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Mesozooplankton vertical pattern was studied down to 2000 m along an east-west transect in the oligotrophic Mediterranean Sea in June 1999. Depth-stratified samples were collected by a WP-2 net (200 µm mesh size). In addition, the copepod grazing impact on autotrophs was assessed in the epipelagic zone by measurement of small and large copepod gut content by day-night sampling. An east-west increasing trend of integrated total biomass and abundance over the 0–1000 m was detected. Similar vertical pattern of standing stock was found along the transect, with subtle differences at stations positioned over cyclonic and anti-cyclonic gyres. No significant differences were found between stations in the epipelagic and upper mesopelagic zone, except at the westernmost station (NW Mediterranean); discrimination of assemblages between the eastern and the western basins were revealed in the lower mesopelagic and bathypelagic zones. Estimated copepod grazing impact on primary production was found overall higher in the western than in the eastern basin. The entire Mediterranean Sea was dominated by copepod functional groups characterized by reproduction strategies (mostly sac spawners), trophic preferences (mainly omnivores and detritivores) and feeding modes (ambush and cruise feeding), facilitating their adaptation and efficiency in the oligotrophic to very oligotrophic environment of the Mediterranean Sea.

The Eastern Mediterranean Sea (EMS) is an ultraoligotrophic semi-enclosed sea with low nutrient levels, low primary production rate, impoverished phytoplankton populations and low zooplankton standing stocks. The Cretan Passage in the western Levantine Sea is one of the least explored areas of the EMS. We measured the mesozooplankton biomass, abundance and taxon composition as well as biochemical parameters like Electron Τransport System (ETS) activity and Aminoacyl-tRNA Synthetases (AARS) activities in late spring of 2016 (28 March–17 April 2016). The sampling area is influenced by the presence of the permanent Rhodes Cyclonic Gyre east of Crete and a series of smaller permanent or recurrent gyres south of Crete which influence the pathway of the Atlantic Water from the Ionian Sea towards the Levantine Basin at surface and subsurface layers. Mesozooplankton abundance showed slightly increasing values from western to eastern stations of the Cretan Passage and although subsurface maxima were recorded in the 50–100 m layer, the abundance at all stations generally decreased with depth. The community was dominated by copepods at all stations. Juveniles of Clausocalanus followed by juveniles of Oithona and the Oncaea media group (Oncaea waldemari, Oncaea scottodicarloi, Oncaea media) were the most abundant taxa in the upper layer, whereas Mormonilla minor, Haloptilus longicornis and Subeucalanus monachus were mainly found below 100 m. ETS, AARS and biomass for carbon measurements were conducted in the same area for the 0–100 m and 0–500 m layers. ETS was highest at LV13 with 10.56 ± 1.33 μl O2 h−1 mg protein−1 in 0–500 m and carbon biomass was highest at LV18 with 4.99 mg m−3 in 0–100 m. Lowest values of 4.52 ± 1.46 μl O2 h−1 mg protein−1 at LV18 in 0–100 m were detected for ETS and 0.91 mg m−3 at LV10 in 0–500 m for carbon biomass. Specific AARS activities showed no significant differences among sites. On the basis of ETS and AARS, we calculated that the zooplankton carbon requirements follow the west to east trend of zooplankton biomass, however, the zooplankton is not well fed and the organisms are living under oligotrophic stress.

Combined analyses of Fatty Acid Trophic Markers (FATM) and Stable Isotopes (SI) were used to characterize food preferences among copepod species/taxa and to trace their food sources in the ultra-oligotrophic Cretan Passage of the Eastern Mediterranean Sea (EMS). FATMs are based on the conservation and transfer of specific source Fatty Acids (FAs) through the food web, providing a qualitative indicator of assimilated food types. SI provide information about trophic levels and food sources of specific organisms. Most species/taxa exhibited low δ15N ratios caused by low δ15N-PON base values at the study site. A cluster analysis based on the mean Fatty Acid (FA) compositions of nine copepods from all stations identified three distinct groups (group 1: Clausocalanus lividus and Euchaeta spp.; group 2: Pleuromamma spp., Haloptilus longicornis, Lucicutia spp., Clausocalanus spp. and Corycaeus spp.; group 3: Pareucalanus attenuatus and Oncaea spp.), coinciding with similarities in feeding behavior. H. longicornis, however, revealed similar trophic positions but different food resources by FATM and SIA analyses. Overall, we conclude that omnivory was the prevailing feeding mode, demonstrating a high degree of opportunistic feeding of copepods in these ultra-oligotrophic waters. This study shed first light on the zooplankton taxa/species life strategies with regard to feeding preferences and lipid storage mechanisms in the oligotrophic EMS, where lack of relevant information exists. Although further investigation is required, the good agreement between FA and SI in some species of copepods emphasize the applicability of lipid trophic markers. Results benefit from the coupling of these indices even in oligotrophic regions of the world ocean.

