Genetic heterogeneity in populations of the Mediterranean shore crab, Carcinus aestuarii (Decapoda,Portunidae), from the Venice Lagoon

Heterogeneity in genetic composition among recruits, mostly due to a large variance in reproductive success mediated by oceanographic processes, has been reported for marine species but is less understood in coastal lagoons' organisms. Temporal genetic variation in natural populations of the Mediterranean shore crab Carcinus aestuarii was quantified over a multi-year sample. A total of 486 adult crabs were collected at eight different sites of the Venice Lagoon during the period 2005–2007 and screened for genetic variation using 11 microsatellite loci. Two additional samples (N = 115) from neighbouring sites, located approximately 100 km North and South to the Venice Lagoon, were included for the sake of comparison. Our results show significant differences in allelic frequencies at the micro-geographic scale of the Venice Lagoon, observed between sites of collection, typologies of habitat, and areas with different class of ecological risk or pattern of hemocyanin expression. However, this pattern was not constant between years, with significant differences observed mainly in 2005 and 2006, but not in 2007. Our results indicate significant temporal differences suggesting the existence of dynamic processes that act on the genetic pool of this species. Although natural selection and gene flow might play a role, we suggest that genetic drift linked to large variation in the reproductive success of individuals is the most probable scenario to explain the local genetic patterns of differentiation in the Mediterranean shore crab. Our study, by providing the first evidence for the existence of genetic differences in this species at the micro-geographic scale, suggests that a better comprehension of the link between reproduction, recruitment and oceanography is critical to understand how colonization and maintenance of genetic variation is achieved in ephemeral and vulnerable environments such as coastal lagoons.     

Differences in the neighborhood: Structural variations in the carapace of shore crabs Carcinus maenas (Decapoda: Portunidae)

The present study compared the carapace structure of Carcinus maenas in two nearby sites (2 km apart) within Minho estuary, submitted to different physicochemical and ecological conditions (water temperature, pH, crabs’ density and sex ratio). The carapace structure of the carapace and chelae of the crabs presented significant differences between sampling sites (t-test; p < 0.01). The SIMPER analysis revealed that the Weight/CW and Thickness/CW ratios explained all the dissimilarities found among sites. Overall, the male carapace was proportionally thicker at station 2 (t-test; p < 0.01), while the female carapace was proportionally thicker at station 1 (t-test; p < 0.001). A thicker carapace can be advantageous when competing for food or a sexual partner. We hypothetized that, since at station 2, the density of individuals was twice higher than at station 1, it is likely that agonistic encounters are more frequent, thus favouring a thicker carapace.     

Polycyclic aromatic hydrocarbons in surface seawater and in indigenous mussels (Mytilus galloprovincialis) from coastal areas of the Saronikos Gulf (Greece)

Polycyclic aromatic hydrocarbons (PAHs) were identified and measured in surface seawater and in the tissues (gills and mantle) of indigenous black mussels, Mytilus galloprovincialis, collected from three coastal sites of Saronikos Gulf (Greece), a gulf that exhibits high levels of pollution. The total PAHs measured by spectrofluorometry in the surface seawater were found in the range of 425–459 ng L⁻¹ at the most polluted sites 1 and 2 (Elefsis Bay–Salamis Island) and in the range of 103–124 ng L⁻¹ at site 3 (Aegina Island). PAHs' sources in seawater were identified by application of specific PAH ratios, such as phenanthrene/anthracene and fluoranthene/pyrene. Levels of PAHs in soft tissues (gills and mantle) of indigenous mussels were much higher than those reported for seawater. Total PAH concentrations in mantle tissues were in the range of 1300–1800 ng g⁻¹ dry weight (dw) tissue at sites 1 and 2 and approximately 380 ng g⁻¹ dw at site 3. In gill tissues total PAH concentrations were in the range of 1480–2400 ng g⁻¹ dw at sites 1 and 2 and approximately 430 ng g⁻¹ dw at site 3. PAHs composition was dominated by two-, three- and four-ring compounds in seawater, where 17 different PAH compounds were identified and measured in mussel tissues. Mussels can be used as sentinel organisms to monitoring PAHs' contamination, since they concentrate PAHs from the surrounding water media and therefore making the chemical analysis simpler and less prone to error than that for water. In surface seawater possible weathering and photodegradation due to hot climates contribute to reduced PAHs concentrations.     

