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The Eastern Boundary Upwelling Systems (EBUS) sustains some of the most productive marine systems on Earth. Within each of these systems, the upwelling process exhibits spatial and temporal variation resulting in marked differences in upwelling intensity and seasonality along extensive coastlines. The study of this variation is well needed, given the magnitude of the services provided by upwelling, and the impending impacts of global warming on EBUS. The critical link between the physical variability associated with upwelling intensity and its consequences on socio-ecological variables remain severely understudied. This study aimed to address such a gap by exploring the influence of coastal upwelling intensity on socio-ecological attributes of co-management units named Marine Exploitation Areas for Benthic Resources (MEABRs), along one the most productive ecosystems of the world: The southern Humboldt Current system. We evaluated the non-linear influence of upwelling on 1) the harvest of economically important resources, 2) the number of exploited species, 3) number and gender distribution of fishers involved, and 4) fishery activities. Our data indicated that on the one hand the annual harvest of commercial species, and all the exploited resources combined, were consistently higher in MEABRs associated with intermediate to high upwelling intensities. On the other hand, the harvest of kelp, the number of species harvested, and the number of fishers per MEABR increased towards low upwelling intensities, showing signals of fishery diversification. Interestingly, representation of female fishers increased towards high upwelling intensities, suggesting that multiple factors account for the variation in this, and other socio-ecological variables examined. Our study provides first-hand information about harvest levels and the allocation of fishery activities and gender distribution when MEABRs associated with different upwelling intensities are compared. Such information will assist in the identification of ecological and social vulnerabilities in a global warming scenario., Competing Interests: Declaration of competing interest We wish to confirm that there are no known conflicts of interest associated with this publication and that the grants associated to resources to make experiments has been indicated in acknowledgement section., (Copyright © 2025 Elsevier Ltd. All rights reserved.)

Upwelling, as a large oceanographic phenomenon, increases coastal productivity and influences all levels of biological complexity. Despite decades of research on it, much remains to be understood about the impact of upwelling on the feeding behavior and thermal tolerance of important groups such as fish. Hence, our aim was to investigate how upwelling conditions modify the feeding behavior and thermal tolerance of a prominent intertidal fish, Girella laevifrons. We collected purple mussels (Perumytilus purpuratus) from upwelling (U) and downwelling sites (DU) in central Chile, and used them as prey in feeding trials and measuring the concentration of organic matter and proteins in their tissues. We assessed fish consumption rates and growth in fish collected from the same U and DU sites, feeding on either U or DU mussels. Lastly, we assessed the thermal tolerance of U and DU fish fed with the aforementioned U vs DU mussels. We found that U mussels held higher concentrations of organic matter and proteins compared to their DU counterparts. U mussels were also selected and consumed in larger amounts than DU mussels, although the origin of the fish also influenced consumption rates. Thermal tolerance assays revealed that U fish exhibited higher maximum performance (Max.pf) and critical thermal maxima (Ct max ) and lower sensitivity to temperature changes (as measured by Q 10 ), compared to DU fish. Altogether, these results point to a strong influence of upwelling on the quality of organisms' tissues, indirectly altering key aspects of fish feeding behavior and thermal tolerance. These findings also contribute to understanding the physiological adjustments organisms make in productive upwelling systems, and how they may change in the future with ongoing climate events., Competing Interests: Declaration of competing interest The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: Jose Pulgar reports a relationship with FONDECYT that includes: funding grants. If there are other authors, they declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Eastern Boundary Upwelling Systems (EBUS) deliver cold, nutrient-rich waters, influencing coastal biota from the molecular to the ecosystem level. Although local upwelling (U) and downwelling (DU) conditions are often known, their influence on body attributes of relevant species has not been systematically compared within and between EBUS (i.e., below and above regional scales). Hence, we compared the physical-chemical characteristics of U and DU sites in the Humboldt Current system (Chile) and the Iberian Current system (Portugal). We then assessed the influence of U and DU upon eight body attributes in purple mussels (Perumytilus purpuratus) and Mediterranean mussels (Mytilus galloprovincialis), from the Humboldt and Iberian systems, respectively. We hypothesized that bivalves from U sites display better fitness, as