mesozooplankton community parameters as indicators of water masses. In order to reveal spatial patterns of zooplankton communities during summer, a large scale oceanic transect study was conducted. Two transects were taken in the southwest East China Sea region, covering for the first time the China shelf, slope, and the estuaries of the Yangtze river and of the Minjiang river, the northern Taiwan Strait, and the Kuroshio Current region. A total of 77 copepod species were quantified. Copepod abundance was significantly higher in the estuary of the Yangtze River runoff mixture waters and lowest at the Kuroshio Current Region. The calanoid Parvocalanus crassirostris was the most frequently occurring and abundant species retrieved from 27 samples of a total of 39 samples. The use of multivariate cluster analysis separated the Mainland China Shelf from the northern Taiwan Strait and the Kuroshio Current Region at the first hierarchical level. The use of an indicator value method (IndVal) associated with each cluster of stations revealed characteristic species assemblages. Two hierarchical levels defined 4 assemblages within geographical sectors representing copepod assemblages of the Kuroshio Current Region, of the northern Taiwan Strait and the southern China Shelf near the estuary of the Minjiang River and northern stations near the estuary of the Yangtze River. Overall, there was a strong correspondence between the distribution of certain copepod species and water masses. Differences between the Mainland China shelf, the northern Taiwan Strait and the Kuroshio Current Region were characterized by differences in species composition and abundance. Water mass boundaries in the study area were exclusively indicated by distinct differences in species composition, emphasizing a correlation between copepod communities and water masses of the southwest East China Sea in summer.     

Distribution of echinoderms and their larvae in the southwestern Peter the Great Bay, Sea of Japan

Based on results of processing planktonic and benthic samples collected in 1996 and 1997, a spatial distribution of echinoderms in the bottom and of their larvae in the plankton were collated for the water area of the Southern region of the Far Eastern State Biosphere Marine Reserve. Some correlation between distributions of the adult and larval sea star Asterias amurensis in July was revealed. At the same time, there was no correlation between distributions of larvae and adult individuals of the brittle star Ophiura sarsi and sea urchin Echinocardium cordatum, which are most abundant in the area. The size structure of bottom populations of the brittle stars O. sarsi and Amphiodia fissa in the studied area was assessed. The correlation coefficient between the distribution of young-of-the-year and the population density was 0.47 in O. sarsi and 0.74 in A. fissa respectively, which implied a selective settling of larvae of those species in areas inhabited by adult brittle stars. Recruitment of bottom populations from 1996 spawning was 5% in O. sarsi and 3.3% in A. fissa.     

Changes in community structure of active protistan assemblages from the lower Pearl River to coastal Waters of the South China Sea

Protists make up an important component of aquatic ecosystems, playing crucial roles in biogeochemical processes on local and global scales. To reveal the changes of diversity and community structure of protists along the salinity gradients, community compositions of active protistan assemblages were characterized along a transect from the lower Pearl River estuary to the open waters of the South China Sea (SCS), using high-throughput sequencing of the hyper-variable V9 regions of 18S rRNA. This study showed that the alpha diversity of protists, both in the freshwater and in the coastal SCS stations was higher than that in the estuary. The protist community structure also changed along the salinity gradient. The relative sequence abundance of Stramenopiles was highest at stations with lower salinity and decreased with the increasing of salinity. By contrast, the contributions of Alveolata, Hacrobia and Rhizaria to the protistan communities generally increased with the increasing of salinity. The composition of the active protistan community was strongly correlated with salinity, indicating that salinity was the dominant factor among measured environmental parameters affecting protistan community composition and structure.     

Prokaryotic assemblages and metagenomes in pelagic zones of the South China Sea

Background

Prokaryotic microbes, the most abundant organisms in the ocean, are remarkably diverse. Despite numerous studies of marine prokaryotes, the zonation of their communities in pelagic zones has been poorly delineated. By exploiting the persistent stratification of the South China Sea (SCS), we performed a 2-year, large spatial scale (10, 100, 1000, and 3000 m) survey, which included a pilot study in 2006 and comprehensive sampling in 2007, to investigate the biological zonation of bacteria and archaea using 16S rRNA tag and shotgun metagenome sequencing.