中国海域南褐虾属种类记述(甲壳动物亚门, 十足目, 褐虾科)

为了搞清中国海域存在的南褐虾(Philocheras)的生物多样性和地理分布,作者通过形态学特征系统研究了中国科学院海洋研究所海洋生物标本馆馆藏的南褐虾标本,此标本主要采自中国海域,也检查了部分采自菲律宾海域的标本。本研究和相关文献的结果表明:中国海域存在的南褐虾共计7种,包括作者描绘的4种以及在其他文献中提到的3种。作者对本次研究的4种南褐虾提供了详细的鉴别特征、地理分布、外部整体图和附肢结构图,并对中国海域存在的7种南褐虾提供了检索表。     

Lifespan mercury accumulation pattern in Liza aurata: Evidence from two southern European estuaries

Mercury accumulation throughout the lifespan of Liza aurata (Risso, 1810) was analysed in four tissues (muscle, gills, liver and brain) in two southern European coastal ecosystems with distinct mercury contamination. Specimens from four to five age classes were captured in two sampling sites in the Ria de Aveiro (Laranjo bay and Mira), a system historically contaminated by industrial mercury, and in one site in the Mondego estuary, assumed as a mercury-free ecosystem. Mercury concentration in all tissues was found to be significantly higher in the Ria de Aveiro (Laranjo bay) compared to the Mondego, in accordance with the environmental contamination (water, sediments and suspended particulate matter). Significant differences inside the Ria de Aveiro (between the Mira and Laranjo bay) were only detected in the liver. This tissue registered the highest levels of mercury (ranging from 0.11 to 4.2 μg g⁻¹) in all sampling sites, followed by muscle, brain, and gills. In all sampling sites and tissues was denoted a mercury dilution pattern along the lifecycle (except in liver at the Mondego, the reference area where the concentrations are always very low). An exponential trend was found in the metal age variation patterns in Laranjo (the most contaminated area) and a linear trend in the Mira and the Mondego (the least contaminated areas). Organic mercury concentration in muscle generally accounted for over 95% of total mercury concentration, and followed the same accumulation pattern of total mercury. This fish species is of lesser importance in mercury transfer to adjacent coastal areas and although the consumption of fish from Laranjo may present some risk for the humans, this risk decreases with fish age/size.     

中国东海和南海4个真虾(甲壳动物亚门:十足目:真虾下目)新记录种

采集自中国东海和南海的4个真虾物种,经研究发现为中国海域的新记录物种,即凯氏鞭腕虾（Lysmata kempi Chace,1997）,利普克鞭腕虾（Lysmata lipkei Okuno&Fiedler,2010）;透明近绿虾（Chlorocurtis jactans（Nobili,1904））和黑斑活额虾（Rhynchocinetes conspiciocellus Okuno&Takeda,1992）。本文描述了这四个物种的形态特征,编制了检索表,并对其分类地位进行了讨论。     

Reproductive biology and seasonality of the Indo-Australasian mysid Mesopodopsis orientalis (Crustacea: Mysida) in a tropical mangrove estuary,Malaysia

A year-round survey of the tropical shallow-water mysid Mesopodopsis orientalis (Tattersall, 1908) (Crustacea, Mysidacea) was conducted in the Merbok mangrove estuary, northwestern Peninsular Malaysia. The mysid formed dense aggregations at the river's edge close to the mangrove forest during the daytime, but very few were captured elsewhere in the estuary system. The sampled population was found in a wide range of salinities from 16 to 32, demonstrating broad euryhalinity, and the number of the catch at the littoral zone ranged from 11.8 to 2273 ind m⁻². The overall annual mean was 709.2 ind m⁻². Females predominated over males in the entire population, and brooding females were present at every monthly sample, indicating that reproduction is continuous year round. The clutch size positively correlated with female body length. The diameter of eggs (Stage I embryos) was unaffected by the seasonality and independent of the maternal size within an observed size range. The life history pattern of the estuarine population of M. orientalis showed close similarity to that of the coastal counterpart. However, the former was found to produce fewer but larger eggs, and the specimens in this population were larger than those in the coastal population at the embryo, juvenile, and adult stages. This evidence indicates that the life history features of the estuarine population would differ to some degree from those of the coastal counterpart.     

