Responses of Dragonflies (Odonata) to Habitat Integrity and Environmental Heterogeneity in Amazonian Streams: lessons for conservation

Responses of Dragonflies (Odonata) to Habitat Integrity and Environmental Heterogeneity in Amazonian Streams: lessons for conservation | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Responses of Dragonflies (Odonata) to Habitat Integrity and Environmental Heterogeneity in Amazonian Streams: lessons for conservation Jorge Luiz Silva, Juan Mateo Rivera-Pérez, Francisco Maciel Barbosa-Santos, and 6 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8066154/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 9 You are reading this latest preprint version Abstract Rapid landscape transformation in the Amazon, driven by human activities such as deforestation and intensive land use, has deeply affected the conservation of their aquatic ecosystems, especially small forest streams (i.e. “igarapés”). These environments rely heavily on riparian vegetation to maintain the physical and chemical conditions that support biodiversity. Among the many threatened aquatic organisms, insects of the order Odonata stand out as sensitive bioindicators of habitat alteration. We here evaluated the effects of environmental variables on the distribution of adult Odonata species in streams of the Eastern Amazon. We expected more intact streams to harbor higher Zygoptera richness, whereas less intact streams would show higher Anisoptera richness. We sampled 30 streams along a gradient of environmental alteration. We found that better-preserved streams were also better at preserving Zygoptera diversity while more degraded environments were dominated by tolerant Anisoptera as predicted. However, Anisoptera richness did not significantly increase in low-HII streams, although their relative abundance was higher there, suggesting a more pronounced species turnover than diversity loss. In addition, limnological variables, such as electrical conductivity and coarse particulate detritus, played central roles in structuring communities. These results demonstrate the efficiency of adult Odonata as bioindicators of environmental change and reinforce their use as an effective tool for their conservation and assessing environmental quality in Amazonian streams. Odonata stream ecology land uses impact bioindicators conservation Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction The Amazon region is known as one of the most biodiverse on the planet, harboring an extensive network of interconnected aquatic ecosystems (Garda et al., 2010 ; Peres et al. 2010 ; Latrubesse et al. 2017 ; Metzger et al. 2019 ). These ecosystems play a fundamental role in global stability, contributing to carbon sequestration, rainfall formation, and the storage of the world’s largest reserve of fresh water (Brasil et al. 2017 ). Despite this importance, the Amazon has undergone rapid landscape transformation driven by the expansion of cattle ranching, agriculture, mining, and timber extraction (Fearnside et al. 2005; Faria et al. 2024 ). The reduction or simplification of environmental conditions and ecosystem integrity directly affects resource availability, creating habitat patches with low capacity to meet the ecophysiological requirements of many species, including several groups of aquatic insects (Oliveira-Junior et al. 2022; Rocha et al., 2023 ). The above environmental changes, especially those affecting riparian vegetation, directly impact aquatic insect communities (Luiza-Andrade et al. 2017 ) and promote the replacement of sensitive species by more tolerant ones (Verberk et al. 2010 ; Heino & Grönroos 2014 ). Such effects tend to be stronger in species with lower dispersal capacity (Lutinski et al. 2021 ), since they depend heavily on local environmental conditions (Juen et al. 2016 ; Brasil et al. 2020 ). Similar patterns of shifts in the structure of aquatic insect communities have been observed in Neotropical environments and are mediated by changes in stream limnological characteristics and physical habitat structure (Dias-Silva et al. 2010 ; Carvalho et al. 2013 ; Monteiro-Júnior et al. 2015 ). Among aquatic insects, the order Odonata has been widely used in ecological and biomonitoring studies because