The interplay of key abiotic factors with the occurrence and abundance of mosquito larvae in bromeliads

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This study investigates the effect of abiotic factors, specifically temperature, rainfall, water volume, and physicochemical parameters, on the occurrence and abundance of mosquitoes developing in bromeliads within the Brazilian Atlantic Forest. Larval samples and abiotic parameters were collected from nine bromeliads between 2015 and 2017 in a forest remnant in São Paulo, Brazil. Relationships between variables were tested using generalized linear mixed-effects models and structural equation modeling. Results indicate that rainfall and temperature influence the water volume of bromeliad tanks, which in turn affect physicochemical parameters, particularly pH. These variations affect the richness, abundance, and occurrence of mosquito species, including the vector of malaria parasites, Anopheles cruzii . The study highlights a cascade effect where abiotic factors alter the microhabitat conditions, thereby affecting mosquito populations. Understanding these interactions is crucial for predicting the impacts of climate change on mosquito-borne diseases in tropical regions. Biological sciences/Ecology/Climate change ecology Biological sciences/Ecology/Ecological epidemiology Biological sciences/Ecology/Ecological genetics Biological sciences/Ecology/Freshwater ecology Health sciences/Diseases/Infectious diseases/Malaria Mosquitoes Bromeliads Climate change Cascading effect Anopheles cruzii Figures Figure 1 Figure 2 Figure 3 INTRODUCTION Abiotic factors are the non-living components of the ecosystem that influence the survival, growth, reproduction and behavior of organisms [1]. Temperature and rainfall are abiotic components that significantly influence aquatic organisms, particularly those in lentic environments, such as insect larvae [2, 3]. While temperature variation is the main factor responsible for structuring aquatic insect assemblages in temperate environment [4], in tropical regions, where temperature varies less, rainfall can play a more influential role by altering both the availability of sites for colonization and the physicochemical factors of the environment [5, 6]. For immature forms of mosquitoes (Diptera: Culicidae), the physicochemical conditions of water are important constraints that influence their survival, growth and adaptation [7, 8, 9]. Seasonal patterns or extreme weather events that alter rainfall and temperature can lead to changes in the physicochemical parameters of lentic environments where mosquitoes breed. For example, dry and warmer periods favor the loss of water by evaporation, reducing the water body volume and leading to an increase of insolation and ionic concentration with changes in primary production that affect pH, salinity and dissolved oxygen [5, 10]. While pH and salinity directly impact the ionic and osmotic regulation of mosquitoes, dissolved oxygen primarily affects the associated microfauna and flora in breeding sites [11, 12]. Bromeliads (Bromeliaceae) are a family of neotropical plants with more than 3,700 described species [13]. Most bromeliads are epiphytic or terrestrial, with long leaves arranged in a rosette shape that form tanks capable of retaining water and nutrients, thereby supporting the development of biotic communities [14]. In the Brazilian Atlantic Forest, there is a high diversity of bromeliads that harbor many species of aquatic insects, including immature forms of mosquitoes, some of which have epidemiological importance, such as Anopheles ( Kerteszia ) cruzii , the main vector of malaria parasites in this region [15, 16, 17]. Given that changes in temperature and rainfall patterns driven by climate change are key factors that will impact the biodiversity of freshwater communities in the coming decades [18], we hypothesize that variations in temperature and precipitation affect the volume of bromeliad water tanks, thereby promoting changes in the physicochemical parameters of the water, which in turn influence the patterns of occurrence, abundance, and richness of mosquito assemblages developing in these aquatic ecosystems. To investigate the effect of these abiotic factors on mosquito assemblages in bromeliads, we conducted a study with nine bromeliads in a protected area of the Atlantic Forest between 2015 and 2017, collecting larval samples and measuring abiotic parameters. The data obtained demonstrate the effect of average maximum temperature and accumulated rainfall on the water volume of the bromeliad tanks and the effect of this variation on the pH, which in turn, either alone or in interaction with salinity, influenced the occurrence, abundance, and richness of mosquitoes in these microhabitats. METHODS The study was performed in the Capivari-Monos Environmental Protection Area (EPA), a remnant of Atlantic Forest situated in the southern region of the city of São Paulo, Brazil. The EPA extends over the first hills near the crest of the Serra do Mar Mountain range at altitudes varying from 740 to 850 m above sea level. The climate is predominantly super-humid, oceanic, and tropical with average annual temperatures of around 19°C and rainfall of between 1600 and 2200 mm [19]. A total of ten field collections were carried out between November 2015 and April 2017 (Nov-Dec/2015, Jan-Oct-Nov-Dec/2016, Jan-Feb-Mar-Apr/2017). Nine bromeliads from three different sites in the EPA were selected and monitored throughout the study: four bromeliads at Cachoeira (C1, C2, C3, and C4), a private property adjacent to a waterfall with a visitation area (23° 56.378′ S / 46°41. 659′ W); two bromeliads at Siloé (S1 and S2), a private property near Marsilac village, constituting a transitional area between a rural environment and the EPA forest (23° 54.556′ S / 46° 42.167′ W); and three bromeliads at Borracharia (B1, B2, and B3), situated in a small village surrounded by the EPA forest (23° 54.395′ S / 46° 42.486′ W) (Fig. 1 ). All the bromeliads were epiphytes (ranging from one to seven meters high) and were identified as Vriesia spp. (Bromeliaceae: Tillandsioideae). At each collection, the water content present in each bromeliad was extracted using suction samplers and examined. The total volume of water, pH, salinity (practical salinity unit - PSU), and dissolved oxygen (parts per million - PPM) were measured for each sample using a multiparameter meter (Hanna® HI-9828). The specimens present were separated using pipettes and stored in plastic pots for later identification. After this, the remaining water content was returned to the bromeliad to minimize interference with its biotic community. The collected immature forms (larvae and pupae) were transported to the Entomology Laboratory, School of Public Health, University of São Paulo (LESP/FSP/USP), where they were monitored until the adult stage, after which the adults were identified morphologically. Daily rainfall and temperature data for the study sites were obtained from the Centro de Gerenciamento de Emergências de São Paulo (CGE). For temperature, the average maximum and minimum (ºC) for the 30 days prior to collection were considered. For rainfall, the accumulated rainfall (mm³) for the 30 days prior to collection was considered. The average maximum and minimum temperatures were highly correlated, so only the average maximum temperature was kept as a predictor variable, as it demonstrated a superior fit in the models. To analyze the data, a cascade effect hypothesis was tested, in which an initial disturbance (variation in rainfall and/or temperature) affects other components in a sequence of causally connected processes [1, 20, 21, 22]. First, the effect of accumulated rainfall and average monthly temperatures on the volume of water in the bromeliad tanks was analyzed. Next, the relationship between water volume and the variation in physicochemical parameters was analyzed. Finally, the relationship between physicochemical parameters and the occurrence, richness, and abundance of mosquitoes in the bromeliads was investigated. Direct effects of precipitation and temperature on physicochemical parameters and water volume on mosquitoes were also explored. The relationships between the variables were verified using generalized linear mixed-effect models (GLMM) with Gaussian, binomial, or Poisson error distributions, depending on the type of response variable. The different sites and bromeliads studied were considered as random effects. To fit the best model, the backward elimination strategy was used, starting with the full model (fixed effect variables + fixed effect interactions + random effect) and eliminating the least significant fixed effect variables until the best fit was obtained. To investigate the relationship between abiotic variables and the occurrence of mosquitoes, only species that occurred five or more times during the study period were considered. Finally, we performed a path analysis using piecewise structural equation modeling (piecewise SEM) [23] to investigate the direct and indirect effects of the abiotic variables on species richness, total abundance, and occurrence of An. cruzii in bromeliads. The goodness-of-fit of piecewise SEM models was evaluated using Fisher’s C test [24]. The values of C can be compared to a chi-squared distribution with 2k degrees of freedom, and values of p above the chosen significance threshold (α = 0.05) indicate a good fit of the model (the hypothesized relationships are consistent with the data). The R computer environment [25] was used to analyze the data using the packages lme4 [26], ggplot2 [27], sjPlot [28], and piecewiseSEM [23]. RESULTS Throughout the study period, the accumulated rainfall for the 30 days prior to collection ranged from 61 to 429 (mm³) while the temperature for the 30 days prior to collection ranged from 21.6 to 29.5ºC for the average daily maximum and from 13.3 to 18.9ºC for the average daily minimum. The volume of water sampled per bromeliad ranged from 95 to 1,050 ml. For the physicochemical parameters, pH ranged from 3.57 to 6.02, salinity from 0.006 to 0.36 PSU and dissolved oxygen from 0.2 to 8.56 ppm. A total of 529 individuals belonging to 20 species were collected, including specimens of An. cruzii and specimens of the genera Culex (mostly of the subgenus Microculex ) and Wyeomyia of the subgenus Phoniomyia . The number of species per bromeliad ranged from 7 to 14 (bromeliads B3 and S1, respectively). The most abundant species were those of the subgenus Microculex , especially Cx. pleuristriatus with 129 individuals collected. In terms of occurrence, only 10 species occurred five or more times throughout the study, with Cx. imitator and Wy. davisi being the most frequent (29 and 19 times, respectively) and the only ones to be present in all the bromeliads investigated (Table 1 ). Table 1 Number of individuals and occurrences per mosquito species found in nine bromeliads investigated at Capivari-Monos EPA between November 2015 and April 2017. Species Bromeliad Total individuals Number of occurrences B1 B2 B3 C1 C2 C3 C4 S1 S2 Anopheles (Ker.) cruzii 7 0 1 0 3 3 6 1 3 24 13 Culex (Cux.) eduardoi 0 0 3 0 0 0 0 0 0 3 1 Culex (Mcx.) albipes 10 7 17 8 0 4 0 32 5 83 17 Culex (Mcx.) daumasturus 5 2 0 0 0 0 0 2 0 9 4 Culex (Mcx.) dubitans 2 0 0 0 0 5 2 2 1 12 7 Culex (Mcx.) fuscatus 0 0 0 0 0 2 0 2 6 10 4 Culex (Mcx.) imitator 8 7 6 8 2 21 14 4 2 72 29 Culex (Mcx.) inimitabilis 0 0 0 0 0 0 0 2 2 4 2 Culex (Mcx.) neglectus 1 0 4 0 1 2 1 3 0 12 9 Culex (Mcx.) pleuristriatus 19 31 73 0 0 4 1 1 0 129 16 Culex (Mcx.) worontzowi 1 5 0 1 4 7 2 7 0 27 14 Culex ocellatus 3 0 0 0 0 0 2 0 0 5 2 Wyeomyia (Pho.) quasilongirostris 0 0 0 0 0 0 0 2 0 2 1 Wyeomyia (Pho.) davisi 11 9 5 5 4 9 3 5 1 52 19 Wyeomyia (Pho.) edwardsi 0 0 0 1 0 0 0 0 0 1 1 Wyeomyia (Pho.) incaudata 2 1 0 1 1 4 5 6 0 20 11 Wyeomyia (Pho.) pallidoventer 0 0 0 0 3 0 0 0 0 3 1 Wyeomyia (Pho.) pilicauda 2 0 0 3 0 0 0 0 0 5 2 Wyeomyia (Pho.) splendida 0 0 0 0 0 4 0 0 0 4 1 Wyeomyia (Pho.) theobaldi 5 5 0 5 6 10 0 19 2 52 17 Total individuals 76 67 109 32 24 75 36 88 22 529 Total species 13 8 7 8 8 12 9 14 8 20 Regarding the best-fit models, the averages of maximum temperature and accumulated rainfall showed a significant influence on the volume of water in the bromeliads, with rainfall positively related and temperature negatively related to the water volume (Table 2 and Fig. 2 A). Rainfall was also positively related to pH, while temperature was negatively related to dissolved oxygen. The volume of water was positively related to the increase in pH (Fig. 2 B) and dissolved oxygen, while the increase in volume was negatively related to the abundance of mosquitoes in the bromeliads (Table 2 ). No significant relationship was found between water volume and the occurrence probability of any mosquito species. Both the richness and abundance of mosquitoes varied as a function of pH, salinity, and the interaction between these two physicochemical parameters. For lower salinities (predicted to 0.025 and 0.05 PSU), richness and abundance increased as a function of increasing pH, while for higher salinities (predicted to 0.1 PSU), richness and abundance decreased in response to increasing pH (Table 2 and Fig. 2 C). In relation to the probability of species occurrence, pH was the parameter most associated with the presence of five of the ten species tested, including the vector An. cruzii , which showed a positive association with pH (Table 2 and Fig. 2 D). Table 2 Models that showed the best fit in the study. The intercept and slope values refer to the fixed effect parameters of the GLMMs. The standard error of the estimate is shown in brackets. Explanatory variables Response variables Intercept Slope p Rain + Temp_max Water volume 786.7 (246.2) 0.674 (0.216) 0.003 -22.93 (10.78) 0.04 Rain pH 4.35 (0.14) 0.002 (0.0005) 0.0005 Temp_max Dissolved oxygen 9.46 (2.25) -0.241 (0.09) 0.007 Water volume pH 4.36 (0.157) 0.001 (0.0004) 0.002 Water volume Dissolved oxygen 1.82 (0.56) 0.004 (0.001) 0.007 Water volume Total abundance 2.32 (0.15) -0.0007(0.0003) 0.037 pH An. cruzii -11.04 (4.35) 1.938 (0.846) 0.022 pH Cx. albipes -12.34 (4.88) 2.36 (1) 0.018 Salinity 166.42 (83.6) 0.046 pH:Salinity -36.25 (18.68) 0.052 pH Cx. dubitans -24.23 (10.53) 4.26 (1.94) 0.028 pH Cx. neglectus -11.61 (4.3) 1.963 (0.832) 0.018 pH Wy. incaudata 7.06 (3.85) -1.824 (0.835) 0.029 pH Species richness -1.38 (1.02) 0.521 (0.2) 0.013 Salinity 28.5 (16.1) 0.075 pH:Salinity -6.65 (3.53) 0.059 pH Total abundance 0.017 (0.63) 0.477 (0.13) 0.0002 Salinity 30.73 (10.54) 0.003 pH:Salinity -7.6 (2.35) 0.001 Piecewise SEM analysis indicated that the data were satisfactorily fitted for the three models tested: species richness (Fisher’s C = 13.5, df = 12, p = 0.33), total abundance (Fisher’s C = 10, df = 8, p = 0.26), and An. cruzii presence (Fisher’s C = 4.6, df = 8, p = 0.8). Rainfall had direct effects on water volume, pH, and total abundance, while temperature had direct effects on water volume and total abundance. The model for total abundance showed a high explanatory power (R² = 0.63), followed by An. cruzii presence (R² = 0.32), and species richness (R² = 0.13) (Fig. 3 ). DISCUSSION The observed results illustrate the interaction between abiotic factors and the occurrence and abundance of mosquito larvae in bromeliads, supporting the hypothesis of a cascade effect where variations in rainfall and temperature lead to changes in the water volume of bromeliad tanks, which in turn influence physicochemical parameters and subsequently affect the richness, abundance, and occurrence of mosquitoes in these microhabitats. While rainfall increases the volume of water retained in the bromeliads, rising temperatures decrease this volume due to greater evaporation. As the volume of water in the bromeliad tanks increases, there is a tendency for pH to increase. In turn, the variation in pH influences the occurrence of some species and, when associated with salinity, induces variations in the richness and abundance of mosquitoes in the bromeliads. Rainfall and temperature have been shown to have diverse effects on the biotic communities of bromeliads. Previous studies have observed that changes in the hydrological regime, particularly extreme events leading to droughts or prolonged rains, affect the food web [29], the relative abundance and biomass of functional groups of invertebrates [30, 31], and the biomass and composition of microorganisms [32, 33, 34]. For mosquitoes, studies conducted in the Atlantic Forest of southeastern Brazil have shown a positive relationship between rainfall, water volume in bromeliad tanks and the presence of some species, including An. cruzii and mosquitoes of the genera Culex and Wyeomyia [17, 35]. In a study conducted in a semi-arid region of northeastern Brazil, it was found that the probability of mosquitoes being present in bromeliads was inversely proportional to the average temperature for the 30 days prior to collection, because under high temperatures there was greater evaporation of water and the bromeliad tanks dried out more quickly [36]. As observed in this study, pH was a key factor associated with changes in the richness, abundance, and occurrence of some species. The water in the bromeliad tank tends to have low pH values and this has been associated with acids resulting from microbial activity [37], leaf litter accumulated in the tanks [38], accumulation of carbonic acid due to CO2 produced in respiration by microbes [39], and the activity of proton pumps in the leaves as mechanisms of tank water acidification [40]. In a study conducted in Costa Rica, it was observed that the average pH of tank water was lower than that of rainwater for six of the eight bromeliad species investigated [40]. Our data corroborates these studies, indicating that the water present in bromeliad ponds tends to be acidic and that the increase in the volume of water in the ponds due to precipitation induces a reduction in the concentration of H + ions. Interestingly, the interaction between increased pH and salinity was observed to reduce the abundance and richness of mosquitoes in bromeliads, suggesting a suboptimal environment for species that colonize this ecosystem. The mosquito species present in bromeliads seem to have some degree of predilection for more acidic environments with low salinity [41]. As for physiological factors, the homeostasis mechanisms that allow mosquito larvae to maintain the ionic composition and pH of the hemolymph are still poorly understood, although it is known that ions and acid-base equivalents are absorbed or excreted directly between the hemolymph and the surroundings through ion-permeable epithelia [42]. The anal papillae appear to function as the main organ of ion exchange, absorbing Na+, Cl−, and K + and excreting