Results

Alphaproteobacteria dominated the bacterial community in the surface SCS, where the abundance of Betaproteobacteria was seemingly associated with climatic activity. Gammaproteobacteria thrived in the deep SCS, where a noticeable amount of Cyanobacteria were also detected. Marine Groups II and III Euryarchaeota were predominant in the archaeal communities in the surface and deep SCS, respectively. Bacterial diversity was higher than archaeal diversity at all sampling depths in the SCS, and peaked at mid-depths, agreeing with the diversity pattern found in global water columns. Metagenomic analysis not only showed differential %GC values and genome sizes between the surface and deep SCS, but also demonstrated depth-dependent metabolic potentials, such as cobalamin biosynthesis at 10 m, osmoregulation at 100 m, signal transduction at 1000 m, and plasmid and phage replication at 3000 m. When compared with other oceans, urease at 10 m and both exonuclease and permease at 3000 m were more abundant in the SCS. Finally, enriched genes associated with nutrient assimilation in the sea surface and transposase in the deep-sea metagenomes exemplified the functional zonation in global oceans.

Conclusions

Prokaryotic communities in the SCS stratified with depth, with maximal bacterial diversity at mid-depth, in accordance with global water columns. The SCS had functional zonation among depths and endemically enriched metabolic potentials at the study site, in contrast to other oceans.

Electronic supplementary material

The online version of this article (doi:10.1186/s12864-015-1434-3) contains supplementary material, which is available to authorized users.     

南海北部表层沉积物中原核微生物多样性

[目的]为研究南海北部沉积物原核微生物的多样性和群落结构.[方法]从南海北部XSCS13站位表层沉积物中扩增古菌和细菌的16S rDNA并构建文库,随机挑出阳性克隆子进行测序,选出所有的OTU构建系统进化树,进行系统发育学分析.[结果]多数克隆子来自于未培养原核微生物,沉积物中的古菌分属3大门类:泉古菌(Crenarchaeota)、奇古菌(Thaumarchaeota)和广古菌(Euryarchaeota),其中泉古菌(Crenarchaeota)为主要门类,占71％;广古菌(Euryarchaeota)最少,只有3个克隆子.泉古菌(Crenarchaeota)又以MG Ⅰ为主要类群,占61％.细菌多样性明显高于古菌,共9个门类:变形杆菌(Proteobacteria)(32.6％)、疣微菌(Verrucomicrobia)(3.0％)、拟杆菌门(Bacteroidete)(5.2％)、酸杆菌(Acidobacteria)(4.4％)、绿弯菌(Chloroflexi)(6.0％)、厚壁菌(Firmicute)(3.7％)、浮霉菌(Planctomycete)(5.2％)、芽单胞菌(Gemmatimonadete)(11.1％)、放线细菌(Actinobacteria)(4.4％).变形杆菌为优势类群(包括α-Proteobacteria、γy-Proteobacteria和δ-Proteobacteria 3个亚群),其中γ-Proteobacteria是Proteobacteria中的优势种群,占54.5％.另外所有原核微生物总共有超过50％的克隆子与硫酸盐的还原以及甲烷的形成相关.[结论]结果表明南海北部XSCS13站位表层沉积物中原核微生物的多样性非常丰富,其中蕴含大量未知的微生物资源;另外古菌和细菌群落结构表明该位点可能处于富含甲烷的冷泉活动区.     

A 5-year study of the influence of the northeast and southwest monsoons on copepod assemblages in the boundary coastal waters between the East China Sea and the Taiwan Strait

Danshuei River in the northwestern tip of Taiwan dischargesinto the boundary coastal waters between the East China Seaand the Taiwan Strait. We conducted a 5-year study between October1998 and September 2003 to assess the influence of the northeast(NE) and the southwest (SW) monsoons on copepod assemblagesin the area. A total of 110 copepod species were identified.Temora turbinata, Paracalanus aculeatus, Acrocalanus gibber,Parvocalanus crassirostris and Oithona rigida comprised 80%of the copepod assemblages that consisted of coastal speciesfrom the East China Sea and species associated with the KuroshioBranch Current. The effect of the NE monsoon was observed duringa short period of each winter when species such as Calanus sinicuswere transported into the study area by the China Coastal Current.In summer, species such as A. gibber, Acrocalanus gracilis andCanthocalanus pauper may be transported into the study areafrom Southern Taiwan by the combined effect of the SW monsoonand the Kuroshio Current. Influence of the NE monsoon on thecopepod assemblages in terms of introduction of species fromthe Yellow Sea and the East China Sea during winter was observedevery year but was unlikely to represent an important carbonsource into the region due to its short-term nature. Local speciessuch as T. turbinata, P. crassirostris and O. rigida form amajor component of the copepod assemblage. The high copepoddiversity in the area was caused by the year-round presenceof many local species and the aggregation of different speciesfrom subtropical, tropical, and temperate water masses. Monsoon-drivenwater currents and the Kuroshio Branch Current appeared to maskthe effect of river discharge in the region.     