of its specific ecophysiological requirements (e.g., Monteiro-Júnior et al. 2013 ; Carvalho et al. 2013 ; Brasil et al. 2020 ). Members of this order maintain a strong relationship with riparian vegetation structure, with some species restricted to more open-canopy environments, whereas others depend on denser vegetation (Oliveira-Junior et al. 2017 ; Carvalho et al. 2018 ; Oliveira-Junior & Juen 2019). In Amazonian igarapés, Odonata respond rapidly to environmental change owing to the relatively short life cycle of their nymphs, which occupy different aquatic microhabitats (e.g., sandy bottoms, rocky substrates, and submerged vegetation) and exhibit distinct feeding habits (Monteiro-Júnior et al. 2013 ; Carvalho et al. 2013 ; Brasil et al. 2020 ). Thus, odonates play an important role in food webs and in the population control of other aquatic invertebrates (Corbet 1999 ; Carvalho et al. 2013 ). This diversity of niches and ecological functions makes the group highly sensitive to variations in the physicochemical and structural conditions of streams, allowing the detection of subtle changes in the ecosystem (Brasil et al. 2020 ). The clear ecological differentiation between the suborders Anisoptera and Zygoptera provides a natural gradient of response to environmental disturbance (Juen and De Marco 2012 ). Zygoptera exhibit ecophysiological attributes that favor “percher” behavior, spending much of their time on perches to defend territories and to carry out mating and oviposition (Heckman 2008; Heiser and Schmitt, 2010 ; Carvalho et al. 2013 ). In contrast, most Anisoptera are strong fliers with greater physiological robustness. They require more open habitats with higher solar incidence to regulate body temperature and initiate activity (Corbet 1999 ). Owing to differences in ecophysiological requirements between the suborders, conserved igarapés tend to preserve greater Zygoptera diversity, whereas degraded igarapés harbor greater Anisoptera diversity (Monteiro-Júnior et al. 2014 , 2015 ; Oliveira-Junior et al. 2017 ; Oliveira-Junior and Juen 2019). An essential metric for understanding these changes in Odonata communities is beta diversity, which represents the variation in species composition along an environmental gradient (Juen et al. 2007 ; Alves-Martins et al. 2019 ; Juen and De Marco 2011 ). This metric makes it possible to identify whether species replacement (turnover) or species loss (nestedness) is associated with habitat degradation (Legendre and Condit,2019). In Amazonian igarapés, where small variations in limnological and structural conditions can generate large differences in the composition of bioindicator communities (Rivera-Pérez et al. 2024), beta diversity is a robust tool for understanding how habitat integrity and heterogeneity influence the differential distribution of Anisoptera and Zygoptera along disturbance gradients (Brito et al. 2021 ; Bastos et al. 2021 ). Accordingly, we here aimed to: i) describe environmental and structural differences among streams, using limnological and habitat variables to identify which best explain species composition; ii) assess the relationship between variation in beta diversity and environmental variables, testing the hypothesis that streams with greater integrity and physicochemical stability preserve higher Zygoptera richness due to this suborder’s stronger dependence on habitat quality and heterogeneity; and iii) evaluate the relationships between habitat integrity and the distribution of Odonata species, testing the hypothesis that species richness of the suborder Anisoptera is higher in environments with lower integrity and greater structural homogeneity. Material and methods Study area The study was conducted in 30 igarapés located in the municipalities of Barcarena and Abaetetuba, in the northeastern portion of the state of Pará, Brazil, during October 2022 (Fig. 1 ). The regional hydrographic network comprises the Murucupi, Arienga, Arapiranga, Barcarena, and Itaporanga rivers. The climate is humid equatorial, with mean annual precipitation ranging from 1,200 to 1,800 mm and mean annual temperatures between 24°C and 26°C. The predominant vegetation corresponds to sub-evergreen equatorial forest (IBGE 2010). Originally, the region