Negative effects of blue mussel (Mytilus edulis) presence in eelgrass (Zostera marina) beds in Flensborg fjord,Denmark

The effect of blue mussel (Mytilus edulis) presence in eelgrass (Zostera marina) beds was studied from June 2004 to July 2005 in Flensborg fjord, Denmark. The field experiments were conducted at two stations, one with only Z. marina (Eelgrass station) present and one where M. edulis were present in the Z. marina beds (Mixed station). Zostera marina parameters were measured (growth of leaves, shoot density, leaf length, and nutrient content) in combination with epiphyte cover and sediment parameters (sulphate reduction rates, sediment nutrient fluxes, organic content, C, N and P content) to examine possible positive and negative effects of the mussels on eelgrass performance. The fluxes of ammonium from the sediments were stimulated at all sampling dates at the Mixed station, and possibly stimulated epiphyte growth at this station. Further ¹⁵N signals in epiphytes from the Mixed station suggested that excretion products from the mussels were important nitrogen sources at this station. Sulphate reduction rates were enhanced at the Mixed station and also sediment sulphide concentrations increased under mussel influence, which may have resulted in sulphide toxicity and decreased growth of Z. marina at this station. The study indicates that for Z. marina beds in Flensborg Fjord the effects of M. edulis in seagrass beds are primarily negative, and raises the question whether this leads to negative effects on the stability and expansion of Z. marina beds.     

石斑鱼肠道原籍嗜冷杆菌( PsychrobacterSP.)SE6作为益生菌的体内外评价

研究了石斑鱼肠道原籍嗜冷杆菌SE6在体外的抑菌特性及对人工胃肠液的耐受特性,在此基础上研究了饲料中添加SE6(108CFU/g)对斜带石斑鱼生长性能和免疫功能的影响。体外试验结果表明:SE6对副溶血弧菌、哈维氏弧菌、梅氏弧菌和金黄色葡萄球菌等常见水产致病菌均有一定的抑制作用。pH为4.0的人工胃液能促进SE6的生长,处理2和4 h存活率分别为110.7%和173.0%;pH为3.0的人工胃液处理2 h后活菌数为1.59×105 CFU/cm3,存活率为1.36%,但SE6不能耐受pH为2.0的人工胃液。经浓度分别为0.1%,0.3%,0.5%和0.7%的胆盐处理3 h后,SE6的存活率分别为89.17%,88.33%,77.17%和12.33%。动物试验结果显示:投喂SE6 60 d后试验组饵料系数(0.95±0.01)显著低于对照组(1.03±0.02)。试验组血清补体C3在试验全期均高于对照组,补体C4含量在投喂SE6 60 d时显著上升。血清IgM含量在投喂30 d时显著上升。总之,嗜冷杆菌SE6能抑制多种常见病原菌的生长,而且对模拟胃肠道环境具有一定的耐受能力,添加到饲料中能显著提高石斑鱼饵料利用率,增强动物的免疫功能。     

Population ecology of fiddler crab, Uca lactea annulipes (Decapoda: Ocypodidae) in Sirik mangrove estuary,Iran

Fiddler crabs are known as the most conspicuous bioturbating animals in mangrove forest, which by their burrowing activities promote nutrient cycling in ecosystems. Mangroves, crabs and bacteria are connected in the form of positive feedback loop, whereby an increase in the activity of any participant will tend to increase the activity of all others as well. Population structure, growth, mortality, recruitment, biomass and production of fiddler crabs as dominant species of deposit-feeder crabs have been investigated from May 2005 to March 2006 in the Sirik mangrove estuary in the eastern part of Strait of Hormuz. Ten 1 m² squares of the substrate were randomly sampled during low tide. There was a significant change in organic content in the sampling area during study period. The sex ratio was significantly deviated from 1:1 ratio in large sizes, where males were dominated year around. Fiddler crabs showed seasonal growth pattern during a year, which the maximum growth was observed during autumn and early summer and ceased during winter by low temperature. Growth parameters were estimated by Appeldoorn's method and fitted with seasonalized von Bertalanffy growth models. The Kimura likelihood ratio test revealed significant differences in the overall growth curves between males and females, originated from the differences in k$k$ and L_∞$L_{\mathrm{\infty }}$ between sexes. Males were found to exhibit higher growth rate and larger size than females, this could be attributed to the higher investment of energy for reproduction in females. Reproduction was restricted to spring and early summer, while recruitment occurs at the end of summer. Total annual production of males and females was 37.90 and 10.05 g dry weight/10 m² respectively and major production took place during early summer. Because of impact of temperature on the activity of fiddler crabs in subtropical regions, density of crabs has significant positive correlation with temperature (P < 0.01), while growth rate shows significant negative correlation with organic matter of sediment (P < 0.05). In conclusion these crabs showed lower mortality and production rate with the seasonal growth and recruitment in comparison with tropical species.     