Spatial structure of eukaryotic ultraplankton community in the northern South China Sea

Spatial distribution, diversity and composition of eukaryotic ultraplankton community of the northern South China Sea (nSCS) surface water and the relationship with the in situ water environment were investigated by the method of polymerase chain reaction-denaturing gradient gel electrophoresis (PCR-DGGE). A total of 18 DGGE intensive bands were detected and the sequence analysis of these DGGE bands revealed that Alveolata was the dominant eukaryotic ultraplankton group of surface water in the nSCS (50%). Other species belonged to Bicoecea, Bolidophyceae, Polycystinea and Chlorophyta, which accounted for less proportion of eukaryotic ultraplankton in the study area. Unweighted pair group method with arithmetic averages clustering of the sampling stations indicated that all stations were classified mainly based on geographical proximity. Redundancy analysis (RDA) was employed to further investigate the relationships between DGGE band pattern and the environmental variables. Based on the RDA analysis, temperature, salinity, phosphorus and silicate were the important factors to shape the eukaryotic ultraplankton community composition in the nSCS.     

Spatial and temporal distribution of nanoflagellates in the northern South China Sea

Seasonal variation, horizontal and vertical distribution, and cell size of nanoflagellates, together with physico-chemical and biological factors, were studied in the northern South China Sea (SCS). It was found that nanoflagellate abundance ranged from 0.157 × 10³ to 9.193 × 10³ cells/ml (with a mean of 0.891 × 10³) in winter (February, 2004), while it ranged from 0.107 × 10³ to 5.417 × 10³ cells/ml (with a mean of 0.599 × 10³) in summer (July, 2004). Nanoflagellates were more abundant in winter than summer in offshore regions, showing an unique seasonal pattern in this subtropical marginal sea. The abundance of nanoflagellates decreased from the estuary to the offshore region. Vertical distribution of nanoflagellates coupled well with that of bacteria and Chl a. The small size fraction of less than 5 μm dominated the nanoflagellate populations. Wind-driven mixing, eddies, availability of nutrients as well as Chl a and abundance of picoplankton seemed to be the major controlling factors for the spatial distribution and seasonal variation of nanoflagellates in the study area. Handling editor: P. Tasman     

Distribution and Diversity of Microbial Eukaryotes in Bathypelagic Waters of the South China Sea

Little is known about the biodiversity of microbial eukaryotes in the South China Sea, especially in waters at bathyal depths. Here, we employed SSU rDNA gene sequencing to reveal the diversity and community structure across depth and distance gradients in the South China Sea. Vertically, the highest alpha diversity was found at 75‐m depth. The communities of microbial eukaryotes were clustered into shallow‐, middle‐, and deep‐water groups according to the depth from which they were collected, indicating a depth‐related diversity and distribution pattern. Rhizaria sequences dominated the microeukaryote community and occurred in all samples except those from less than 50‐m deep, being most abundant near the sea floor where they contributed ca. 64–97% and 40–74% of the total sequences and OTUs recovered, respectively. A large portion of rhizarian OTUs has neither a nearest named neighbor nor a nearest neighbor in the GenBank database which indicated the presence of new phylotypes in the South China Sea. Given their overwhelming abundance and richness, further phylogenetic analysis of rhizarians were performed and three new genetic clusters were revealed containing sequences retrieved from the deep waters of the South China Sea. Our results shed light on the diversity and community structure of microbial eukaryotes in this not yet fully explored area.     