measured by body attributes, regardless of their origin (EBUS). As expected, waters from U sites in both systems showed lower temperatures and pH, and higher nitrite concentrations. We also found that mussels from U sites showed better fitness than those in DU sites in 12 out of 16 direct U vs DU comparisons. Shell length, shell volume, organic content of soft-tissues, and mechanical properties of the shell averaged consistently higher in mussels from U sites in both Current systems. In addition, total weight, soft-tissue weight, shell weight and shell thickness were all higher in the U site at the Humboldt system but had less consistent differences at the Iberian system. Altogether, most results supported our working hypothesis and indicate that U conditions support better fitted mussels. The few attributes that did not exhibit the expected U vs DU differences in the Iberian system suggest that local and species-specific differences also play a role on the attributes of these species. These results may also serve as a reference point for further studies addressing the influence of upwelling in these productive, critically important systems., Competing Interests: Declaration of competing interest We wish to confirm that there are no known conflicts of interest associated with this publication and that the grants associated to resources to make experiments has been indicated in acknowledgement section., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Upwelling systems deliver nutrient-rich water into coastal ecosystems, influencing primary productivity and potentially altering seaweed-herbivore interactions. Upwelling bottom-up effects on distinct trophic levels are well-known. However, their influence on seaweed biomolecules and on algae-herbivore interactions and growth are less known. The aim of this study was threefold: i) to compare physical-chemical characteristics and nutrient levels in the water of upwelling (U) and downwelling (DU) zones, ii) to quantify their influence on the content of protein and carbohydrates in seaweed tissues of representative U and DU locations, and iii) to experimentally assess their effect on the feeding behavior and growth of a prominent intertidal herbivore, the sea urchin Loxechinus albus. Waters from U zones showed lower temperatures and pH, and higher phosphate concentrations than those from downwelling zones. Similarly, the tissue of seaweeds from a U location had significantly more proteins and carbohydrates than those from a DU location. The origin location of the sea urchins had a significant influence on consumption and growth rates: in general, those coming from a site with U conditions consumed and grew more than those coming from DU conditions. The quality of the algae was a significant factor on consumption rates, although in the case of preference trials, this factor interacted with sea urchin origin location. Our results show that the availability and quality of the food in upwelling zones has an influence on herbivore-seaweed direct interactions. However, these interactions and the growth of the sea urchins were also related to the coastal site and conditions from which the sea urchins came from. These results are relevant considering the expected impact of climate change on the world's oceans, and the importance of U zones as thermal (cold water) refuges for marine ectotherms., Competing Interests: Declaration of competing interest We wish to confirm that there are no known conflicts of interest associated with this publication and that the grants associated to resources to make experiments has been indicated in acknowledgement section., (Crown Copyright © 2022. Published by Elsevier B.V. All rights reserved.)

Environmental variation alters biological interactions and their ecological and evolutionary consequences. In coastal systems, trematode parasites affect their hosts by disrupting their life-history traits. However, the effects of parasitism could be variable and dependent on the prevailing environmental conditions where the host-parasite interaction occurs. This study compared the effect of a trematode parasite in the family Renicolidae (metacercariae) on the body size and the shell organic and mechanical characteristics of the intertidal mussels Perumytilus purpuratus, inhabiting two environmentally contrasting localities in northern and central Chile (ca. 1600 km apart). Congruent with the environmental gradient along the Chilean coast, higher levels of temperature, salinity and pCO 2 , and a lower pH characterise the northern locality compared to that of central Chile. In the north, parasitised individuals showed lower body size and shell resistance than non-parasitised individuals, while in central Chile, the opposite pattern was observed. Protein level in the organic matter of the shell was lower in the parasitised hosts than in the non-parasitised ones regardless of the locality. However, an increase in polysaccharide levels was observed in the parasitised individuals from central Chile. These results evidence that body size and shell properties of P. purpuratus vary between local populations and that they respond differently when confronting the parasitism impacts. Considering that the parasite prevalence reaches around 50% in both populations, if parasitism is not included in the analysis, the true response