Distribution of amphipods Ischyrocerus on the red king crab, Paralithodes camtschaticus: Possible interactions with the host in the Barents Sea

The determination of costs and benefits experienced by crustaceans as a result of occupation by their symbionts has received increased attention from marine ecologists. However, the interactions between some important species and their associates remain unclear. We examined the distribution of amphipods in the genus Ischyrocerus on the red king crab Paralithodes camtschaticus, a commercially important species, in two areas of the Barents Sea. Ischyrocerus commensalis was found on 30.5% of crabs in Dalnezelenetskaya Bay (DZB) with the mean number per crab being 55.1, in Dolgaya Bay (DLB) these rates were 28.6% with 19.3 specimens per crab. Sympatric species Ischyrocerus anguipes was found on 13.5% of crabs in DZB with a mean of 7.3 individuals per host, in DLB it had much lower occurrence (1.3% and 1.5 specimens per host). There were no significant differences between proportions of male and female crabs infested by amphipods in both areas examined. Prevalence of amphipods was similar among years examined except for I. commensalis on small crabs (carapace length CL <90 mm) and I. anguipes on large crabs (CL >90 mm) in DZB. We found that I. commensalis and I. anguipes are not egg predators of P. camtschaticus in the Barents Sea, at least in summer. High numbers of I. commensalis occur in crab gills, and both mean intensity of the amphipods and their empty tubes increased with crab size. In the gills, I. commensalis predominated in the section nearest the mouth parts. Possible negative impacts for the hosts due to gills infestation are discussed. In contrast, I. anguipes were predominately found on the carapace and limbs of crabs and appears to be a less specific symbiont of P. camtschaticus. Both amphipod species seem to be commensals, however possible negative impacts for the host could not be excluded.     

Stage-V copepodites of Calanus sinicus and Calanus jashnovi (Copepoda: Calanoida) in mesopelagic zone of Sagami Bay as identified with genetic markers, with special reference to their vertical distribution

We applied genetic makers to identify Calanus species occurring in Sagami Bay, Japan, in order to investigate their vertical distribution in the upper 1000 m. First, interspecific genetic distances of three gene loci, mitochondrial small ribosomal RNA (srRNA), nuclear internal transcribed spacers 1 (ITS1) and 2 (ITS2), were estimated from morphologically distinguishable adult females of Calanus sinicus, Calanus jashnovi and Calanus pacificus that were collected from Sagami Bay, the Kuroshio Extension and the Oyashio region, respectively. The highest levels of interspecific genetic distance were observed in srRNA, followed by ITS1 and ITS2. The intraspecific genetic distances within C. sinicus were much lower than the interspecific genetic distances, indicating that DNA sequences in these loci are consistent with the morphological differences. This information was used as a criterion for species identification based on DNA sequence variation, and allowed us to identify the fifth copepodites (CVs) or younger stages of these species. Next, the vertical distribution of Calanus species was investigated in Sagami Bay in May 2006, on the basis of a stratified sampling in the upper 1000 m. By applying the genetic markers, 23 individuals comprising all copepodite stages were allocated into either C. sinicus or C. jashnovi, and the small- and large-sized CVs were identified as C. sinicus and C. jashnovi, respectively. The total abundance of C. sinicus was highest at 0-50 m and decreased with depth. On the contrary, CV individuals of C. sinicus were abundant not only in 0-50 m but also below 200 m with minimum occurrences in 150-200 m depth. C. jashnovi was much less abundant than C. sinicus and comprised of only CIV and CV which occurred in the upper 100 m and deeper than 50 m depths, respectively. The abundance of C. sinicus in the 1000-m water column of Sagami Bay was at a level comparable to that in shelf waters, suggesting the importance of off-shelf individuals in the biological production and organic transport in the respective areas.     