Distribution of polycystine radiolarians in the northern South China Sea in September 2005

We studied the distribution of polycystine radiolarians in the northern South China Sea (SCS) in September 2005 in order to document the relation between radiolarian assemblages and environmental parameters. Vertical plankton tows were collected at depth-intervals from 0 to 250 m at thirteen stations, using a closing net (62 µm mesh). The highest abundance of polycystine radiolarians generally occurred at depths of 0–75 m, i.e., above the chlorophyll-a maximum and thermocline. In contrast, the maximum polycystine abundance occurred below the chlorophyll-a maximum at station E105 (75–150 m), possibly as a result of down-welling of warm waters, and the maximum fell within the chlorophyll-a maximum at station E702, as a result of upwelling of intermediate cold waters.During our sampling period, the southwestern monsoon caused upwelling in the northern SCS (the Taiwan Bank region), as shown by the high abundance of Tetrapyle octacantha, Acanthodesmia vinculata, Cladococcus cervicoris, Octopyle octospinosa, Spongaster tetras, Streblacantha circumtexta and Spongotrochus glacialis. The patterns of distribution and the abundances of the species Botryocyrtis scutum, Pterocorys hertwigii, Collosphaera tuberosa and Didymocyrtis tetrathalamus tetrathalamus suggested that western Equatorial Pacific waters did not yet influence the region at the time of sampling; the first two species were dominant in the northern SCS during our sampling. The species Cornutella profunda and Cyrtopera laguncula were rare at depths of 150–250 m in the tropical region of the SCS, and assemblages in northern and southern SCS were very similar. The relative abundance of B. scutum and P. hertwigii is an indicator for seasonal changes in monsoonal circulation.     

Seasonal and interannual variation of subtidal fish assemblages in Wakasa Bay with reference to the warming trend in the Sea of Japan

A bi-monthly underwater visual census was conducted in the subtidal habitat of Nagahama, Wakasa Bay, Sea of Japan, over 5 years from January 2002 to December 2006. A total of 83 fish species and 63,328 individuals were recorded over 120 1-h visual censuses composed of three 2 × 200 m transects. The number of both total individuals (abundance) and species (richness) were strongly correlated with the bottom-water temperature. Species richness was greatest in summer (July–September), with 10–20 species per 400 m², when the sea-bottom temperature was highest (24–28°C), and lowest in winter (January–March), with 1–8 species per 400 m², when the temperature was lowest (10–12°C). Five species, i.e., rockfish, Sebastes inermis, wrasse, Pseudolabrus sieboldi, two species of goby, Acentrogobius pflaumii and Tridentiger trigonocephalus, and pygmy filefish, Rudarius ercodes, were frequently observed and present in more than 200 of the total of 360 total transects. While the presence of most fish species was dependent on water temperature, some species were independent of water temperature, including anchovy, Engraulis japonicus, S. inermis, wasp fish, Hypodytes rubripinnis, A. pflaumii and pufferfish, Takifugu poecilonotus. Jack mackerel, Trachurus japonicus, was the most abundant fish, being most typically seen from spring through autumn, with a total of 21,850 individuals observed over the whole census period. Some species showed strong annual fluctuations in abundance. The interannual stability in fish abundance and species richness was probably because the fish assemblage was composed of a sufficiently large number of species, but with the number of each species fluctuating over time. A comparison of the present survey to one in 1970–72 revealed that over the last 30 years southern fish species have significantly increased whereas those indigenous to northern waters have decreased.

南海北部浮游植物生长对营养盐的响应

2004年夏季作者在南海北部海域研究了浮游植物生长的营养动力学,结合物理-化学过程对浮游植物生物量分布的影响与机制进行了研究,阐明了水平对流和中尺度涡对营养盐分布的影响及浮游植物生长和现存生物量对其的响应。受西南季风和东向沿岸流作用所形成的Ekman输送的影响,南海北部海岸带表层海水作离岸运动,使深层富含营养盐的冷水爬坡涌升到表层来补充,激发浮游植物生物量迅速增长。海区反气旋涡使海水辐聚下沉,造成水体具高温、低盐、高溶解氧浓度、低营养盐浓度和低浮游植物生物量。同时通过现场营养盐加富试验,发现该海域营养盐是浮游植物生长的主要限制因子,而且是多种营养元素共同限制了浮游植物的生长,添加单一的营养盐并不能促进浮游植物的生长。在生物量出现增长的试验组中,营养盐添加不仅促使浮游植物生物量的增长,而且也改变了浮游植物的粒级结构和群落结构。例如,在站S1008,培养前叶绿素a浓度为0.28 mg.m-3,加富培养60 h后浮游植物生物量在NP和NPSi的试验组中有显著的增加,叶绿素a浓度分别达1.07 mg.m-3和1.19 mg.m-3;培养前粒度分级叶绿素a主要以Pico级份占优势,而加富试验结束后,在NP和NPSi的试验组以Nano级份占优势,其它试验组仍以Pico级份占优势;同时,在培养后生物量出现增长的试验组,浮游植物群落的优势类群从甲藻向硅藻演替。     

