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A uniform response of the pollen grains towards storage conditions was registered in all five shrubs investigated with a conspicuous decline in germination percentage and pollen tube length after storage. Pollen tube growth reacted more sensitively to storage than germination. The most profound reductions in pollen viability traits were observed in samples stored at + 4 °C. The germination percentage of freshly collected pollen of individual shrubs ranged between 67.3 and 88.6%, whereas that in stored pollen was between 18.0 and 39.6%. In relative terms, storage represented a 49.3–73.2% decline in germination (Fig. 1). The same tendency was also observed in pollen tube growth, when freshly collected pollen possessed 248.0–367.3 µm long pollen tubes, and pollen stored at + 4 °C was characterised by 93.9–218.5 µm long pollen tubes. The corresponding decline reached 32.5–68.7%.Figure 1Graphical illustrations of variation in pollen germination percentage (a) and pollen tube length (b) of individual shrubs revealed in fresh pollen and in pollen under storage. Different letters refer to the statistical significance of the differences between tested individuals and storage variants, resulting from Duncan’s pairwise tests.Full size imageContrary to storage at + 4 °C, pollen stored at − 20 °C had an increased germination by 0.3% in shrub no. 1 and 0.6% in shrub no. 5 as compared with fresh pollen. A more conspicuous increase in pollen germinability was registered in individual no. 4, exhibiting 70.0% germination in fresh pollen and 93.6% in pollen stored at − 20 °C. In the remaining two shrubs (no. 2, 3), only a negligible decline in pollen germination was recorded. The deviation from freshly collected pollen varied within 0.5–16.8%. In general, the germination characteristics of pollen stored at − 20 °C were comparable with those of the fresh pollen and varied between 67.6 and 93.6%. As a second viability trait, pollen tube growth deviated more profoundly from that of fresh pollen than germination. On average, the pollen tube length of pollen stored at − 20 °C ranged from 163.0 to 286.6 µm, which represents a 11.4–45.7% decline compared to fresh pollen (Figs. 1, S1). ANOVA and Duncan`s grouping confirmed the highly significant differences between tested shrubs in both pollen germination percentage (P  More

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We untangled how the direct and indirect paths of forest cover and water quality variables interact and shape anopheline assemblages in two seasons. Although previous studies determined how environmental variables at different spatial extents affected anopheline distributions in Amazônia, most studies focused on a single effect of an environmental variable or focused on single habitat types (terrestrial or aquatic)22,23,41,42. Our most important finding is that seasonality modulates the direct and indirect effects of forest cover on Amazônian anopheline larval distributions. In particular, we found that forest cover had stronger direct and indirect influence on larval anopheline assemblage composition in the rainy season than the dry season.The different paths and strengths of forest cover influences on anopheline assemblages during the rainy and dry seasons can be associated with the responses of adults and larvae to forest characteristics. Forest cover influences water quality variables of ponds by shading, organic matter inputs and erosion processes43. These effects have consequences for pond water quality44 and favor the establishment of different culicid species45. We showed that during the rainy season, forest cover directly and indirectly influenced site water quality. Greater forest cover in the rainy season directly and indirectly affected A. nimbus and the secondary malaria vectors A. triannulatus and A. braziliensis positively. In the dry season, greater forest cover positively but marginally affected A. peryassui, A. nuneztovari and A. albitarsis, but only indirectly through water quality. Some species like A. triannulatus, A. nuneztovari and A. braziliensis coexist with the malaria vector, A. darlingi, in breeding sites46, and these species have been positively associated with pH, dissolved oxygen and total suspended solids in natural and artificial habitats20,21,22,23,24,25,26,27,28,29,30,31,32,33,34,35,36,37,38,39,40,41,42,43,44,45,46,47, which are environmental conditions favored by greater forest cover. The marginal indirect effect of forest cover on anopheline assemblage in the dry season suggests that we need caution in the interpretation of this result and long-term temporal data is required to confirm if this effect is corroborated.Forest conditions influence mosquito vectors and their hosts. For example, some mosquitoes are zoophiles that feed on the blood of birds, reptiles, and mammals48, which are often more abundant in conserved areas. Other species are anthropophilic and prefer to feed on human blood49 and altered environments can force these species to migrate and, consequently, to change hosts48. In our study, A. triannulatus and A. minbus were more abundant in sites with more natural characteristics, whereas A. darlingi and A. nuneztovari were more abundant in altered landscapes. In addition, urbanization and deforestation increase the proximities of humans and domestic animals to mosquito vectors and their hosts, thereby maintaining and increasing transmission cycles50.Forest conditions influence anopheline diversity by different paths, which may alter the strength of their seasonal effects. During the dry season, mosquito survival is also affected by altered microclimate (e.g., lower humidity)51 and lentic habitats contain less water, increased nutrient concentrations and decreased abundance and richness of mosquitoes52,53. We observed that rainfall plays an important role in the larval abundance of Anopheles in artificial larval habitats in Manaus. In addition, climatic factors such as rainfall and river levels are strongly associated with vector abundance and malaria cases in the region54,55. During the rainy season, increased water volume in artificial habitats provides more areas for distribution and development of mosquito species56 and we detected a significant increase in abundance of A. triannulatus, A. darlingi and A. nuneztovari. These observations may partially explain why we found a direct effect of forest cover on mosquitoes only during the rainy season.Our results add more evidence that managing and conserving forest cover is important to control anophelines, thereby decreasing the contact of potential vectors (e.g., A. darlingi) with humans. In general, our results support the idea that mosquitoes are directly affected by the loss