The influence of upwelling on key bivalves from the Humboldt and Iberian current systems.

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.

Upwelling enhances seaweed nutrient quality, altering feeding behavior and growth rates in an intertidal sea urchin, Loxechinus albus.

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.

Parasitism by metacercariae modulates the morphological, organic and mechanical responses of the shell of an intertidal bivalve to environmental drivers.

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 pCO2, 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.

Morphological, physiological and behavioral responses of an intertidal snail, Acanthina monodon (Pallas), to projected ocean acidification and cooling water conditions in upwelling ecosystems.

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 pCO2 (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 pCO2 or the interaction between pCO2 and temperature. Instead, metabolic rates were driven by pCO2, but a significant interaction with temperature suggests that in cooler conditions, metabolic rates will increase when associated with high pCO2 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.

Ocean acidification alters anti-predator responses in a competitive dominant intertidal mussel.

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 pCO2 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.

Exposure to artificial light at night (ALAN) alters RNA:DNA ratios in a sandy beach coleopteran insect.

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.

The combined effects of ocean acidification and warming on a habitat-forming shell-crushing predatory crab.

In mid rocky intertidal habitats the mussel Perumytilus purpurarus monopolizes the substratum to the detriment of many other species. However, the consumption of mussels by the shell-crushing crab Acanthocyclus hassleri creates within the mussel beds space and habitat for several other species. This crab uses its disproportionately large claw to crush its shelled prey and plays an important role in maintaining species biodiversity. This study evaluated the consequences of projected near-future ocean acidification (OA) and warming (OW) on traits of A. hassleri linked with their predatory performance. Individual A. hassleri were maintained for 10-16 weeks under contrasting pCO2 (~500 and 1400 µatm) and temperature (~15 and 20 °C) levels. We compared traits at the organismal (oxygen consumption rate, survival, calcification rate, feeding rates, crusher claw pinching strength, self-righting speed, sarcomere length of the crusher claw muscles) and cellular (nutritional status ATP provisioning capacity through citrate synthase activity, expression of HSP70) level. Survival, calcification rate and sarcomere length were not affected by OA and OW. However, OW increased significantly feeding and oxygen consumption. Pinching strength was reduced by OA; meanwhile self-righting was increased by OA and OW. At 20 °C, carbohydrate content was reduced significantly by OA. Regardless of temperature, a significant reduction in energy reserves in terms of protein content by OA was found. The ATP provisioning capacity was significantly affected by the interaction between temperature and pCO2 and was highest at 15 °C and present day pCO2 levels. The HSP70 levels of crabs exposed to OW were higher than in the control crabs. We conclude that OA and OW might affect the amount and size of prey consumed by this crab. Therefore, by reducing the crab feeding performance these stressors might pose limits on their role in generating microhabitat for other rocky intertidal species inhabiting within mussel beds.

Geographical variability and parasitism on body size, reproduction and shell characteristics of the keyhole limpet Fissurella crassa (Mollusca: Vetigastropoda).

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).

Seawater-temperature and UV-radiation interaction modifies oxygen consumption, digestive process and growth of an intertidal fish.

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.

Increased parasitism of limpets by a trematode metacercaria in fisheries management areas of central Chile: effects on host growth and reproduction : management areas and parasitism.

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.