the acid-base equivalents H + , NH4 + , and HCO3 − [43]. More than 200 mosquito species have been described inhabiting bromeliads in the Neotropical region, many of which are bromeliad specialists [14, 35]. An example of a specialist is the Anopheles of the subgenus Kerteszia , particularly An. cruzii , which is primarily responsible for maintaining the circulation of malaria plasmodia between humans and non-human primates in the Atlantic Forest [44, 45]. For this reason, the autochthonous malaria that occurs in this region is also known as “Bromeliad-malaria”. In the mid-1940s, the southeastern and southern regions of Brazil were considered endemic areas for bromeliad malaria, with an annual incidence of close to 4,000 cases per 100,000 inhabitants [46]. Once it was confirmed that Kerteszia mosquitoes were the transmitters of malaria in the Atlantic Forest, various control efforts were simultaneously adopted. The measures consisted of applying larvicides and insecticides, treating patients with antimalarials, destroying bromeliads by removing them or using herbicides, and even deforesting areas close to transmission hotspots [47]. Currently, bromeliad malaria is hypoendemic in the region and few cases have been recorded annually. Data from the Centro de Vigilância Epidemiológica of the State of São Paulo show that between January 2017 and November 2024, 77 autochthonous cases of malaria were confirmed in the State [48]. Climate change is expected to have profound effects on freshwater communities, particularly by altering hydrological cycles, temperature regimes, and nutrient dynamics [49]. These environmental shifts can significantly impact mosquito populations, as changes in rainfall and temperature influence breeding site availability, larval development rates, and adult survival [50]. For instance, increased temperatures can accelerate the life cycle of vectors such as Aedes aegypti and Culex quinquefasciatus , leading to higher population densities and prolonged transmission seasons for arboviruses, including dengue, Zika, West Nile virus, and Chikungunya [51]. Similarly, malaria transmission, primarily driven by Anopheles mosquitoes, is also projected to shift due to climate change. Studies have shown that regions in East Africa and South America may become increasingly suitable for malaria transmission, while some highly endemic areas could experience declines due to excessive warming [52]. Additionally, extreme weather events, such as floods and droughts, may expand mosquito habitats into previously unsuitable areas, facilitating the introduction of vector-borne diseases into new ecological and epidemiological contexts [53]. These complex interactions highlight the need for investigating the interplay between abiotic factors and mosquitoes’ biodiversity. Understanding these dynamics is crucial for predicting the impacts of climate change on mosquito-borne diseases and for developing effective strategies to mitigate the associated public health risks. This study has provided evidence of how changes in precipitation and temperature promote a cascade effect that leads to changes in the richness, abundance, and composition of mosquitoes, using the microcosm of bromeliads as a model. Given this evidence, it is necessary for future studies to seek to understand how ongoing climate change, associated with land use change and loss of biodiversity, will affect the occurrence, distribution, and abundance of mosquitoes, and the impacts of these changes on human health and ecosystems in tropical regions of the planet. Declarations Competing interests The authors declare no competing interests. Author Contribution A.R.M-S: Conceptualization, Investigation, Methodology, Visualization, Formal analysis, and Writing - Original Draft. R. O-C: Conceptualization, Investigation and Data-curation. W. C-J: Investigation and Data-curation. K.M.B-N: Investigation. E.E: Investigation. R.W-S: Investigation. L.F.M: Conceptualization and Investigation. M.B.P: Data-curation. M.T.M: Conceptualization, Project administration, Funding acquisition and Supervision. All authors reviewed the manuscript. Acknowledgement We thank the State of São Paulo Research Foundation (FAPESP Grants Nos. 2014/50444-5, 2023/11212-0 and 2023/17044-2) and the National Council for Scientific and Technological Development, Brazil (CNPq 309872/2021-9) for providing financial support. We would also like to express our gratitude to the field and laboratory teams at the Department for the Control of Endemic Diseases, São Paulo Zoonosis Control Center, and the School of Public Health, University of São Paulo: Ana Maria Ribeiro de Castro Duarte, João Carlos do Nascimento, Paulo Frugoli dos Santos, Luis Milton Bonafé, Antônio Waldomiro de Oliveira, Laércio Molinari, Gabriel Marcelino Neto, Luiz Sposito Jr, Renildo Souza Teixeira, Daniel Pagotto Vendrami, Gabriela Cristina de Carvalho, and Amanda Alves Camargo. Data Availability The datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request. 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Bacterial communities as bioindicators of climate change in freshwater ecosystems: Tank bromeliads as model systems. Ecological Indicators, 171, 113161. Cardoso, C. A. A., Lourenço-de-Oliveira, R., Codeço, C. T., & Motta, M. A. (2015). Mosquitoes in bromeliads at ground level of the Brazilian Atlantic Forest: the relationship between mosquito fauna, water volume, and plant type. Annals of the Entomological Society of America, 108(4), 449–458. Marteis, L. S., Natal, D., Sallum, M. A. M., Medeiros-Sousa, A. R., & La Corte, R. (2017). Mosquitoes of the Caatinga: 2. Species from periodic sampling of bromeliads and tree holes in a dry Brazilian forest. Acta Tropica, 171, 114–123. Jocque, M., & Kolby, J. E. (2012). Acidity of tank bromeliad water in a cloud forest, Cusuco National Park, Honduras. International Journal of Plant Physiology and Biochemistry, 4(4), 59–70. Laessle, A. M. (1961). A micro-limnological study of Jamaican bromeliads. Ecology, 42(3), 499–517. Benzing, D. H., Derr, J. A., & Titus, J. E. (1972). The water chemistry of microcosms associated with the bromeliad Aechmea bracteata. American Midland Naturalist, 60–70. North, Gretchen B., et al. Acid waters in tank bromeliads: Causes and potential consequences. American Journal of Botany 110.1 (2023): e16104. Multini, L. C., Oliveira-Christe, R., Medeiros-Sousa, A. R., Evangelista, E., Barrio-Nuevo, K. M., Mucci, L. F., ... & Marrelli, M. T. (2021). The influence of the pH and salinity of water in breeding sites on the occurrence and community composition of immature mosquitoes in the Green Belt of the city of São Paulo, Brazil. Insects, 12(9), 797. Durant, A. C., & Donini, A. (2018). Evidence that Rh proteins in the anal papillae of the freshwater mosquito Aedes aegypti are involved in the regulation of acid–base balance in elevated salt and ammonia environments. Journal of Experimental Biology, 221(23), jeb186866. Donini, A., & O'Donnell, M. J. (2005). Analysis of Na+, Cl-, K+, H + and NH4 + concentration gradients adjacent to the surface of anal papillae of the mosquito Aedes aegypti: application of self-referencing ion-selective microelectrodes. Journal of Experimental Biology, 208(4), 603–610. Deane, L. M. (1992). Simian malaria in Brazil. Memórias do Instituto Oswaldo Cruz, 87, 1–20. Buery, J. C., Alencar, F. E. C. D., Duarte, A. M. R. D. C., Loss, A. C., Vicente, C. R., Ferreira, L. M., ... & Cerutti Junior, C. (2021). Atlantic Forest Malaria: A review of more than 20 years of epidemiological investigation. Microorganisms, 9(1), 132. Deane, L. M. Malaria studies and control in Brazil. The American journal of tropical medicine and hygiene, v. 38, n. 2, p. 223–230, 1988. Barata, J. M. S. (1973). Estado atual da bromélia-malária no Brasil. (Master Dissertation) – School of Public Health, University of São Paulo, São Paulo, 1973. CVE – Centro de Vigilância Epidemiológica. (2024). Casos Autóctones de Malária no Estado de São Paulo. Available in https://www.saude.sp.gov.br/resources/cve-centro-de-vigilancia-epidemiologica/areas-de-vigilancia/doencas-de-transmissao-por-vetores-e-zoonoses/dados/malaria/malaria_dados.pdf. Accessed: Fev 6 2025. Woodward, G., Perkins, D. M., & Brown, L. E. (2010). Climate change and freshwater ecosystems: impacts across multiple levels of organization. Philosophical Transactions of the Royal Society B: Biological Sciences, 365(1549), 2093–2106. Rocklöv, J., & Dubrow, R. (2020). Climate change: An enduring challenge for vector-borne disease prevention and control. Nature Immunology, 21(5), 479–483. Mordecai, E. A., Ryan, S. J., Caldwell, J. M., Shah, M. M., & LaBeaud, A. D. (2020). Climate change could shift disease burden from malaria to arboviruses in Africa. The Lancet Planetary Health, 4(9), e416-e423. Caminade, C., Kovats, S., Rocklov, J., Tompkins, A. M., Morse, A. P., Colón-González, F. J., ... & Lloyd, S. J. (2014). Impact of climate change on global malaria distribution. Proceedings of the National Academy of Sciences, 111(9), 3286–3291. Caminade, C., McIntyre, K. M., & Jones, A. E. (2019). Impact of recent and future climate change on vector-borne diseases. Annals of the New York Academy of Sciences, 1436(1), 157–173. Additional Declarations No competing interests reported. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6184497","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":431568278,"identity":"8237b7ff-bf16-4026-915a-7374d91e47da","order_by":0,"name":"Antônio Ralph Medeiros-Sousa","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA0UlEQVRIiWNgGAWjYHACxgOMDQwM/CBmQgGResBaJBtAWgxI0WJwAMQkRgv/7MMPDhfu2GZvfH514ocHBgzy/GIH8GuROJdmcHjmmduJ22683SwBdJjhzNkJBKw5w2BwmLftdoLZjbMbQFoSDG4T0CJ/hv0DSIu98Yyzm38QpcXgDA/YFsYN/L3biLPF8AxPweGZbbcTZ9zg3WaRYCBB2C9yZ9g3Pi4EOoy//+zmmz8qbOT5pQloAQFmMCkBVilBWDlCC/8B4lSPglEwCkbByAMAlZdKuDH8AQoAAAAASUVORK5CYII=","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":true,"prefix":"","firstName":"Antônio","middleName":"Ralph","lastName":"Medeiros-Sousa","suffix":""},{"id":431568279,"identity":"fcb78951-121a-4a6c-bb6c-d7205da27724","order_by":1,"name":"Rafael Oliveira-Christe","email":"","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Rafael","middleName":"","lastName":"Oliveira-Christe","suffix":""},{"id":431568280,"identity":"0b6f539b-942d-4ea4-acbc-d4c5c18bfe35","order_by":2,"name":"Walter Ceretti-Junior","email":"","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Walter","middleName":"","lastName":"Ceretti-Junior","suffix":""},{"id":431568281,"identity":"c70c01d1-a136-4d47-acae-3aa7dcded8bf","order_by":3,"name":"Karolina Morales Barrio-Nuevo","email":"","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Karolina","middleName":"Morales","lastName":"Barrio-Nuevo","suffix":""},{"id":431568282,"identity":"bc44ad3d-ff47-4a68-8a30-f1d85017d955","order_by":4,"name":"Eduardo Evangelista","email":"","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Eduardo","middleName":"","lastName":"Evangelista","suffix":""},{"id":431568283,"identity":"373c422a-088a-47d6-a3a8-3f679bc0fd76","order_by":5,"name":"Ramon Wilk-da-Silva","email":"","orcid":"","institution":"Instituto Butantan","correspondingAuthor":false,"prefix":"","firstName":"Ramon","middleName":"","lastName":"Wilk-da-Silva","suffix":""},{"id":431568284,"identity":"49ad3143-d5a3-447f-b3b6-c40d035f606d","order_by":6,"name":"Luis Filipe Mucci","email":"","orcid":"","institution":"São Paulo State Department of Health","correspondingAuthor":false,"prefix":"","firstName":"Luis","middleName":"Filipe","lastName":"Mucci","suffix":""},{"id":431568285,"identity":"93a447a0-4264-40d5-a9ce-b4bc26288018","order_by":7,"name":"Márcia Bicudo Paula","email":"","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Márcia","middleName":"Bicudo","lastName":"Paula","suffix":""},{"id":431568286,"identity":"b843b8ff-5980-430e-88d5-1fee04984963","order_by":8,"name":"Mauro Toledo Marrelli","email":"","orcid":"","institution":"Universidade de São Paulo","correspondingAuthor":false,"prefix":"","firstName":"Mauro","middleName":"Toledo","lastName":"Marrelli","suffix":""}],"badges":[],"createdAt":"2025-03-08 14:08:04","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6184497/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6184497/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1038/s41598-025-15514-7","type":"published","date":"2025-08-19T16:29:39+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":79128092,"identity":"8f306f30-3a73-487b-950f-95b7ab9978b8","added_by":"auto","created_at":"2025-03-24 17:56:02","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":144285,"visible":true,"origin":"","legend":"\u003cp\u003eStudy sites in the Capivari-Monos EPA, São Paulo, Brazil: Cachoeira, Siloé, and Borracharia. Green areas represent the remnants of the Atlantic Forest. Circles indicate collection points. The map was created using QGIS v3.34 (\u003ca href=\"http://www.qgis.org/\"\u003ehttp://www.qgis.org\u003c/a\u003e).\u003c/p\u003e","description":"","filename":"Picture1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6184497/v1/fe32dd8fe9198a3a6c7c714f.jpg"},{"id":79128090,"identity":"467b4729-e1d6-4a3e-888b-ca4599661f2b","added_by":"auto","created_at":"2025-03-24 17:56:02","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":138322,"visible":true,"origin":"","legend":"\u003cp\u003eGraphs showing four models that suggest a cascade effect: (A) rainfall and temperature influence water volume in bromeliads, (B) water volume influences pH, (C) pH in interaction with salinity influences mosquito abundance, and (D) the probability of \u003cem\u003eAnopheles cruzii\u003c/em\u003e presence is influenced by pH. The shaded areas in A and C represent the 95% confidence interval for the estimate.\u003c/p\u003e","description":"","filename":"Picture2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6184497/v1/e3dc82819af20dd3bfc38e16.jpg"},{"id":79127558,"identity":"76c6c1a5-35a6-4958-b8c0-eb984c6edcc9","added_by":"auto","created_at":"2025-03-24 17:48:02","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":72958,"visible":true,"origin":"","legend":"\u003cp\u003eStructural equation models exploring the effects of abiotic variables on mosquitoes collected in bromeliads: (A) species richness, (B) total abundance, and (C) \u003cem\u003eAnopheles cruzii\u003c/em\u003e presence. Relationships between variables are indicated by arrows. The values on the arrows show the scaled estimated slope parameters.\u003c/p\u003e","description":"","filename":"Picture3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6184497/v1/5222f122011897982b086b26.jpg"},{"id":89847642,"identity":"0abe2a3a-27a2-4422-b888-cdaf5ab34cc2","added_by":"auto","created_at":"2025-08-25 16:43:53","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1112156,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6184497/v1/a1e16a09-ea06-47a3-8356-bf6f73994390.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"The interplay of key abiotic factors with the occurrence and abundance of mosquito larvae in bromeliads","fulltext":[{"header":"INTRODUCTION","content":"\u003cp\u003eAbiotic factors are the non-living components of the ecosystem that influence the survival, growth, reproduction and behavior of organisms [1]. Temperature and rainfall are abiotic components that significantly influence aquatic organisms, particularly those in lentic environments, such as insect larvae [2, 3]. While temperature variation is the main factor responsible for structuring aquatic insect assemblages in temperate environment [4], in tropical regions, where temperature varies less, rainfall can play a more influential role by altering both the availability of sites for colonization and the physicochemical factors of the environment [5, 6].\u003c/p\u003e \u003cp\u003eFor immature forms of mosquitoes (Diptera: Culicidae), the physicochemical conditions of water are important constraints that influence their survival, growth and adaptation [7, 8, 9]. Seasonal patterns or extreme weather events that alter rainfall and temperature can lead to changes in the physicochemical parameters of lentic environments where mosquitoes breed. For example, dry and warmer periods favor the loss of water by evaporation, reducing the water body volume and leading to an increase of insolation and ionic concentration with changes in primary production that affect pH, salinity and dissolved oxygen [5, 10]. While pH and salinity directly impact the ionic and osmotic regulation of mosquitoes, dissolved oxygen primarily affects the associated microfauna and flora in breeding sites [11, 12].\u003c/p\u003e \u003cp\u003eBromeliads (Bromeliaceae) are a family of neotropical plants with more than 3,700 described species [13]. Most bromeliads are epiphytic or terrestrial, with long leaves arranged in a rosette shape that form tanks capable of retaining water and nutrients, thereby supporting the development of biotic communities [14]. In the Brazilian Atlantic Forest, there is a high diversity of bromeliads that harbor many species of aquatic insects, including immature forms of mosquitoes, some of which have epidemiological importance, such as \u003cem\u003eAnopheles\u003c/em\u003e (\u003cem\u003eKerteszia\u003c/em\u003e) \u003cem\u003ecruzii\u003c/em\u003e, the main vector of malaria parasites in this region [15, 16, 17]. Given that changes in temperature and rainfall patterns driven by climate change are key factors that will impact the biodiversity of freshwater communities in the coming decades [18], we hypothesize that variations in temperature and precipitation affect the volume of bromeliad water tanks, thereby promoting changes in the physicochemical parameters of the water, which in turn influence the patterns of occurrence, abundance, and richness of mosquito assemblages developing in these aquatic ecosystems.\u003c/p\u003e \u003cp\u003eTo investigate the effect of these abiotic factors on mosquito assemblages in bromeliads, we conducted a study with nine bromeliads in a protected area of the Atlantic Forest between 2015 and 2017, collecting larval samples and measuring abiotic parameters. The data obtained demonstrate the effect of average maximum temperature and accumulated rainfall on the water volume of the bromeliad tanks and the effect of this variation on the pH, which in turn, either alone or in interaction with salinity, influenced the occurrence, abundance, and richness of mosquitoes in these microhabitats.\u003c/p\u003e"},{"header":"METHODS","content":"\u003cp\u003eThe study was performed in the Capivari-Monos Environmental Protection Area (EPA), a remnant of Atlantic Forest situated in the southern region of the city of S\u0026atilde;o Paulo, Brazil. The EPA extends over the first hills near the crest of the Serra do Mar Mountain range at altitudes varying from 740 to 850 m above sea level. The climate is predominantly super-humid, oceanic, and tropical with average annual temperatures of around 19\u0026deg;C and rainfall of between 1600 and 2200 mm [19].