of native forest cover57 in the rainy season. Mosquitoes associated with serious human diseases (e.g., malaria, yellow fever, dengue, leishmaniasis) are more abundant in areas with low levels of native forest cover14,15,16,17,18,19,20,21,22,23,24,25,26,27,28,29,30,31,32,33,34,35,36,37,38,39,40,41,42,43,44,45,46,47,48,49,50,51,52,53,54,55,56,57,58. This is a critically important finding because recent studies have shown that forest cover plays an important role in the vector dynamics of mosquitoes and forest conservation keeps pathogens within the forest, avoiding spillover to human settlements59. On the other hand, deforestation provides favorable conditions for these vectors, thereby increasing malária cases and decreasing scores of the Human Development Index60. In addition, there is a positive correlation between mosquito abundance in fragmented forests and the prevalence of Plasmodium, the protozoan that causes malaria61.Artificial larval habitats promote conditions for malaria vectors in Amazônia62,63. Therefore, the best way to develop control techniques would be to understand larval ecology in these habitats, where they are more sensitive to infections by pathogens, parasites, predation, larvicides and growth regulators64. This information is necessary to minimize failures in programs to control or eradicate the vector and the disease. Under this perspective, our study adds a new piece in the puzzle of mosquito control in Amazônia. For example, during the rainy season when forest cover directly and indirectly influences larval habitats, control programs can strengthen the control of key limnological variables, habitat structure, and entomological aspects, intensifying the environmental filter, particularly in areas with little forest cover and greater human concentrations near those habitats. The limnological study of Anopheles larval habitats is still far from complete, as each case has peculiarities inherent to them. Despite attempts, anophelines demonstrate versatility in relation to abiotic parameters20,21,22,23,24,25,26,65,66. However, we can use approaches that modify the larval environments. For example, more efficient management of water levels in fish farming ponds could decrease larval numbers and anopheline reproduction, Similarly, greater rationing of fish feed would decrease the supply of food resources for mosquito larvae. It is also worth mentioning that some variables are related to the efficiency of others. Regarding biological control via entomopathogenic bacteria, environmental factors (solar radiation) and water quality (amounts of total suspended solids and organic matter), can interfere with the effectiveness of the formulated Bacillus sphaericus applied in habitats for vector control62,63,64,65,66,67. Furthermore, eutrophication decreased the assemblages of aquatic invertebrates predating mosquito larvae.Another alternative is the use of physical control (removal of grasses and macrophytes from the edge of habitats), helping to reduce microhabitats that provide larval refuges. Also, increased light and water temperature at the edges favor natural predation and biological control processes from potential fish and macroinvertebrates. The conservation of natural enemies and the use of biotic agents in the population control of vector mosquitoes have been recommended in small and medium-sized natural and artificial breeding sites19,20,21,22,23,24,25,26,27,30,31,32,33,34,35,36,37,38,39,40,41,42,43,44,45,46,47,48,49,50,51,52,53. A combination of techniques that shape the important environmental variables for the establishment of these species are essential for vector control.The analytical approach used here opens some windows of opportunity for improvements that are important to be recognized. First, our model did not incorporate important complexity of natural systems, such as ecological interactions among vectors and hosts, including human behavior. Agent-based models, including different host behavior, could provide important insights in this way. Second, our study is very limited in terms of temporal climatic variability. Additional information is needed to better understand the effects of long-term changes in land-use, water quality and climate and their interactions with mosquito assemblages in the region, particularly considering an ecological-evolutionary perspective. Third, it is important to highlight that the magnitude of effects of the estimated drivers were not the same in the rainy and dry seasons. Also, they may not remain constant in coming decades, especially considering potential regional process on mosquito assemblages, such as spillover effects, mass effects and host changes. Fourth, our study was carried out in an area of Amazonia that has experienced, a relatively old land use conversion from forest to urban areas (urban expansion rate of around 12% per year for the past 34 years)68. Beginning in the 1970s, human population increased at a rate of around 23% per decade and 25% in Manaus11. Therefore, the region we studied is very relevant in terms of historical interactions among human populations, mosquitoes and land use changes. However, understanding the effect of these changes on mosquito assemblages in areas with different land-use change dynamics, provides us with important information69, particularly those with very rapid urbanization processes, such as in the Arch of Deforestation70. Lastly, we need studies that consider the nexus among climate and land use changes, human and animal population health, economic conditions, and ecosystem services provided by these forest-urban transitional regions. Such information would facilitate including mosquito information in land use planning and climate mitigation programs based on forest management in and around cities.Therefore, identifying ecological factors and paths that affect the composition of species of epidemiological importance are essential because they inform vector integrated management strategies. We emphasize that larval control in lentic habitats requires knowledge about larval ecology and the effects of biotic and abiotic variables on larvae, especially when it comes to biological controls. The application of integrated pest management can be conducted in both dry and rainy seasons. However, we recommend focusing on the dry season when larval habitats are more limited, in smaller volumes and more accessible for entry and application of vector control techniques. These are critically important considerations because over 2 million people live in Amazonas state11 and anophelines transmitted over 59,637 malaria cases in the Amazon region in the first half of 2020, and about 44.4% came from the state of Amazonas71. More

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