According to the methodological standards established by Marine Strategy Framework Directive, the assessment for the pelagic habitat under the Biodiversity Descriptor should be carried out at the regional or sub-regional level. In the case of Mediterranean Sea, the sub-regional assessment seems optimal to take into account biogeographic differences in species composition and functional characteristics. Previous research has shown that phytoplankton diversity indicators are efficient for reliable environmental assessments, although more effort has been recommended to test these indicators on a wide spatial scale to cover wider gradients of natural and anthropogenic pressures. In this work, a set of eight diversity indices was tested against the pressure levels within a common data set of the structure and abundance of phytoplankton communities from the Adriatic, Ionian and Aegean Seas. Expert knowledge was used to define four categories of impacts that take into account partial pressures, such as point and non-point pollution, industry, ports and fisheries. At the level of the common data set, most of the diversity, evenness and dominance indices could only distinguish between the highest level of impact and the rest of impact categories. These indices maintained the distinction between two levels of subsequently dichotomised impacts (no to low impact vs. high impact) across latitudinal and longitudinal gradients. On average, the indices were less sensitive to impacts in the northernmost and westernmost areas than in the southernmost and easternmost areas, although they still showed a significant response. The results also suggest that phytoplankton communities become more uniform and less dominated by a single taxon as sampling depth increases at sites with low impact, while evenness and dominance at impacted sites remain similar at all depths. In order to establish meaningful definitions of good environmental status and targets for pelagic habitats in the Mediterranean Sea, it is necessary to establish spatially specific thresholds by additional examination of indices of good performance.

Since 2007, the non-indigenous calanoid copepod Pseudodiaptomus marinus Sato, 1913 has been increasingly recorded in numerous European sites, spreading at an unexpectedly fast pace over a short time-span. This species presents specific biological and behavioural traits which make it of particular interest for ecological and applied research topics. On 29–30 January 2018, 29 scientists from nine European Countries established the EUROBUS (Towards a EURopean OBservatory of the nonindigenous calanoid copepod Pseudodiaptomus marinUS) Working Group (WG). This WG aimed at creating a European network of institutions and researchers working on the various aspects of the biology and ecology of P. marinus, with an open forum where sharing experience and know-how among WG participants. This brought to an updated distribution map of P. marinus in European waters, as well as to the identification of priority research lines and potential joint initiatives under the WGEUROBUS umbrella. This contribution, stemming from the experts participating at the WG, represents the manifesto of the current and future initiatives developed within WGEUROBUS. info:eu-repo/semantics/publishedVersion

In order to identify how ocean acidification will influence biological interactions and fluxes among planktonic organisms and across trophic levels, a large-scale mesocosm experiment was performed in the oligotrophic Northwestern Mediterranean Sea in the framework of the European MedSeA project. Nine mesocosms were deployed in the Bay of Calvi (Corsica, France) in summer 2012. Six mesocosms were subjected to different levels of CO2 partial pressures (pCO2; 550, 650, 750, 850, 1000 and 1250 μatm) covering the range of atmospheric pCO2 anticipated for the end of this century depending on future emission scenarios, and the last three mesocosms were unaltered (ambient pCO2 of ∼450 μatm). During this 21-day experiment, we monitored copepod egg and naupliar stocks, estimated copepod (Acartia clausi and Centropages typicus) feeding rates and determined the abundance and taxonomic composition of the mesozooplankton community at the start and at the completion of the experiment. This community was clearly dominated by copepods and its final composition slightly varied between mesocosms most likely due to natural and experimental variability that cannot be related to CO2 conditions. The abundances of eggs and nauplii as well as feeding rates of A. clausi and C. typicus on diatoms, dinoflagellates and ciliates showed no significant differences among CO2 levels. The above findings suggest that the experimental set-up especially for the specific trophic conditions and the short duration of the experiment did not provide the information on the effect of acidification that was expected. The acidification might have an effect on planktonic communities and even worsen the problems imposed by food limitation, therefore on this short time scale experiment and under the extreme ologotrophic conditions the signal that dominates was the food limitation.