Myrionecta rubra (Mesodinium rubrum) bloom initiation in the Columbia River estuary

To better understand the development of the annually recurring late summer red water blooms of the phototrophic ciliate Myrionecta rubra in the Columbia River estuary we examined its standing stocks and measured its growth rates both in the estuary main channels and in Baker Bay, a peripheral embayment situated near the river mouth. Data collected during two summers show a biphasic development of M. rubra blooms, with an initial phase when the protist was only detected in Baker Bay, followed by an established phase when red waters were observed throughout the lower estuary. Ilwaco harbor (Baker Bay’s seaward-end) is at least one of the locations where the bloom starts since M. rubra was detected there at concentrations >100s cell L⁻¹ before Chinook harbor (Baker Bay’s upriver-end) or the estuary main channels. In 2010, this initial phase lasted about 1.5 months, spanning the neap tide of early July to the beginning of the neap tide of mid-August. While high growth rates were measured in Ilwaco harbor during the initial phase (1.2–3.1 d⁻¹) and in the estuary main channels in both surface red (0.7 d⁻¹) and adjacent non-red (1.1 d⁻¹) waters during the established period, growth of the ciliate was not detected in Ilwaco harbor during this second phase. Growth rate data obtained during the established bloom phase also suggest that M. rubra cells in the estuary mostly divide during the daytime and that red water patches might experience self-shading.     

Fate of the Black Sea Acartia clausi and Acartia tonsa (Copepoda) penetrating into the Marmara Sea through the Bosphorus

In October 2005 spatial distribution of live and dead Acartia clausi and Acartia tonsa was studied in the Black and Marmara Seas and near the Marmara Sea inlet of the Bosphorus, in order to understand their fate upon transportation between two seas. The morphometric characteristics in both species from all studied areas, and the decreased abundance of A. clausi and A. tonsa from the Black Sea towards the Marmara Sea indicate that the Marmara Sea Acartia populations are formed by recruitment from the Black Sea. We observed mass mortality of A. clausi in the Marmara Sea near the Prince Islands. The majority of carcasses (66% of total A. clausi numbers in the Marmara Sea) were found in the salinity gradient layer.     

漳州西施舌线粒体基因组全序列:腔蛤蜊属存在新种的证据

利用长PCR扩增漳州西施舌线粒体DNA(ZZ-mtDNA),用引物步移法测序,获得线粒体基因组DNA全序列,研究其基因组特点。结合双壳类49个物种线粒体全基因组,分析基因间核苷酸和蛋白质氨基酸序列的差异,构建系统进化树,探讨漳州西施舌系统演化地位。研究结果表明:漳州西施舌线粒体基因组DNA全长17 199 bp,A+T含量为64.2%,编码36个基因,其中12个蛋白质编码基因,22个转运RNA基因(tRNAs),2个核糖体RNA基因(lrRNA 和srRNA),全部基因均位于重链上,tRNASer为单拷贝,tRNAMet为双拷贝;非编码区占10.9%(1 882 bp/17 199 bp),其中,主非编码区为882 bp,与RZ-mtDNA主非编码区差异明显,另有一个较大(400 bp)的非编码区为漳州西施舌特异性非编码区;以双壳纲帘蛤目文蛤属3种贝类、贻贝目贻贝属4种贝类、珍珠贝目牡蛎科的6种贝类为参照,对编码基因的核苷酸序列和蛋白质氨基酸序列进行差异分析显示,漳州西施舌与日照西施舌达到了种间差异水平。线粒体基因组编码的基因、tRNA组成、非编码区均揭示漳州西施舌是腔蛤蜊属的一个新种。     

Seasonal variability in somatic and reproductive investment of the bivalve Scrobicularia plana (da Costa, 1778) along a latitudinal gradient