\u003c/p\u003e \u003cp\u003eA total of ten field collections were carried out between November 2015 and April 2017 (Nov-Dec/2015, Jan-Oct-Nov-Dec/2016, Jan-Feb-Mar-Apr/2017). Nine bromeliads from three different sites in the EPA were selected and monitored throughout the study: four bromeliads at Cachoeira (C1, C2, C3, and C4), a private property adjacent to a waterfall with a visitation area (23\u0026deg; 56.378\u0026prime; S / 46\u0026deg;41. 659\u0026prime; W); two bromeliads at Silo\u0026eacute; (S1 and S2), a private property near Marsilac village, constituting a transitional area between a rural environment and the EPA forest (23\u0026deg; 54.556\u0026prime; S / 46\u0026deg; 42.167\u0026prime; W); and three bromeliads at Borracharia (B1, B2, and B3), situated in a small village surrounded by the EPA forest (23\u0026deg; 54.395\u0026prime; S / 46\u0026deg; 42.486\u0026prime; W) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). All the bromeliads were epiphytes (ranging from one to seven meters high) and were identified as \u003cem\u003eVriesia\u003c/em\u003e spp. (Bromeliaceae: Tillandsioideae).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eAt each collection, the water content present in each bromeliad was extracted using suction samplers and examined. The total volume of water, pH, salinity (practical salinity unit - PSU), and dissolved oxygen (parts per million - PPM) were measured for each sample using a multiparameter meter (Hanna\u0026reg; HI-9828). The specimens present were separated using pipettes and stored in plastic pots for later identification. After this, the remaining water content was returned to the bromeliad to minimize interference with its biotic community. The collected immature forms (larvae and pupae) were transported to the Entomology Laboratory, School of Public Health, University of S\u0026atilde;o Paulo (LESP/FSP/USP), where they were monitored until the adult stage, after which the adults were identified morphologically.\u003c/p\u003e \u003cp\u003eDaily rainfall and temperature data for the study sites were obtained from the Centro de Gerenciamento de Emerg\u0026ecirc;ncias de S\u0026atilde;o Paulo (CGE). For temperature, the average maximum and minimum (\u0026ordm;C) for the 30 days prior to collection were considered. For rainfall, the accumulated rainfall (mm\u0026sup3;) for the 30 days prior to collection was considered. The average maximum and minimum temperatures were highly correlated, so only the average maximum temperature was kept as a predictor variable, as it demonstrated a superior fit in the models.\u003c/p\u003e \u003cp\u003eTo analyze the data, a cascade effect hypothesis was tested, in which an initial disturbance (variation in rainfall and/or temperature) affects other components in a sequence of causally connected processes [1, 20, 21, 22]. First, the effect of accumulated rainfall and average monthly temperatures on the volume of water in the bromeliad tanks was analyzed. Next, the relationship between water volume and the variation in physicochemical parameters was analyzed. Finally, the relationship between physicochemical parameters and the occurrence, richness, and abundance of mosquitoes in the bromeliads was investigated. Direct effects of precipitation and temperature on physicochemical parameters and water volume on mosquitoes were also explored.\u003c/p\u003e \u003cp\u003eThe relationships between the variables were verified using generalized linear mixed-effect models (GLMM) with Gaussian, binomial, or Poisson error distributions, depending on the type of response variable. The different sites and bromeliads studied were considered as random effects. To fit the best model, the backward elimination strategy was used, starting with the full model (fixed effect variables\u0026thinsp;+\u0026thinsp;fixed effect interactions\u0026thinsp;+\u0026thinsp;random effect) and eliminating the least significant fixed effect variables until the best fit was obtained. To investigate the relationship between abiotic variables and the occurrence of mosquitoes, only species that occurred five or more times during the study period were considered. Finally, we performed a path analysis using piecewise structural equation modeling (piecewise SEM) [23] to investigate the direct and indirect effects of the abiotic variables on species richness, total abundance, and occurrence of \u003cem\u003eAn. cruzii\u003c/em\u003e in bromeliads. The goodness-of-fit of piecewise SEM models was evaluated using Fisher\u0026rsquo;s C test [24]. The values of C can be compared to a chi-squared distribution with 2k degrees of freedom, and values of \u003cem\u003ep\u003c/em\u003e above the chosen significance threshold (α\u0026thinsp;=\u0026thinsp;0.05) indicate a good fit of the model (the hypothesized relationships are consistent with the data). The R computer environment [25] was used to analyze the data using the packages lme4 [26], ggplot2 [27], sjPlot [28], and piecewiseSEM [23].\u003c/p\u003e"},{"header":"RESULTS","content":"\u003cp\u003eThroughout the study period, the accumulated rainfall for the 30 days prior to collection ranged from 61 to 429 (mm\u0026sup3;) while the temperature for the 30 days prior to collection ranged from 21.6 to 29.5\u0026ordm;C for the average daily maximum and from 13.3 to 18.9\u0026ordm;C for the average daily minimum. The volume of water sampled per bromeliad ranged from 95 to 1,050 ml. For the physicochemical parameters, pH ranged from 3.57 to 6.02, salinity from 0.006 to 0.36 PSU and dissolved oxygen from 0.2 to 8.56 ppm.\u003c/p\u003e \u003cp\u003eA total of 529 individuals belonging to 20 species were collected, including specimens of \u003cem\u003eAn. cruzii\u003c/em\u003e and specimens of the genera \u003cem\u003eCulex\u003c/em\u003e (mostly of the subgenus \u003cem\u003eMicroculex\u003c/em\u003e) and \u003cem\u003eWyeomyia\u003c/em\u003e of the subgenus \u003cem\u003ePhoniomyia\u003c/em\u003e. The number of species per bromeliad ranged from 7 to 14 (bromeliads B3 and S1, respectively). The most abundant species were those of the subgenus \u003cem\u003eMicroculex\u003c/em\u003e, especially \u003cem\u003eCx. pleuristriatus\u003c/em\u003e with 129 individuals collected. In terms of occurrence, only 10 species occurred five or more times throughout the study, with \u003cem\u003eCx. imitator\u003c/em\u003e and \u003cem\u003eWy. davisi\u003c/em\u003e being the most frequent (29 and 19 times, respectively) and the only ones to be present in all the bromeliads investigated (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eNumber of individuals and occurrences per mosquito species found in nine bromeliads investigated at Capivari-Monos EPA between November 2015 and April 2017.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"12\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"9\" nameend=\"c10\" namest=\"c2\"\u003e \u003cp\u003eBromeliad\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eTotal individuals\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c12\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eNumber of occurrences\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eB1\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eB2\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eB3\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eC1\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eC2\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eC3\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eC4\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eS1\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eS2\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAnopheles (Ker.) cruzii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCulex (Cux.) eduardoi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCulex (Mcx.) albipes\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCulex (Mcx.) daumasturus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCulex (Mcx.) dubitans\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCulex (Mcx.) fuscatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCulex (Mcx.) imitator\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e29\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCulex (Mcx.) inimitabilis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCulex (Mcx.) neglectus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCulex (Mcx.) pleuristriatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e73\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e129\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCulex (Mcx.) worontzowi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCulex ocellatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eWyeomyia (Pho.) quasilongirostris\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eWyeomyia (Pho.) davisi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eWyeomyia (Pho.) edwardsi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eWyeomyia (Pho.) incaudata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eWyeomyia (Pho.) pallidoventer\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eWyeomyia (Pho.) pilicauda\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eWyeomyia (Pho.) splendida\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eWyeomyia (Pho.) theobaldi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eTotal individuals\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e109\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e529\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eTotal species\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eRegarding the best-fit models, the averages of maximum temperature and accumulated rainfall showed a significant influence on the volume of water in the bromeliads, with rainfall positively related and temperature negatively related to the water volume (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e and Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA). Rainfall was also positively related to pH, while temperature was negatively related to dissolved oxygen. The volume of water was positively related to the increase in pH (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eB) and dissolved oxygen, while the increase in volume was negatively related to the abundance of mosquitoes in the bromeliads (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). No significant relationship was found between water volume and the occurrence probability of any mosquito species.