The rapid warming of the wolrd’s oceans during the last few decades has affected distributional patterns of marine planktonic communities. Here, we analyse links between sea warming and changes in copepod community composition over the last 3 decades (1980−2012). We used zooplankton time-series data which included 79 species of copepods collected at 3 sites in the eastern North Atlantic (Bay of Biscay and the Kattegat Sea) and the Mediterranean Sea (Gulf of Saronikos). First, using community β-diversity metrics, we analysed temporal patterns of copepod community composition changes over time and its relation to local environmental conditions. Second, to test whether the changes in copepod community composition correspond to community thermal preferences, we used the community temperature index (CTI) and compared CTI interannual changes with local temperature trends. The β-diversity analysis reveals a high temporal turnover in the copepod community composition at the 3 sites (30−45%), with a significant similarity decrease over time (‘decay’) associated with both niche descriptors and demographic stochastic processes. CTI results reveal that both in the Kattegat and Saronikos, where the ocean warming rate was the highest amongst sites, copepod community changes are linked to temperature variability, suggesting that the community is tracking their thermal niche over time. Our findings unveil the fundamental role of abiotic factors structuring copepod biodiversity over time and reveal that the local velocity of ocean warming and the species thermal thresholds are key to rearranging copepod community composition in coastal ecosystems.

Mesozooplankton composition including functional groups in relation to hydrological features is studied in the oligotrophic pelagic ecosystem of the eastern Mediterranean Sea. Combining classic/standard microscopy and advanced digital imaging approach copepod growth rates, ingestion and production rates were estimated using allometric relationships. For this purpose, mesozooplankton samples were taken in the epipelagic zone (0–200 m) in June 2016, over a grid of 16 stations in the Cretan Sea, the Cretan Passage and the eastern and western Cretan Straits including one station in the Mirtoan Sea. The sampling area is influenced by the presence of the permanent Rhodes Cyclonic Gyre east of Crete and a series of smaller permanent or recurrent gyres south of Crete that influence the anti-estuarine pathway of the Atlantic Water from the Ionian Sea towards the Levantine Basin at surface and subsurface layers. Mesozooplankton abundance exhibited almost similar values (mean value 312 ind m−3 ± 100 SD) among transects with a higher value at the western Straits (up to 562 ind m−3). Copepods were the dominant group at all stations with almost the same relative contributions among stations (76%) but slightly higher values in the Cretan Passage. The contribution of copepod functional groups in the mesozooplankton community is also highlighted. The importance of small sized copepods was underlined by small ambush feeding carnivores, prevailing at all stations and demonstrating a high degree of opportunistic feeding of copepods in these ultra-oligotrophic waters, with chlorophyll a ranging from 0.05 to 0.20 μg L−1. Copepod vital rates (production, ingestion and respiration) estimated by a growth rate model were generally low, with higher values in the Cretan Sea.

The effects of atmospheric deposition on plankton community structure were examined during a mesocosm experiment using water from the Cretan Sea (Eastern Mediterranean), an area with a high frequency of atmospheric aerosol deposition events. The experiment was carried out under spring-summer conditions (May 2012). The main objective was to study the changes induced from a single deposition event, on the autotrophic and heterotrophic surface microbial populations, from viruses to zooplankton. To this end, the effects of Saharan dust addition were compared to the effects of mixed aerosol deposition on the plankton community over 9 days. The effects of the dust addition seemed to propagate throughout the food-web, with changes observed in nearly all of the measured parameters up to copepods. The dust input stimulated increased productivity, both bacterial and primary. Picoplankton, both autotrophic and heterotrophic capitalized on the changes in nutrient availability and microzooplankton abundance also increased due to increased availability of prey. Five days after the simulated deposition, copepods also responded, with an increase in egg production. The results suggest that nutrients were transported up the food web through autotrophs, which were favored by the Nitrogen supplied through both treatments. Although, the effects of individual events are generally short lived, increased deposition frequency and magnitude of events is expected in the area, due to predicted reduction in rainfall and increase in temperature, which can lead to more persistent changes in plankton community structure. Here we demonstrate how a single dust deposition event leads to enhancement of phytoplankton and microzooplankton and can eventually, through copepods, transport more nutrients up the food web in the Eastern Mediterranean Sea, SI