Monthly investment in soma and gonads in the bivalve Scrobicularia plana is described for three populations along its distributional range: Minho estuary, Portugal; Westerschelde estuary, The Netherlands and Buvika estuary, Norway. Seasonal cycles in body mass (BMI), somatic mass (SMI) and gonadal mass (GMI) indices were observed for all populations. In Portugal, BMI and SMI peaked in mid-autumn, while in The Netherlands both indices were at their highest in mid-spring. Norway showed a different pattern with two distinct peaks: one in mid-autumn and a second peak in spring. GMI reached maximum values in July in Portugal and Netherlands and in June in Norway. Overall, mean BMI and SMI were lower in Portugal while mean GMI was lower in Norway. The spawning period lasted the whole summer in Portugal, but was shorter (only two months) in The Netherlands and Norway. The reproductive investment in The Netherlands was significantly higher than in Portugal and Norway, with the lowest values being observed in Norway. Differences in annual cycles between populations were attributed to environmental factors, namely temperature and food availability. Temperature seems important in shaping the reproductive pattern with more northern populations showing shorter reproductive periods starting later in the year, and a lower reproductive output. In addition, winter water temperatures can explain the lower mean body and somatic mass values observed in Portugal. Food availability influenced the physiological performance of the species with peaks in somatic mass coinciding with phytoplankton blooms. This relation between physiological performance and environmental factors influences S. plana distribution, densities and even survival, with natural consequences on its commercial importance.     

Spatial diversity of rocky midlittoral macro-invertebrates associated with the endangered species Patella ferruginea (Mollusca: Gastropoda) of Tunisian coastline

The present study focuses on horizontal spatial variability of benthic macrofauna associated with Patella ferruginea. Thirty-six samples collected at 12 transects belonging to 4 midlittoral sites along the rocky Tunisian coastline, were examined. A total of 44 species belonging to 5 taxa were found. Multivariate analysis applied on gathered data did not show a horizontal spatial variability at small scale (between transects), but at large scale, between sites as well as sectors. Thus, three groups of communities were identified (GI: Korbous and El Haouaria; GIIa: Zembra Island and GIIb: Kelibia). The distribution of species abundance within these groups revealed that crustaceans were the most abundant taxon, due to the overwhelming dominance of Chthamalus stellatus. This substratum appeared to create favourable micro-habitats for the installation of molluscs including gastropods. Regarding the low diversity index (H') and evenness (J), they seemed to reflect a disturbance and a demographic unbalance within these communities. The heterogeneity of substrate surface, created by C. stellatus specimens appeared to be caused by various complex interactions established between the key components of these communities in particular suspension feeders, predators, herbivorous molluscs and macroalgae. Thus, the dynamic status of each of these communities is the result of these complex interactions.     

中国海域长眼寄居蟹属两个新记录种

在我国南海海域大型底栖生物进行分类和多样性研究中,发现我国长眼寄居蟹属2个新记录种,紫斑长眼寄居蟹(Paguristes gonagrus)和条纹长眼寄居蟹(Paguristes runyanae),并对其进行描述。紫斑长眼寄居蟹采集自海南岛,主要特征为：螯足和步足密布刚毛,第一触角柄未达到角膜基部,螯足腕节、掌节背面刺和突起密集而发达,螯足和第二、三步足长节末端有紫色色斑;条纹长眼寄居蟹采集于西沙群岛,主要特征为：右螯足略大于左螯足,第一触角柄短,长度仅为眼柄长度的1/2,螯足和第二、三步足背腹面均有黄色条纹。     

Feeding preferences of estuarine mysids Neomysis integer and Rhopalophthalmus tartessicus in a temperate estuary (Guadalquivir Estuary,SW Spain)

Mysid shrimps are an important component of estuarine food webs because they play a key role in energy transfer as intermediate prey. We investigated the seasonal, tidal and depth specific variation in the diet of the estuarine mysids Neomysis integer and Rhopalophthalmus tartessicus and explored its implications for the planktonic community structure of a temperate estuary (Guadalquivir Estuary, SW Spain). Neomysis integer is an opportunistic omnivore feeding mainly on mesozooplankton and on members of the detrital–microbial loop, shifting prey seasonally according to availability. In contrast, R. tartessicus showed a more carnivorous diet and shifted its target prey during seasons of low resource availability. Despite statistically significant differences in diet composition, both species shared prey of similar size, particularly juvenile Mesopodopsis slabberi, the most abundant mysid species in this estuary, and copepods. Although these similarities imply inter-specific resource competition, their co-existence is achieved by niche partitioning and spatial segregation: the higher osmoregulatory capacity and foraging plasticity of N. integer confers a broader niche breadth for this species allowing N. integer to inhabit the more stressful oligohaline region of the estuary where R. tartessicus cannot survive. We propose that this mechanism relaxes the potential for competition between N. integer and R. tartessicus.