\u003c/p\u003e \u003cp\u003eBoth the richness and abundance of mosquitoes varied as a function of pH, salinity, and the interaction between these two physicochemical parameters. For lower salinities (predicted to 0.025 and 0.05 PSU), richness and abundance increased as a function of increasing pH, while for higher salinities (predicted to 0.1 PSU), richness and abundance decreased in response to increasing pH (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e and Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eC). In relation to the probability of species occurrence, pH was the parameter most associated with the presence of five of the ten species tested, including the vector \u003cem\u003eAn. cruzii\u003c/em\u003e, which showed a positive association with pH (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e and Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eD).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eModels that showed the best fit in the study. The intercept and slope values refer to the fixed effect parameters of the GLMMs. The standard error of the estimate is shown in brackets.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eExplanatory variables\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eResponse variables\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIntercept\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSlope\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003ep\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eRain\u0026thinsp;+\u0026thinsp;Temp_max\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eWater volume\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e786.7 (246.2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.674 (0.216)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.003\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-22.93 (10.78)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.04\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRain\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e4.35 (0.14)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.002 (0.0005)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.0005\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTemp_max\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDissolved oxygen\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e9.46 (2.25)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-0.241 (0.09)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.007\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater volume\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e4.36 (0.157)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.001 (0.0004)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.002\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater volume\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDissolved oxygen\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1.82 (0.56)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.004 (0.001)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.007\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater volume\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTotal abundance\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2.32 (0.15)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-0.0007(0.0003)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.037\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eAn. cruzii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-11.04 (4.35)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.938 (0.846)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.022\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e\u003cem\u003eCx. albipes\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e-12.34 (4.88)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2.36 (1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.018\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSalinity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e166.42 (83.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.046\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH:Salinity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-36.25 (18.68)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.052\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eCx. dubitans\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-24.23 (10.53)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e4.26 (1.94)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.028\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eCx. neglectus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-11.61 (4.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.963 (0.832)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.018\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eWy. incaudata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e7.06 (3.85)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-1.824 (0.835)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.029\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eSpecies richness\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e-1.38 (1.02)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.521 (0.2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.013\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSalinity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e28.5 (16.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.075\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH:Salinity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-6.65 (3.53)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.059\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eTotal abundance\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003e0.017 (0.63)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.477 (0.13)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.0002\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSalinity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e30.73 (10.54)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.003\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epH:Salinity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-7.6 (2.35)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003ePiecewise SEM analysis indicated that the data were satisfactorily fitted for the three models tested: species richness (Fisher\u0026rsquo;s C\u0026thinsp;=\u0026thinsp;13.5, \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;12, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.33), total abundance (Fisher\u0026rsquo;s C\u0026thinsp;=\u0026thinsp;10, \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;8, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.26), and \u003cem\u003eAn. cruzii\u003c/em\u003e presence (Fisher\u0026rsquo;s C\u0026thinsp;=\u0026thinsp;4.6, \u003cem\u003edf\u003c/em\u003e\u0026thinsp;=\u0026thinsp;8, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.8). Rainfall had direct effects on water volume, pH, and total abundance, while temperature had direct effects on water volume and total abundance. The model for total abundance showed a high explanatory power (R\u0026sup2; = 0.63), followed by \u003cem\u003eAn. cruzii\u003c/em\u003e presence (R\u0026sup2; = 0.32), and species richness (R\u0026sup2; = 0.13) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"DISCUSSION","content":"\u003cp\u003eThe observed results illustrate the interaction between abiotic factors and the occurrence and abundance of mosquito larvae in bromeliads, supporting the hypothesis of a cascade effect where variations in rainfall and temperature lead to changes in the water volume of bromeliad tanks, which in turn influence physicochemical parameters and subsequently affect the richness, abundance, and occurrence of mosquitoes in these microhabitats. While rainfall increases the volume of water retained in the bromeliads, rising temperatures decrease this volume due to greater evaporation. As the volume of water in the bromeliad tanks increases, there is a tendency for pH to increase. In turn, the variation in pH influences the occurrence of some species and, when associated with salinity, induces variations in the richness and abundance of mosquitoes in the bromeliads.