was covered by native tropical forest characterized by large trees, as well as secondary forest formations, riparian forest, and floodplain areas influenced by seasonal inundation (Paz et al. 2011 ). From the 1980s onward, extractives were the main local economic activity, that were later replaced by agriculture and livestock, and more recently by urbanization, port activities, and the establishment of industries. These land uses have caused intense reduction and fragmentation of both terrestrial and aquatic habitats. Biological sampling Sampling was carried out along a fixed 150 m reach in each stream, subdivided into 10 sections of 15 m, delimited by 11 transects labeled A (downstream) to K (upstream). This subdivision of the water course into standardized sections facilitates sampling and helps to control the spatial distribution of specimens in the field (Shimano et al., 2013 ; Brito et al., 2018 ). Adult insects were captured by active search along the described transect using an entomological net with a 50 cm diameter. Sampling lasted 1 h 30 min along the 150 m reach and was conducted between 10:00 and 14:00 (Juen et al., 2025 ). Captured specimens were placed individually in glassine envelopes and properly labeled with site, section, and collection date. The specimens were then submerged in absolute anhydrous ethanol (PA/ACS grade) for a minimum of 18 h to preserve key diagnostic structures such as genitalia, wing venation, and coloration essential for taxonomic identification. After this step, the material was taken to the laboratory, where specimens were identified to species using specialized taxonomic keys (Garrison & Von Ellenrieder 2015 ; Garrison et al. 2006 ; Lencioni 2005, 2006 ), complemented by consultation with specialists and comparison with the reference collection of the Laboratory of Ecology and Conservation at the Federal University of Pará (LABECO/UFPA). The identified specimens were subsequently cataloged, accessioned, and incorporated into LABECO’s scientific collection. Environmental variables In each stream, limnological variables were measured using a multiparameter probe (AKSO-87), including water temperature (°C), pH, electrical conductivity (mS /m⁻¹), and dissolved oxygen (mg/L⁻¹). Measurements were taken at three equidistant transects along the channel (downstream, mid-reach, and upstream), and the mean value for each variable was then calculated for each stream. Habitat integrity was assessed using the Habitat Integrity Index (HII) (Nessimian et al. 2008), which considers physical characteristics and the degree of impact from human activities across 12 parameters related to the structure and physical condition of the stream, its banks, and substrate: land use beyond the riparian zone, riparian forest width, riparian forest preservation state, vegetation condition within a 10 m buffer, retention devices, sediment deposition in the channel, bank structure, undercutting of banks, condition of the streambed, presence of riffles, pools, and meanders, aquatic vegetation, and presence of detritus. The index ranges from 0 (least intact) to 1 (most intact), representing a gradient of environmental degradation. In a recent review, HII was identified as one of the most robust metrics for explaining variation in the composition of aquatic insect communities in the Amazon (Brasil et al. 2020 ). Data analysis Initially, each sampled stream was treated as a sampling unit, totaling 30 observations. To reduce the dimensionality of the environmental variables, we applied a Principal Component Analysis (PCA) (Legendre and Legendre, 2012 ). This analysis was used to visualize relationships among pH, temperature, conductivity, and dissolved oxygen. Variables were standardized using z-scores to remove scale effects (Legendre & Legendre 2012 ). The broken-stick criterion was adopted for principal component retention (Jackson 1993 ), and variables with loadings greater than |0.6| were considered relevant. The structure of the Odonata community was visualized using Principal Coordinates Analysis (PCoA) based on Bray–Curtis distance applied to Hellinger-transformed abundance data, to minimize the effects of rare and abundant species (Legendre & De Cáceres, 2013 ). The Hellinger transformation