夏季南海北部纤毛虫群落组成及其水平分布

在2007年中国科学院南海海洋研究所南海北部开放航次期间,设计了两条纤毛虫采样断面(X断面、Y断面),共13个站位,涵盖了珠江冲淡水区、粤东上升流区和陆坡开阔海区,旨在对纤毛虫的种类组成和空间分布及其影响因素进行探讨。共检出4纲8目22属38种纤毛虫,其中种类数最多的属是砂壳目拟铃虫属(9种),其次是急游目急游属(5种)。本航次中纤毛虫主要是砂壳目纤毛虫(27种),共鉴定出砂壳虫15属27种。纤毛虫优势种为蚤状中缢虫Mesodinium pulex(18.1%),其次是丁丁急游虫Strombidium tintinnodes(9.7%),盾形拟铃虫Tintinnopsis urula(5.8%)。纤毛虫种类数和丰度从近岸向外海逐渐减少,纤毛虫种类数与温度(R2=0.53,P0.05)和盐度(R2=0.43,P0.05)呈负相关,随着温度的升高,纤毛虫种类数逐渐降低。纤毛虫丰度与温度和盐度相关性并不明显,这可能由于受上升流的影响,近岸上升流高盐区存在着纤毛虫丰度较为丰富的情况。叶绿素a浓度与纤毛虫种类数(R2=0.36,P0.05)和纤毛虫丰度(R2=0.36,P0.05)呈正相关,近岸浮游植物生物量高,纤毛虫种类丰富,丰度大,叶绿素a沿纵断面降低,纤毛虫种类数和丰度也降低。     

夏、冬季南海北部浮游植物群落特征

对2009年7月19日—8月16日和2010年1月6—30日南海北部(18°—23.5°N、109°—120°E)两个航次的浮游植物样品应用Utermhl方法进行了分析鉴定。结果如下:夏季样品鉴定浮游植物4门72属150种,浮游植物细胞丰度范围为(0.16—6001.78)×103个/L,平均细胞丰度为26.49×103个/L,硅藻的平均细胞丰度为25.81×103个/L,主要优势种属有铁氏束毛藻(Trichodesmium thiebautii)、菱形海线藻(Thalassionema nitzschioides)、柔弱伪菱形藻(Pseudo-nitzschia delicatissima)及裸甲藻(Gymnodinium spp.)等;冬季样品鉴定浮游植物4门58属168种,浮游植物细胞丰度范围为(0.08—37.52)×103个/L,浮游植物平均细胞丰度为2.69×103个/L,硅藻的平均细胞丰度为2.49×103个/L,主要优势物种为菱形海线藻、伏氏海线藻(Thalassionema frauenfeldii)及短刺角毛藻(Chaetoceros messanensis)等;夏季调查区5m层浮游植物细胞丰度的平面分布由近岸到外海迅速减少,高值区主要在广东东部近岸及海南东部近岸;冬季则在珠江口近岸和海盆区出现较高值。两次调查中,浮游植物细胞丰度在浅水站位(200m)远高于深水站位(200m);较冬季相比,夏季浮游植物平均细胞丰度偏高,但物种丰富度却略偏低。夏、冬季浮游植物的香农-威纳多样性指数平均值分别为2.12和2.36,Pielou均匀度指数平均值分别为0.79和0.81。两个航次调查中,浮游植物细胞丰度均与盐度表现出显著性负相关性;在冬季还与磷酸盐浓度表现出显著性负相关性。     