\u003c/p\u003e \u003cp\u003eRainfall and temperature have been shown to have diverse effects on the biotic communities of bromeliads. Previous studies have observed that changes in the hydrological regime, particularly extreme events leading to droughts or prolonged rains, affect the food web [29], the relative abundance and biomass of functional groups of invertebrates [30, 31], and the biomass and composition of microorganisms [32, 33, 34]. For mosquitoes, studies conducted in the Atlantic Forest of southeastern Brazil have shown a positive relationship between rainfall, water volume in bromeliad tanks and the presence of some species, including \u003cem\u003eAn. cruzii\u003c/em\u003e and mosquitoes of the genera \u003cem\u003eCulex\u003c/em\u003e and \u003cem\u003eWyeomyia\u003c/em\u003e [17, 35]. In a study conducted in a semi-arid region of northeastern Brazil, it was found that the probability of mosquitoes being present in bromeliads was inversely proportional to the average temperature for the 30 days prior to collection, because under high temperatures there was greater evaporation of water and the bromeliad tanks dried out more quickly [36].\u003c/p\u003e \u003cp\u003eAs observed in this study, pH was a key factor associated with changes in the richness, abundance, and occurrence of some species. The water in the bromeliad tank tends to have low pH values and this has been associated with acids resulting from microbial activity [37], leaf litter accumulated in the tanks [38], accumulation of carbonic acid due to CO2 produced in respiration by microbes [39], and the activity of proton pumps in the leaves as mechanisms of tank water acidification [40]. In a study conducted in Costa Rica, it was observed that the average pH of tank water was lower than that of rainwater for six of the eight bromeliad species investigated [40]. Our data corroborates these studies, indicating that the water present in bromeliad ponds tends to be acidic and that the increase in the volume of water in the ponds due to precipitation induces a reduction in the concentration of H\u003csup\u003e+\u003c/sup\u003e ions.\u003c/p\u003e \u003cp\u003eInterestingly, the interaction between increased pH and salinity was observed to reduce the abundance and richness of mosquitoes in bromeliads, suggesting a suboptimal environment for species that colonize this ecosystem. The mosquito species present in bromeliads seem to have some degree of predilection for more acidic environments with low salinity [41]. As for physiological factors, the homeostasis mechanisms that allow mosquito larvae to maintain the ionic composition and pH of the hemolymph are still poorly understood, although it is known that ions and acid-base equivalents are absorbed or excreted directly between the hemolymph and the surroundings through ion-permeable epithelia [42]. The anal papillae appear to function as the main organ of ion exchange, absorbing Na+, Cl\u0026minus;, and K\u0026thinsp;+\u0026thinsp;and excreting the acid-base equivalents H\u003csup\u003e+\u003c/sup\u003e, NH4\u003csup\u003e+\u003c/sup\u003e, and HCO3\u003csup\u003e\u0026minus;\u003c/sup\u003e [43].\u003c/p\u003e \u003cp\u003eMore than 200 mosquito species have been described inhabiting bromeliads in the Neotropical region, many of which are bromeliad specialists [14, 35]. An example of a specialist is the \u003cem\u003eAnopheles\u003c/em\u003e of the subgenus \u003cem\u003eKerteszia\u003c/em\u003e, particularly \u003cem\u003eAn. cruzii\u003c/em\u003e, which is primarily responsible for maintaining the circulation of malaria plasmodia between humans and non-human primates in the Atlantic Forest [44, 45]. For this reason, the autochthonous malaria that occurs in this region is also known as \u0026ldquo;Bromeliad-malaria\u0026rdquo;. In the mid-1940s, the southeastern and southern regions of Brazil were considered endemic areas for bromeliad malaria, with an annual incidence of close to 4,000 cases per 100,000 inhabitants [46]. Once it was confirmed that \u003cem\u003eKerteszia\u003c/em\u003e mosquitoes were the transmitters of malaria in the Atlantic Forest, various control efforts were simultaneously adopted. The measures consisted of applying larvicides and insecticides, treating patients with antimalarials, destroying bromeliads by removing them or using herbicides, and even deforesting areas close to transmission hotspots [47]. Currently, bromeliad malaria is hypoendemic in the region and few cases have been recorded annually. Data from the Centro de Vigil\u0026acirc;ncia Epidemiol\u0026oacute;gica of the State of S\u0026atilde;o Paulo show that between January 2017 and November 2024, 77 autochthonous cases of malaria were confirmed in the State [48].\u003c/p\u003e \u003cp\u003eClimate change is expected to have profound effects on freshwater communities, particularly by altering hydrological cycles, temperature regimes, and nutrient dynamics [49]. These environmental shifts can significantly impact mosquito populations, as changes in rainfall and temperature influence breeding site availability, larval development rates, and adult survival [50]. For instance, increased temperatures can accelerate the life cycle of vectors such as \u003cem\u003eAedes aegypti\u003c/em\u003e and \u003cem\u003eCulex quinquefasciatus\u003c/em\u003e, leading to higher population densities and prolonged transmission seasons for arboviruses, including dengue, Zika, West Nile virus, and Chikungunya [51]. Similarly, malaria transmission, primarily driven by \u003cem\u003eAnopheles\u003c/em\u003e mosquitoes, is also projected to shift due to climate change. Studies have shown that regions in East Africa and South America may become increasingly suitable for malaria transmission, while some highly endemic areas could experience declines due to excessive warming [52]. Additionally, extreme weather events, such as floods and droughts, may expand mosquito habitats into previously unsuitable areas, facilitating the introduction of vector-borne diseases into new ecological and epidemiological contexts [53]. These complex interactions highlight the need for investigating the interplay between abiotic factors and mosquitoes\u0026rsquo; biodiversity. Understanding these dynamics is crucial for predicting the impacts of climate change on mosquito-borne diseases and for developing effective strategies to mitigate the associated public health risks.\u003c/p\u003e \u003cp\u003eThis study has provided evidence of how changes in precipitation and temperature promote a cascade effect that leads to changes in the richness, abundance, and composition of mosquitoes, using the microcosm of bromeliads as a model. Given this evidence, it is necessary for future studies to seek to understand how ongoing climate change, associated with land use change and loss of biodiversity, will affect the occurrence, distribution, and abundance of mosquitoes, and the impacts of these changes on human health and ecosystems in tropical regions of the planet.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eCompeting interests\u003c/h2\u003e \u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eA.R.M-S: Conceptualization, Investigation, Methodology, Visualization, Formal analysis, and Writing - Original Draft. R. O-C: Conceptualization, Investigation and Data-curation. W. C-J: Investigation and Data-curation. K.M.B-N: Investigation. E.E: Investigation. R.W-S: Investigation. L.F.M: Conceptualization and Investigation. M.B.P: Data-curation. M.T.M: Conceptualization, Project administration, Funding acquisition and Supervision. All authors reviewed the manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eWe thank the State of S\u0026atilde;o Paulo Research Foundation (FAPESP Grants Nos. 2014/50444-5, 2023/11212-0 and 2023/17044-2) and the National Council for Scientific and Technological Development, Brazil (CNPq 309872/2021-9) for providing financial support. We would also like to express our gratitude to the field and laboratory teams at the Department for the Control of Endemic Diseases, S\u0026atilde;o Paulo Zoonosis Control Center, and the School of Public Health, University of S\u0026atilde;o Paulo: Ana Maria Ribeiro de Castro Duarte, Jo\u0026atilde;o Carlos do Nascimento, Paulo Frugoli dos Santos, Luis Milton Bonaf\u0026eacute;, Ant\u0026ocirc;nio Waldomiro de Oliveira, La\u0026eacute;rcio Molinari, Gabriel Marcelino Neto, Luiz Sposito Jr, Renildo Souza Teixeira, Daniel Pagotto Vendrami, Gabriela Cristina de Carvalho, and Amanda Alves Camargo.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eThe datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eKumar, G., Pasi, S., Kaur, J., \u0026amp; Singh, H. (2024). Abiotic and Biotic Interactions of Mosquitoes. In Mosquitoes: Biology, Pathogenicity and Management (pp. 223\u0026ndash;235). Springer Nature, Singapore.\u003c/li\u003e\n\u003cli\u003eValdez, L. D., Sibona, G. J., \u0026amp; Condat, C. A. (2018). Impact of rainfall on Aedes aegypti populations. Ecological Modelling, 385, 96\u0026ndash;105.\u003c/li\u003e\n\u003cli\u003eNash, L. N., Zorzetti, L. W., Antiqueira, P. A., Carbone, C., Romero, G. Q., \u0026amp; Kratina, P. (2023). Latitudinal patterns of aquatic insect emergence driven by climate. Global Ecology and Biogeography, 32(8), 1323\u0026ndash;1335.