Atmospheric deposition is assumed to stimulate heterotrophic processes in highly oligotrophic marine systems, controlling the dynamics and trophic efficiency of planktonic food webs, and is expected to be influenced by climate change. In the course of an 8-day mesocosm experiment, we examined the channeling, of the Saharan dust (SD) and mixed aerosols (A) effects on microplankton up to the copepod trophic level, in the highly oligotrophic Eastern Mediterranean Sea. Based on mesocosms with SD and A treatments, we evaluated the feeding response of the dominant copepod Clausocalanus furcatus every other day. We hypothesized that increased food availability under atmospheric deposition would result in increased copepod ingestion rates, selectivity and production. Overall, no robust pattern of food selection was documented, and daily rations on the prey assemblage of all mesocosms were very low indicating severe food limitation of C. furcatus. Although increased food availability was not true, after few days ingestion of ciliates was maximized, followed by egg production, in both the SD and A treatments, indicating their importance in the diet of this copepod as well as a response of C. furcatus feeding performance. Our results help in understanding the trophic efficiency of marine food webs in ultra-oligotrophic environments under atmospheric deposition. We suggest that future mesocosm research in oligotrophic waters should consider more than one copepod species., Sí

Ptacnik, Robert ... et al.-- 10 pages, 4 figures, 1 table, supplementary information https://dx.doi.org/10.1038/srep29286, Mixotrophs combine photosynthesis with phagotrophy to cover their demands in energy and essential nutrients. This gives them a competitive advantage under oligotropihc conditions, where nutrients and bacteria concentrations are low. As the advantage for the mixotroph depends on light, the competition between mixo- and heterotrophic bacterivores should be regulated by light. To test this hypothesis, we incubated natural plankton from the ultra-oligotrophic Eastern Mediterranean in a set of mesocosms maintained at 4 light levels spanning a 10-fold light gradient. Picoplankton (heterotrophic bacteria (HB), pico-sized cyanobacteria, and small-sized flagellates) showed the fastest and most marked response to light, with pronounced predator-prey cycles, in the high-light treatments. Albeit cell specific activity of heterotrophic bacteria was constant across the light gradient, bacterial abundances exhibited an inverse relationship with light. This pattern was explained by light-induced top-down control of HB by bacterivorous phototrophic eukaryotes (PE), which was evidenced by a significant inverse relationship between HB net growth rate and PE abundances. Our results show that light mediates the impact of mixotrophic bacterivores. As mixo- and heterotrophs differ in the way they remineralize nutrients, these results have far-reaching implications for how nutrient cycling is affected by light, The experiment was financed by the European Union Seventh Framework Program MESOAQUA (FP7/2009-2013) coordinated by JCN under grant agreement no. 228224. R.P. further acknowledges financial support by the German Research Council (DFG Pt 5/3-1), JMG by Spanish project REMEI (CTM2015-70340) and AC by project PROTOS (CTM2009-08783) from the Spanish Ministry of Economy and Competitiveness

Dispersal plays a key role to connect populations and, if limited, is one of the main processes to maintain and generate regional biodiversity. According to neutral theories of molecular evolution and biodiversity, dispersal limitation of propagules and population stochasticity are integral to shaping both genetic and community structure. We conducted a parallel analysis of biological connectivity at genetic and community levels in marine groups with different dispersal traits. We compiled large data sets of population genetic structure (98 benthic macroinvertebrate and 35 planktonic species) and biogeographic data (2193 benthic macroinvertebrate and 734 planktonic species). We estimated dispersal distances from population genetic data (i.e., FST vs. geographic distance) and from β-diversity at the community level. Dispersal distances ranked the biological groups in the same order at both genetic and community levels, as predicted by organism dispersal ability and seascape connectivity: macrozoobenthic species without dispersing larvae, followed by macrozoobenthic species with dispersing larvae and plankton (phyto- and zooplankton). This ranking order is associated with constraints to the movement of macrozoobenthos within the seabed compared with the pelagic habitat. We showed that dispersal limitation similarly determines the connectivity degree of communities and populations, supporting the predictions of neutral theories in marine biodiversity patterns., RADIALES (IEO)