Year-to-year variation in larval fish assemblages of the Southern North Sea

In order to test the temporal stability within and the reproducibility of larval fish assemblages between years, the larval fish assemblage at Helgoland Roads, North Sea (NE Atlantic) was quantitatively sampled almost daily from January 2003 to December 2005. The survey resulted in a total of 462 samples containing 50,632 larval fish of at least 42 taxa. In winter the larval fish assemblage was mainly dominated by larvae emerging from demersal eggs. This changed gradually to larvae hatching from pelagic eggs. Larvae from pelagic eggs dominated the ichthyoplankton assemblage in summer. A remarkably stable seasonality in terms of dominance patterns with recurring, season-specific fish assemblages was observed over the 3 years, despite substantial variation in environmental conditions such as a temperature difference of almost 20°C between summer and winter. The lesser sandeel (Ammodytes marinus), was the only species which showed significant fluctuations in abundance between the years. After removal of this species from the analysis, the dominance patterns of the remaining fish species were almost identical between years.     

Mapping biodiversity in three-dimensions challenges marine conservation strategies: The example of coralligenous assemblages in North-Western Mediterranean Sea

Multi-facet diversity indices have been increasingly widely used in conservation ecology but congruence analyses both on horizontal and vertical axes have not yet been explored. We investigated the vertical and horizontal distributions of α and β taxonomic (TD), functional (FD) and phylogenetic diversity (PD) in a three-dimensional structured ecosystem. We focused on the Mediterranean coralligenous assemblages which form complex structures both vertically and horizontally, and are considered as the most diverse and threatened communities of the Mediterranean Sea. Although comparable to tropical reef assemblages in terms of richness, biomass and production, coralligenous assemblages are less known and more rarely studied, in particular because of their location in deep waters. Our study covers the entire range of distribution of coralligenous habitats along the French Mediterranean coasts, representing the most complete database so far developed for this important ecosystem. To our knowledge, this is the first analysis of spatial diversity patterns of marine biodiversity on both horizontal and vertical scales.Our study revealed that taxonomic diversity differed from functional and phylogenetic diversity patterns at the station level, the latter two being strongly structured by depth, with shallower stations generally richer than deeper ones. Considering all stations, phylogenetic diversity was less congruent to taxonomic diversity (Pearson's correlation of r = 0.48) but more congruent to functional diversity (r = 0.69) than randomly expected. Similar congruence patterns were revealed for stations deeper than 50 m (r = 0.44 and r = 0.84, respectively) but no significantly different congruence level than randomly expected was revealed among diversity facets for more shallow stations. Mean functional α- and β-diversity were lower than phylogenetic diversity and even lower than taxonomic α- and β-diversity for both vertical and horizontal scales. Low FD and PD values at both α- and β-diversity indicated functional and phylogenetic clustering. Community dissimilarities (β-diversity) increased over depth especially in central and eastern part of the French Mediterranean littoral and in northern Corsica, indicating coralligenous vertical structure within these regions. Overall horizontal β-diversity was higher within the 50–70 m depth belts.We conclude that taxonomic diversity alone is inadequate as a basis for setting conservation goals for this ecosystem and additional information, at least on phylogenetic diversity, is needed to preserve the ecosystem functioning and coralligenous evolutionary history. Our results highlight the necessity of considering different depth belts as a basis for regional scale conservation efforts. Current conservation approaches, such as the existing marine protected areas, are insufficient in preserving coralligenous habitats. The use of multi-facet indices should be considered, focusing on preserving local diversity patterns and compositional dissimilarities, both vertically and horizontally.     

Due Diligence and Overlooked Evidence in the South China Sea Arbitration: A Note

Abstract

In the South China Sea Arbitration, the Tribunal decided that China had not breached the due diligence obligation to protect and preserve the marine environment under Articles 192 and 194(5) of the United Nations Convention on the Law of the Sea concerning Chinese fishers fishing with explosives, but that China had breached the same obligation regarding Chinese fishers harvesting endangered species. This article looks at how the Tribunal interpreted and applied the due diligence obligation and argues, from a Chinese perspective, that there were facts overlooked by the Tribunal that China could have presented to counter the evidence of the Philippines, which might have been enough to affect the decision on destructive fishing had China participated in the Arbitration.