\u003c/li\u003e\n\u003cli\u003eWard, J. V., \u0026amp; Stanford, J. A. (1982). Thermal responses in the evolutionary ecology of aquatic insects. Annual review of entomology, 27(1), 97\u0026ndash;117.\u003c/li\u003e\n\u003cli\u003eWilliams, D. D. (1996). Environmental constraints in temporary fresh waters and their consequences for the insect fauna. 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(2016). piecewiseSEM: Piecewise structural equation modelling in r for ecology, evolution, and systematics. Methods in Ecology and Evolution, 7(5), 573\u0026ndash;579.\u003c/li\u003e\n\u003cli\u003eShipley, B. (2000). A new inferential test for path models based on directed acyclic graphs. Structural Equation Modeling, 7(2), 206\u0026ndash;218.\u003c/li\u003e\n\u003cli\u003eR Development Core Team (2024). R - a language and environment for statistical computing. Vienna: R Foundation for Statistical Computing. https://www.r-project.org\u003c/li\u003e\n\u003cli\u003eBates, D., Maechler, M., Bolker, B., Walker, S., Christensen, R. H. B., Singmann, H., ... \u0026amp; Bolker, M. B. (2015). Package \u0026lsquo;lme4\u0026rsquo;. Convergence, 12(1), 2.\u003c/li\u003e\n\u003cli\u003eWickham, H., Chang, W., Henry, L., Pedersen, T. L., Takahashi, K., Wilke, C., ... \u0026amp; Dunnington, D. (2020). Package \u0026lsquo;ggplot2\u0026rsquo;: create elegant data visualisations using the grammar of graphics. R package version, 3(0).\u003c/li\u003e\n\u003cli\u003eL\u0026uuml;decke, M. D. (2024). Package \u0026lsquo;sjPlot\u0026rsquo;. Data Visualization for Statistics in Social Science. Available at: https://strengejacke.github.io/sjPlot/\u003c/li\u003e\n\u003cli\u003eRomero, G. Q., Marino, N. A., MacDonald, A. A. M., C\u0026eacute;r\u0026eacute;ghino, R., Trzcinski, M. K., Mercado, D. A., ... \u0026amp; Srivastava, D. S. (2020). Extreme rainfall events alter the trophic structure in bromeliad tanks across the Neotropics. Nature communications, 11(1), 3215.\u003c/li\u003e\n\u003cli\u003eDezerald, O., Leroy, C., Corbara, B., Dejean, A., Talaga, S., \u0026amp; C\u0026eacute;r\u0026eacute;ghino, R. (2017). Environmental drivers of invertebrate population dynamics in Neotropical tank bromeliads. Freshwater Biology, 62(2), 229\u0026ndash;242.\u003c/li\u003e\n\u003cli\u003eSrivastava, D. S., C\u0026eacute;r\u0026eacute;ghino, R., Trzcinski, M. K., MacDonald, A. A. M., Marino, N. A., Mercado, D. A., ... \u0026amp; Campos, A. B. (2020). Ecological response to altered rainfall differs across Neotropics. Ecology, 101(4), e02984.\u003c/li\u003e\n\u003cli\u003eBuosi, P. R. B., Utz, L. R. P., de Meira, B. R., da Silva, B. T. S., Lansac-T\u0026ocirc;ha, F. M., Lansac-T\u0026ocirc;ha, F. A., \u0026amp; Velho, L. F. M. (2014). Rainfall influence on species composition of the ciliate community inhabiting bromeliad phytotelmata. Zoological Studies, 53, 1\u0026ndash;12.\u003c/li\u003e\n\u003cli\u003ePires, A. P. F., Leal, J. D. S., \u0026amp; Peeters, E. T. (2017). Rainfall changes affect the algae dominance in tank bromeliad ecosystems. PLoS One, 12(4), e0175436.\u003c/li\u003e\n\u003cli\u003ePenna, D. D. P. S., Romero, G. Q., Nessel, M. P., Gonz\u0026aacute;lez, A. L., \u0026amp; Oliveira, V. M. (2025). Bacterial communities as bioindicators of climate change in freshwater ecosystems: Tank bromeliads as model systems. Ecological Indicators, 171, 113161.\u003c/li\u003e\n\u003cli\u003eCardoso, C. A. A., Louren\u0026ccedil;o-de-Oliveira, R., Code\u0026ccedil;o, C. T., \u0026amp; Motta, M. A. (2015). Mosquitoes in bromeliads at ground level of the Brazilian Atlantic Forest: the relationship between mosquito fauna, water volume, and plant type. Annals of the Entomological Society of America, 108(4), 449\u0026ndash;458.\u003c/li\u003e\n\u003cli\u003eMarteis, L. S., Natal, D., Sallum, M. A. M., Medeiros-Sousa, A. R., \u0026amp; La Corte, R. (2017). Mosquitoes of the Caatinga: 2. Species from periodic sampling of bromeliads and tree holes in a dry Brazilian forest. Acta Tropica, 171, 114\u0026ndash;123.\u003c/li\u003e\n\u003cli\u003eJocque, M., \u0026amp; Kolby, J. E. (2012). Acidity of tank bromeliad water in a cloud forest, Cusuco National Park, Honduras. 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R., Ferreira, L. M., ... \u0026amp; Cerutti Junior, C. (2021). Atlantic Forest Malaria: A review of more than 20 years of epidemiological investigation. Microorganisms, 9(1), 132.\u003c/li\u003e\n\u003cli\u003eDeane, L. M. Malaria studies and control in Brazil. The American journal of tropical medicine and hygiene, v. 38, n. 2, p. 223\u0026ndash;230, 1988.\u003c/li\u003e\n\u003cli\u003eBarata, J. M. S. (1973). Estado atual da brom\u0026eacute;lia-mal\u0026aacute;ria no Brasil. (Master Dissertation) \u0026ndash; School of Public Health, University of S\u0026atilde;o Paulo, S\u0026atilde;o Paulo, 1973.\u003c/li\u003e\n\u003cli\u003eCVE \u0026ndash; Centro de Vigil\u0026acirc;ncia Epidemiol\u0026oacute;gica. (2024). Casos Aut\u0026oacute;ctones de Mal\u0026aacute;ria no Estado de S\u0026atilde;o Paulo. 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The Lancet Planetary Health, 4(9), e416-e423.\u003c/li\u003e\n\u003cli\u003eCaminade, C., Kovats, S., Rocklov, J., Tompkins, A. M., Morse, A. P., Col\u0026oacute;n-Gonz\u0026aacute;lez, F. J., ... \u0026amp; Lloyd, S. J. (2014). Impact of climate change on global malaria distribution. Proceedings of the National Academy of Sciences, 111(9), 3286\u0026ndash;3291.\u003c/li\u003e\n\u003cli\u003eCaminade, C., McIntyre, K. M., \u0026amp; Jones, A. E. (2019). Impact of recent and future climate change on vector-borne diseases. Annals of the New York Academy of Sciences, 1436(1), 157\u0026ndash;173.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Mosquitoes, Bromeliads, Climate change, Cascading effect, Anopheles cruzii","lastPublishedDoi":"10.21203/rs.3.rs-6184497/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6184497/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eBromeliads provide a unique habitat for many insect species, including immature forms of mosquitoes. This study investigates the effect of abiotic factors, specifically temperature, rainfall, water volume, and physicochemical parameters, on the occurrence and abundance of mosquitoes developing in bromeliads within the Brazilian Atlantic Forest. Larval samples and abiotic parameters were collected from nine bromeliads between 2015 and 2017 in a forest remnant in S\u0026atilde;o Paulo, Brazil. Relationships between variables were tested using generalized linear mixed-effects models and structural equation modeling. Results indicate that rainfall and temperature influence the water volume of bromeliad tanks, which in turn affect physicochemical parameters, particularly pH. These variations affect the richness, abundance, and occurrence of mosquito species, including the vector of malaria parasites, \u003cem\u003eAnopheles cruzii\u003c/em\u003e. The study highlights a cascade effect where abiotic factors alter the microhabitat conditions, thereby affecting mosquito populations. Understanding these interactions is crucial for predicting the impacts of climate change on mosquito-borne diseases in tropical regions.\u003c/p\u003e","manuscriptTitle":"The interplay of key abiotic factors with the occurrence and abundance of mosquito larvae in bromeliads","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-03-24 17:47:57","doi":"10.21203/rs.3.rs-6184497/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-04-23T03:50:03+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-04-06T18:39:23+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-04-04T19:12:32+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"268167901637027911781905151175769581932","date":"2025-03-28T16:56:45+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"58349671099464435539199738381609508355","date":"2025-03-25T12:08:20+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-03-20T11:56:41+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-03-14T12:32:36+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2025-03-14T10:36:11+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-03-14T06:24:20+00:00","index":"","fulltext":""},{"type":"submitted","content":"Scientific Reports","date":"2025-03-08T13:52:07+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"21cf00b4-6889-457a-8860-0e9fe4d139e2","owner":[],"postedDate":"March 24th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[{"id":45967176,"name":"Biological sciences/Ecology/Climate change ecology"},{"id":45967177,"name":"Biological sciences/Ecology/Ecological epidemiology"},{"id":45967178,"name":"Biological sciences/Ecology/Ecological genetics"},{"id":45967179,"name":"Biological sciences/Ecology/Freshwater ecology"},{"id":45967180,"name":"Health sciences/Diseases/Infectious diseases/Malaria"}],"tags":[],"updatedAt":"2025-08-25T16:40:42+00:00","versionOfRecord":{"articleIdentity":"rs-6184497","link":"https://doi.org/10.1038/s41598-025-15514-7","journal":{"identity":"scientific-reports","isVorOnly":false,"title":"Scientific Reports"},"publishedOn":"2025-08-19 16:29:39","publishedOnDateReadable":"August 19th, 2025"},"versionCreatedAt":"2025-03-24 17:47:57","video":"","vorDoi":"10.1038/s41598-025-15514-7","vorDoiUrl":"https://doi.org/10.1038/s41598-025-15514-7","workflowStages":[]},"version":"v1","identity":"rs-6184497","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6184497","identity":"rs-6184497","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}