also makes the composition matrix suitable for PCoA, preventing negative eigenvalues (Legendre & Legendre 2012 ). The HII index was overlaid onto the ordination to visually assess the influence of habitat integrity on assemblage composition. To quantify how much of the variation in the Odonata community is explained by environmental variables, we applied Redundancy Analysis (RDA) (Legendre & Legendre 2012 ). The response matrix consisted of Hellinger-transformed abundances, and predictor variables included physicochemical and structural attributes, such as conductivity and amount of detritus. All variables were standardized by z-scores. Variable selection was performed by forward selection with 9,999 permutations. Multicollinearity was assessed using the Variance Inflation Factor (VIF), adopting a threshold of VIF ≥ 3 as indicative of collinearity (Zuur et al. 2010 ); all included variables had VIF < 3. The significance of the final model was tested with 9,999 permutations. Additionally, we applied Pearson’s correlation test to evaluate the relationship between the HII and two community metrics: species richness and the total abundance of Odonata per stream. All statistical analyses were conducted in R (R Core Team, 2020) using the vegan package (Oksanen et al., 2018 ), with a 5% significance level. Results Community Description A total of 519 individuals were collected, representing 57 species distributed across eight families (Table 1 ). The suborder Zygoptera was the most abundant and diverse, with 409 individuals (78.8% of the total) distributed among eight family and 32 species. Within this suborder, the family Protoneuridae was the most numerous, with Epipleoneura metallica (Rácenis, 1955) as the most abundant species (58 individuals), followed by Mnesarete aenea (Selys, 1853) with 42 individuals and Mnesarete williamsoni (Garrison, 2006) with 36 individuals. The suborder Anisoptera contributed 110 individuals (21.2% of the total) distributed among 25 species. The family Libellulidae was the most representative, with Erythrodiplax basalis (Kirby, 1897) as the most abundant species (13 individuals), followed by Erythrodiplax fusca (Rambur, 1842) with 11 individuals. In addition, five species stood out as potential indicators of the ecological conditions of the sampled streams: Aeolagrion dorsale (Burmeister, 1839), Perithemis tenera (Say, 1840), Acanthagrion kennedii (Williamson, 1916), Chalcopteryx rutilans (Rambur, 1842), and Erythrodiplax castanea (Burmeister, 1839). These species exhibit distinct preferences regarding habitat structure and environmental quality. Whereas C. rutilans and A. dorsale are associated with more preserved, shaded reaches, E. castanea and P. tenera occur more frequently in open, human-altered environments. A. kennedii , in turn, tends to occupy transitional areas, reflecting intermediate conditions. Taken together, these species provide a valuable ecological interpretation of the environmental status of the studied igarapés (Table 1 ). Table 1 List of Odonata species collected, and their abundance. TAXON ZYGOPTERA / Family Genus Species / author Abundance Calopterygidae Hetaerina H. laesa Hagen in Selys, 1853 7 H. sanguínea Selys, 1853 6 Hetaerina sp. Hagen in Selys, 1853 1 H. westfalli Rácenis, 1968 1 Mnesarete M. aenea Selys, 1853 42 Mnesarete sp. Cowley, 1934 11 M. williamsoni Garrison, 2006 36 Coenagrionidae Acanthagrion A. adustum Williamson, 1916 2 Acanthagrion sp. Selys, 1876 5 A. ascendens Calvert, 1909 12 A. kennedii Williamson, 1916 3 Acanthallagma A. luteum Willianson & Willianson, 1924 1 Aeolagrion A. dorsale Burmeister, 1839 1 Argia A. collata Selys, 1865 30 A. indicatrix Calvert, 1909 15 A. infumata Selys, 1865 7 A. oculata Hagen in Selys, 1865 10 Argia sp. Rambur, 1842 8 A. tinctipennis Selys, 1865 20 Dicteriadidae Dicterias D. atrosanguinea Selys, 1862 8 Heteragrionidae Heteragrion H. silvarum Sjöstedt, 1918 30 Heteragrion sp. Selys, 1862 3 Oxystigma O. petiolatum Selys, 1862 14 Oxystigma sp. Selys, 1862 1 Polythoridae Chalcopteryx C. rutilans Rambur, 1842 8 Protoneuridae Epipleoneura Epipleoneura sp. 11 E. metallica Rácenis, 1955 58 E. susanae Pessacq, 2014 27 Neoneora N. luzmarina De Marmels, 1989 10 Phasmoneura P. exigua Selys, 1886 1 Protoneura P. tenuis Selys, 1860 16 Psaironeura P. tenuissima Selys. 