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Protist plankton can be divided into three main groups: phytoplankton, zooplankton, and mixoplankton.In situmethods for studying phytoplankton and zooplankton are relatively straightforward since they generally target chlorophyll/photosynthesis or grazing activity, while the integration of both processes within a single cell makes mixoplankton inherently challenging to study. As a result, we understand less about mixoplankton physiology and their role in food webs, biogeochemical cycling, and ecosystems compared to phytoplankton and zooplankton. In this paper, we posit that by merging conventional techniques, such as microscopy and physiological data, with innovative methods likein situsingle-cell sorting and omics datasets, in conjunction with a diverse array of modeling approaches ranging from single-cell modeling to comprehensive Earth system models, we can propel mixoplankton research into the forefront of aquatic ecology. We present eight crucial research questions pertaining to mixoplankton and mixotrophy, and briefly outline a combination of existing methods and models that can be used to address each question. Our intent is to encourage more interdisciplinary research on mixoplankton, thereby expanding the scope of data acquisition and knowledge accumulation for this understudied yet critical component of aquatic ecosystems. 

Abstract Phago-mixotrophy, the combination of photoautotrophy and phagotrophy in mixoplankton, organisms that can combine both trophic strategies, have gained increasing attention over the past decade. It is now recognized that a substantial number of protistan plankton species engage in phago-mixotrophy to obtain nutrients for growth and reproduction under a range of environmental conditions. Unfortunately, our current understanding of mixoplankton in aquatic systems significantly lags behind our understanding of zooplankton and phytoplankton, limiting our ability to fully comprehend the role of mixoplankton (and phago-mixotrophy) in the plankton food web and biogeochemical cycling. Here, we put forward five research directions that we believe will lead to major advancement in the field: (i) evolution: understanding mixotrophy in the context of the evolutionary transition from phagotrophy to photoautotrophy; (ii) traits and trade-offs: identifying the key traits and trade-offs constraining mixotrophic metabolisms; (iii) biogeography: large-scale patterns of mixoplankton distribution; (iv) biogeochemistry and trophic transfer: understanding mixoplankton as conduits of nutrients and energy; and (v) in situ methods: improving the identification of in situ mixoplankton and their phago-mixotrophic activity. 

Abstract Mixotrophic protists combine photosynthesis and phagotrophy to obtain energy and nutrients. Because mixotrophs can act as either primary producers or consumers, they have a complex role in marine food webs and biogeochemical cycles. Many mixotrophs are also phenotypically plastic and can adjust their metabolic investments in response to resource availability. Thus, a single species's ecological role may vary with environmental conditions. Here, we quantified how light and food availability impacted the growth rates, energy acquisition rates, and metabolic investment strategies of eight strains of the mixotrophic chrysophyte,Ochromonas. All eightOchromonasstrains photoacclimated by decreasing chlorophyll content as light intensity increased. Some strains were obligate phototrophs that required light for growth, while other strains showed stronger metabolic responses to prey availability. When prey availability was high, all eight strains exhibited accelerated growth rates and decreased their investments in both photosynthesis and phagotrophy. Photosynthesis and phagotrophy generally produced additive benefits: In low‐prey environments,Ochromonasgrowth rates increased to maximum, light‐saturated rates with increasing light but increased further with the addition of abundant bacterial prey. The additive benefits observed between photosynthesis and phagotrophy inOchromonassuggest that the two metabolic modes provide nonsubstitutable resources, which may explain why a tradeoff between phagotrophic and phototrophic investments emerged in some but not all strains. 

Abstract AimCoffee is an important export for many developing countries, with a global annual trade value of $100 billion, but it is threatened by a warming climate. Shade trees may mitigate the effects of climate change through temperature regulation that can aid in coffee growth, slow pest reproduction, and sustain avian insectivore diversity. The impact of shade on bird diversity and microclimate on coffee farms has been studied extensively in the Neotropics, but there is a dearth of research in the Paleotropics. LocationEast Africa. MethodsWe created current and future regional Maxent models for avian insectivores in East Africa using Worldclim temperature data and observations from the Global Biodiversity Information Database. We then adjusted current and future bioclimatic layers based on mean differences in temperature between shade and sun coffee farms and projected the models using these adjusted layers to predict the impact of shade tree removal on climatic suitability for avian insectivores. ResultsExisting Worldclim temperature layers more closely matched temperatures under shade trees than temperatures in the open. Removal of shade trees, through warmer temperatures alone, would result in reduction of avian insectivore species by over 25%, a loss equivalent to 50 years of climate change under the most optimistic emissions scenario. Under the most extreme climate scenario and removal of shade trees, insectivore richness is projected to decline from a mean of 38 to fewer than 8 avian insectivore species. Main conclusionsWe found that shade trees on coffee farms already provide important cooler microclimates for avian insectivores. Future temperatures will become a regionally limiting factor for bird distribution in East Africa, which could negatively impact control of coffee pests, but the effect of climate change can be potentially mediated through planting and maintaining shade trees on coffee farms.