of the host species would be masked by the effects of the parasite, skewing our understanding of how environmental variables will affect marine species. Considering parasitism and identifying its effects on host species faced with environmental drivers is essential to understand and accurately predict the ecological consequences of climate change., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Artificial Light at Night (ALAN) is expanding worldwide, and the study of its influence remains limited mainly to documenting impacts, overlooking the variation in key characteristics of the artificial light such as its intensity. The potential dose-response of fitness-related traits to different light intensities has not been assessed in sandy beach organisms. Hence, this study explored dose-responses to ALAN by exposing the intertidal sandy beach isopod Tylos spinulosus to a range of light intensities at night: 0 (control), 20, 40, 60, 80 and 100 lx. We quantified the response of this species at the molecular (RNA:DNA ratios), physiological (absorption efficiency) and organismal (growth rate) levels. Linear and non-linear regressions were used to explore the relationship between light intensity and the isopod response. The regressions showed that increasing light intensity caused an overall ~ threefold decline in RNA:DNA ratios and a ~ threefold increase in absorption efficiency, with strong dose-dependent effects. For both response variables, non-linear regressions also identified likely thresholds at 80 lx (RNA:DNA) and 40 lx (absorption efficiency). By contrast, isopod growth rates were unrelated (unaltered) by the increase in light intensity at night. We suggest that ALAN is detrimental for the condition of the isopods, likely by reducing the activity and feeding of these nocturnal organisms, and that the isopods compensate this by absorbing nutrients more efficiently in order to maintain growth levels., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Widespread intertidal mussels are exposed to a variety of natural and anthropogenic stressors. Even so, our understanding of the combined influence of stressors such as predation risk and ocean acidification (OA) on these species remains limited. This study examined the response of the purple mussel (Perumytilus purpuratus), a species distributed along Pacific southeastern rocky shores, to the effects of predation risk and OA. Using a laboratory 2 × 2 cross design, purple mussels were either devoid or exposed to predator cues from the muricid snail Acanthina monodon, while simultaneously exposing them to current (500 ppm) or projected OA conditions (1500 ppm). The response of purple mussels to these factors was assessed using growth, calcification, clearance, and metabolic rates, in addition to byssus production. After 60 d, the presence of predator cues reduced mussel growth in width and length, and in the latter case, OA enhanced this response making the effects of predator cues more severe. Calcification rates were driven by the interaction between the two stressors, whereas clearance rates increased only in response to OA, likely explaining some of the growth results. Mussel byssus production also increased with pCO 2 but interacted with predation risk: in the absence of predator cues, byssus production increased with OA. These results suggest that projected levels of OA may alter and in some cases prevail over the natural response of purple mussels to predation risk. Considering the role played by this mussel as a dominant competitor and ecosystem engineer in rocky shores, these results have community-wide implications., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Ocean acidification (OA) is expected to rise towards the end of the 21st century altering the life history traits in marine organisms. Upwelling systems will not escape OA, but unlike other areas of the ocean, cooling effects are expected to intensify in these systems. Regardless, studies evaluating the combined effects of OA and cooling remain scarce. We addressed this gap using a mesocosm system, where we exposed juveniles of the intertidal muricid snail Acanthina monodon to current and projected pCO 2 (500 vs. 1500 ppm) and temperature (15 vs. 10 °C) from the southeast Pacific upwelling system. After 9 weeks of experimental exposure to those conditions, we conducted three estimations of growth (wet weight, shell length and shell peristomal length), in addition to measuring calcification, metabolic and feeding rates and the ability of these organisms to return to the normal upright position after being overturned (self-righting). Growth, feeding and calcification rates increased in projected cooling conditions (10 °C) but were unaffected by pCO 2 or the interaction between pCO 2 and temperature. Instead, metabolic rates were driven by pCO 2 , but a significant interaction with temperature suggests that in cooler conditions, metabolic rates will increase when associated with high pCO 2 levels. Snail self-righting times were not affected across treatments. These results suggest that colder temperatures projected for this area would drive this species growth, feeding and calcification, and consequently, some of its population biology and productivity. However, the snails may need to compensate for the increase in metabolic rates under the effects of ocean