The short-time scale evolution of plankton carbon partitioning and downward flux in the modified Black Sea water (BSW) mass entering the northeast Aegean Sea was studied using a Lagrangian approach (6-10 April 2008). The free-drifting sediment trap positioned at the bottom of the BSW layer and the control drifter, followed the same path within the anticyclone that circulates the BSW in the area. Zooplankton biomass increased (from 159 to 292 mg C m -2 ), as did faecal pellet production (from 5 to 8 mg C m -2 day -1 ), whereas a generally decreasing trend was displayed by particulate organic carbon (POC) (from 2099 to 1440 mg C m -2 ), net primary production and biomass of plankton cells >5 μm (from 32 to 11 mg C m -2 day -1 and from 153 to 124 mg C m -2 , respectively). At the same time, the organic carbon flux increased (from 131 to 311 mg C m -2 day -1 ), due to the contribution of zooplankton detritus (from 30 to 165 mg C m -2 day -1 ). Normalized biomass-size spectra slopes suggest an elevated grazing pressure upon microplankton cells and a non-steady-state ecosystem. Moreover, both the overall shallow slope values and their high correlation to organic carbon flux indicate an increased efficiency of energy transfer to higher trophic levels.

Special issue: Towards a Broader Perspective on Ocean Acidification Research.-- 9 pages, 4 figures, 3 tables, Rising levels of atmospheric CO2 are responsible for a change in the carbonate chemistry of seawater with associated pH drops (acidification) projected to reach 0.4 units from 1950 to 2100. We investigated possible indirect effects of seawater acidification on the feeding, fecundity, and hatching success of the calanoid copepod Acartia grani, mediated by potential CO2-induced changes in the nutritional characteristics of their prey. We used as prey the autotrophic dinoflagellate Heterocapsa sp., cultured at three distinct pH levels (control: 8.17, medium: 7.96, and low: 7.75) by bubbling pure CO2 via a computer automated system. Acartia grani adults collected from a laboratory culture were acclimatized for 3 d at food suspensions of Heterocapsa from each pH treatment (ca. 500 cells ml−1; 300 μg C l−1). Feeding and egg production rates of the preconditioned females did not differ significantly among the three Heterocapsa diets. Egg hatching success, monitored once per day for the 72 h, did not reveal significant difference among treatments. These results are in agreement with the lack of difference in the cellular stoichiometry (C : N, C : P, and N : P ratios) and fatty acid concentration and composition encountered between the three tested Heterocapsa treatments. Our findings disagree with those of other studies using distinct types of prey, suggesting that this kind of indirect influence of acidification on copepods may be largely associated with interspecific differences among prey items with regard to their sensitivity to elevated CO2 levels, The participation of ES andCPwas helped from funding by the Spanish Ministry of Economy and Competitivity through projects CTM2011-23480 and CTM2012-32017, respectively. This work was supported by the project CROA(LS8-1893), implemented within the framework of the Action “Supporting Postdoctoral Researchers” of the Operational Program “Education and Lifelong Learning” (Action’s Beneficiary: General Secretariat for Research and Technology), and is co-financed by the European Social Fund (ESF) and the Greek State

12 pages, 9 figures, 2 tables, Plankton biomass and composition in the pelagic zone of oceans is exposed to changes in availability of light and nutrients due to large-scale ocean circulation and water column stratification. We hypothesized that displacement of plankton from surface to deeper darker waters would not only favor heterotrophy over time, as previously suggested, but also first rapidly affect the level of mixotrophy and, consequently, overall microbial grazing in plankton food webs. To test this in an oligotrophic marine system we incubated Eastern Mediterranean water (from 10 m depth north of Crete in September 2010) in 2.8 m3 mesocosms simulating two different light intensities at the sampling station, surface waters (ca. 10 m; mesocosms L1) and deeper layers (ca. 50–60 m; mesocosms L4). The biomass and abundance of the main planktonic groups were monitored either daily or every second day, depending on the group. Microzooplankton grazing rates and the contribution of mixotrophic feeding were estimated by a combination of dilution experiments and incubations with live fluorescently labeled algae (LFLA). Although no nutrients were added to the mesocosms the chlorophyll a increased during the first 2 days of the experiment in both treatments. This increase resulted from phytoplankton growth in the light L1-mesocosm (autotrophic biomass was ca. doubled in L1 compared to L4), but was mostly due to photoadaptation of the algae in the L4-mesocosm, as indicated by lower carbon to chlorophyll a ratios. By the end of the experiment, the total biomass of protozoan and metazoan grazers in L1 was ca. twofold higher than in L4. The microzooplankton responded within the first 24 h, showing different grazing activity in L1 than in L4. Microzooplankton grazing rates on total Chl a were similar in both treatments; however, phytoplankton instantaneous growth rates were higher in the more illuminated mesocosm. This resulted in a closer coupling between both rates in L4, where all production was grazed daily, than in L1. Nevertheless, the overall flux of carbon through the microzooplankton was 33–60% higher in L1 compared to L4 throughout the experiment. The fraction of mixotrophy in the ciliate community varied in L1 (20–50%), but decreased over time in L4 from 50% to 30%. Our results do not support studies from freshwater, postulating that reduced light and nutrient limitation may increase herbivore production due to stoichiometric effects. Finally, we discuss how mixotrophy may bias rate estimates in dilution experiments, This work was funded through projects CTM2009-08783 from the SpanishMinistry of Science and Innovation assigned to A.C. and projects LightDynamix, LiMic and LightCopFed from the European Union Seventh Framework Program (FP7/2007-2013) under grant agreement no. 228224, MESOAQUA, assigned to R.P. (LightDynamix), A.C., R.A.M., A.S., A.G. (LiMic) and S.I. (LightCopFed)

We analyzed and compared Mediterranean mesozooplankton time series spanning 1957–2006 from six coastal stations in the Balearic, Ligurian, Tyrrhenian, North and Middle Adriatic and Aegean Sea. Our analysis focused on fluctuations of major zooplankton taxonomic groups and their relation with environmental and climatic variability. Average seasonal cycles and interannual trends were derived. Stations spanned a large range of trophic status from oligotrophic to moderately eutrophic. Intra-station analyses showed (1) coherent multi-taxa trends off Villefranche sur mer that diverge from the previous results found at species level, (2) in Baleares, covariation of zooplankton and water masses as a consequence of the boundary hydrographic regime in the middle Western Mediterranean, (3) decrease in trophic status and abundance of some taxonomic groups off Naples, and (4) off Athens, an increase of zooplankton abundance and decrease in chlorophyll possibly caused by reduction of anthropogenic nutrient input, increase of microbial components, and more efficient grazing control on phytoplankton. (5) At basin scale, the analysis of temperature revealed significant positive correlations between Villefranche, Trieste and Naples for annual and/or winter average, and synchronous abrupt cooling and warming events centered in 1987 at the same three sites. After correction for multiple comparisons, we found no significant correlations between climate indices and local temperature or zooplankton abundance, nor between stations for zooplankton abundance, therefore we suggest that for these coastal stations local drivers (climatic, anthropogenic) are dominant and that the link between local and larger scale of climate should be investigated further if we are to understand zooplankton fluctuations, Sí

This study includes the first information on the combined effect of low pH and raised temperature on egg production rate (EP), hatching success (HS), excretion and respiration of the Mediterranean copepod Acartia clausi. Adult individuals of A. clausi and fresh surface seawater were collected at a coastal station in Saronikos Gulf during April 2012. Four different conditions were applied: two different pH levels (present: 8.09 and future: 7.83) at two temperature values (present: 16°C and present+4 °C= 20°C). EP and HS success decreased significantly over the duration of exposure at future pH at both temperature conditions. However, the analysis of the combined effect of pH, T, chlorophyll α and the duration of the experiments on EP and HS revealed that ocean acidification had no discernible effect, whereas warming; food and the duration of exposure were more significant for the reproductive output of A. clausi. Temperature appeared to have a positive effect on respiration and excretion. Acidification had no clear effect on respiration, but a negative effect on the A. clausi excretion was observed. Acidification and warming resulted in the increase of the excretion rate and the increase was higher than that observed by warming only. Our findings showed that a direct effect of ocean acidification on copepod’s vital rates was not obvious, except maybe in the case of excretion. Therefore, the combination of acidification with the ambient oligotrophic conditions and the warming could result in species being less able to allocate resources for coping with multiple stressors.