acidification. Although A. monodon ability to adjust to individual or combined stressors will likely account for some of the changes described here, our results point to a complex dynamic to take place in intertidal habitats associated with upwelling systems., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Coastal habitats worldwide, including sandy beaches, are becoming increasingly exposed to Artificial Light at Night (ALAN). Despite the spread of this global stressor, research assessing ALAN potential impacts remain scarce, particularly at the molecular level. This study addressed this gap by assessing the influence of ALAN on the physiological condition of the sandy beach insect Phalerisida maculata Kulzer (Coleoptera, Tenebrionidae). RNA:DNA ratios were used here as a proxy of the insect's nutritional condition in laboratory trials that lasted 20 d. Insects were exposed to two representative ALAN conditions (either 60 or 120 lx) and compared with those maintained in a natural daylight/night cycle (0 lx at nigth; control). After the trial, organisms from each treatment were frozen in liquid nitrogen and standard protocols were followed to estimate RNA, DNA and RNA:DNA ratios. Estimates of RNA:DNA ratios from insects maintained in control conditions were significantly higher (P < 0.05) than those from insects exposed to ALAN. The reduced nutritional condition of insects exposed to light pollution is explained by the lower in situ biosynthetic capacity in these organisms resulting from a reduction in their feeding. ALAN likely altered P. maculata normal locomotor activity, which takes place primarily at night, forcing the insects to remain buried in the sand for extended periods of time. As ALAN continues to spread along coastlines worldwide, there is a likelihood of growing impacts on these and other species living on sandy beaches and other coastal habitats., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Environmental variation may alter biological interactions and their ecological consequences. For instance, in marine ecosystems hosts and parasites are subject to environmental variability across latitudinal gradients, and their co-evolutionary dynamics may be the result of the interplay with local physical-chemical variables in seawater. Thus, assessing the environmental conditions required for a host in order to improve their survival is essential to understand the host-parasite interaction and dynamics. In this study, we evaluated the impact of parasitism by Proctoeces humboldti on the body size and reproduction of the intertidal keyhole limpet Fissurella crassa collected from three populations spanning ca. 1500 km along the latitudinal gradient of the Chilean coast. In addition, for the first time, we explore whether the effect of parasitism can be extended to changes in the organic composition and mechanical properties of the host shell. Our results show that parasitism prevalence and intensity, and body size of F. crassa increased in central Chile (ca. 33°S). Unlike body size, which was greater in parasitized limpets than in non-parasitized limpets at the three study sites, reproductive performance followed this trend only in central Chile populations, with no differences between parasitized and non-parasitized limpets collected in the northern Chilean (ca. 23°S), and lower in parasitized than non-parasitized individuals from the south-central Chile (ca. 37°S). The organic composition of F. crassa shells showed significant differences between parasite conditions (e.g. polysaccharides and water decreased in parasitized limpets) and across sites (e.g. proteins levels increase in shell of parasitized limpets from central Chile, but decreased at south-central Chile). However, variability in shell mechanical properties (e.g. toughness and elastic module) do not showed significant differences across sites and parasitism condition. These results suggest the interplay of both parasitism and environmental fluctuations upon the reproductive performance and morphology of the host. In addition, our result highlight that the host may also trade-offs reproduction, growth and shell organic composition to maintain the shell functionality (e.g. protection for mechanical forces and durophagous predators)., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Parasites alter the reproductive performance of their hosts, limit their growth, and thereby modify the energy budget of these hosts. Experimental studies and theoretical models suggest that the outcome of the host-parasite interactions could be determined by ecological factors such as food availability levels in the local habitats. Nutrient inputs may affect the host's food resource availability with positive or negative effects on parasite infection rates and tolerance of infection, however this has not been specifically evaluated in natural systems. In this study, we evaluate the effects of parasitism by Proctoeces humboldti on body size, gonadosomatic index (GSI), and metabolic rate (oxygen consumption) of their second intermediate host Fissurella crassa limpets, under contrasting natural conditions of productivity (upwelling center vs upwelling shadow sites). Our results evidenced that parasitized limpets collected from the intertidal habitat influenced by coastal upwelling site showed greater shell length, muscular foot biomass and GSI as compared to non-parasitized limpets collected in the same site, and compared to parasitized and non-parasitized limpets collected from the sites under the influence of upwelling shadow conditions. Oxygen consumption was lower in parasitized limpets collected from the upwelling-influenced site than in the other groups, independent of age, suggesting reduced metabolic stress in infected individuals inhabiting these productive sites. Our results suggest that increased productivity in upwelling sites could mitigate the conflict for resources in the P. humboldti - F. crassa system, influencing where such interaction is found in the continuum between parasitism and mutualism. Since parasitism is ubiquitous in natural systems, and play important roles in ecological and evolutionary processes, it is important to analyze host-parasite interaction across a variety of ecological conditions, especially in biological conservation., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Biological interactions and environmental constraints alter life-history traits, modifying organismal performances. Trematode parasites often impact their hosts by inducing parasitic castration, frequently correlated with increased body size in the host (i.e., gigantism hypothesis), which is postulated to reflect the re-allocation of energy released by the reduction in the reproductive process. In this study, we compared the effect of a trematode species on shell size and morphology in adult individuals of the intertidal mussels Perumytilus purpuratus (>20 mm) collected from two local populations of contrasting environmental regimes experienced in central-southern Chile. Our field data indicates that in both study locations, parasitized mussels evidenced higher body sizes (shell length, total weight and volume) as compared with non-parasitized. In addition, parasitized mussels from the southern location evidenced thinner shells than non-parasitized ones and those collected from central Chile, suggesting geographical variation in shell carbonate precipitation across intertidal habitats of the Chilean coast. In laboratory conditions, mussels collected from a local population in central Chile were exposed to two temperature treatments (12 and 18 °C). Parasitized mussels showed higher growth rates than non-parasitized, regardless of the seawater temperature treatments. However, the metabolic rate was not influenced by the parasite condition or the temperature treatments. Our field and laboratory results support the parasite-induced gigantism hypothesis, and suggest that both the thermal environment and geographic location explain only a portion of the increased body size, while the parasitic condition is the most plausible factor modulating the outcome of this host-parasite interaction.

UV-radiation (UVR) and temperatures have increased substantially over recent decades in many regions of the world. Both stressors independently have shown to affect the metabolism and growth in fish. However, because increase of both stressors are occurring concomitantly, to better understand their influences on marine species, their combined effects were evaluated. We test the hypothesis that UVR and temperature act synergistically affecting the metabolism, digestive process and growth of an intertidal fish. Two UVR conditions (with and without UVR) and two temperature levels (20° C and 25° C) were used. UVR increase the oxygen consumption and this was associated to opaque feces production. The absorption efficiency was higher without UVR at high temperatures (25 °C) and with UVR at low temperatures (20 °C). Finally, independent of UVR treatment, fish subjected to low temperature have higher biomass than those of high temperature. The interaction between UVR and temperature may influence on the physiology and growth of animals that inhabit in extreme habitats as upper intertidal, it could pose significant functional for aquatic animal survivorship., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

The rapid increase in body size and abundance of most species inside Management and Exploitations Areas for Benthic Resources (MEABRs) has led to the proposal of these areas as a good complement for achieving the conservation objectives of Marine Protected Areas (MPAs). However, when evaluating MEABRs and MPAs as conservation and/or management tools, their impact upon parasite populations has rarely been considered, despite the fact that epidemiological theory suggests an increased susceptibility to parasitism under high population abundance. We evaluated the effects of MEABRs on the parasite abundance of Proctoeces lintoni and its impact on the growth of the host limpet Fissurella crassa in central Chile. Parasitic magnitude was higher inside MEABRs than in Open-Access Areas, and parasitized limpets showed a greater shell length, muscular foot biomass, and gonadosomatic index compared to non-parasitized limpets of the same age. Our results suggest that the life cycle of P. lintoni and, consequently, its trophic links have been strengthened inside MEABRs. The increased growth rate could reduce the time required to reach the minimum catch size and increase the reproductive and muscular output of the host population. Thus, parasitism should be considered in the conservation and management of economically important mollusk hosts.