1886 4 ANISOPTERA Gomphidae Zonophora Z. calippus klugi Schmidt, 1941 1 Libellulidae Argyrothemis A. argentea Ris, 1909 8 Brachymesia B. herbida Gundlach, 1889 2 Diastatops D. obscura Fabricius, 1775 5 Erythemis E. peruviana Rambur, 1842 1 Erythemis sp. Hagen, 1861 1 Erythrodiplax E. basalis Kirby, 1897 13 E. castânea Burmeister, 1839 9 E. fusca Rambur, 1842 11 Erythrodiplax sp. Brauer, 1868 9 Fylgia F. amazonica amazônica Kirby, 1889 10 Gynothemis G. pumila Karsch, 1890 1 Gynothemis sp. Calvert, 1909 1 Macrothemis M. hahneli Ris, 1913 1 Micrathyria M. artemis Ris, 1911 1 Micrathyria sp. Kirby, 1889 1 Oligoclada Oligoclada sp. Karsch, 1890 1 Orthemis Orthemis biolleyi Calvert, 1906 1 O. cultriformis Calvert, 1899 1 Perithemis P. lais Perty, 1834 4 P. tenera Say, 1840 3 P. thais Kirby, 1889 4 Uracis U. imbuta Burmeister, 1839 8 Zenithoptera Z. lanei Santos, 1941 10 Zenithoptera_ sp. Selys, 1869 3 Richness and abundance In the sampled streams, species richness ranged from 4 to 17 (mean ± SD = 8.07 ± 2.95). Despite this variation, no significant correlation was observed between the HII and total richness (r = − 0.146; p = 0.440), indicating that habitat integrity did not consistently influence the number of recorded species. Abundance showed a wider range, from 4 to 41 individuals (10.82 mean ± 17.30 SD). As with richness, the correlation between HII and abundance was not significant (r = 0.216; p = 0.252), suggesting that the habitat integrity gradient did not determine differences in the number of individuals among streams. Environmental variables of streams The PCA explained 72.52% of the total variation of the set of environmental variables in the first two axes (Component 1 = 46.44%; Component 2 = 25.98%). Dissolved oxygen (DO) showed the highest positive correlation with the first axis, while conductivity exhibited the highest negative correlation (Table 2 ; Fig. 2 ). Visually, streams with higher HII values tended to cluster on the positive side of axis 1, associated with higher DO, whereas streams with lower HII values clustered on the negative side, associated with higher conductivity (Table 2 ). This pattern indicates that the first axis represents a habitat-integrity gradient, in which more intact environments are characterized by higher oxygenation and lower conductivity (Fig. 2 ). On the second axis, pH and temperature showed the strongest positive correlations, contributing similarly to the formation of this component. Table 2 Loadings of environmental variables on the first two axes of the PCA conducted with the set of streams limnological variables. Variables with loadings >|0.6| are in bold. Variables PCA 1 PCA 2 pH 0.321 0.794 Conductivity -0.903 -0.129 Dissolved oxygen 0.872 0.164 Temperature -0.372 0.706 Eigenvalue 2.322 1.299 Broken-stick 2.283 1.283 % Explanation 46.44 25.98 Species composition The ordination obtained by Principal Coordinates Analysis (PCoA), based on Bray–Curtis distance, explained 25.82% of the total variation in species composition across the first two axes (PCoA1 = 13.85%; PCoA2 = 11.97%). Axis 1 indicated a partial separation of Odonata assemblages along the habitat-integrity gradient. Reaches with higher HII values tended to cluster on the left side of the ordination, associated with species characteristic of more shaded, structurally heterogeneous environments, such as Epipleoneura susanae , Mnesarete williamsoni , M. aenea , and Heteragrion silvarum . Conversely, reaches with lower HII values were positioned to the right, showing greater representation of heliophilous species tolerant of open, modified environments, such as Argia indicatrix and Zenithoptera lanei . The second axis reflected an additional organization of the Odonata community along the environmental gradient, albeit with lower explanatory power. Overall, the ordination indicated that variation in species composition is driven primarily by habitat integrity and the structural heterogeneity of the streams (Fig. 3 ). Relationship between environmental variables and species composition RDA indicated that higher conductivity and lower amounts of detritus were the environmental variables that best explained variation in the structure of the Odonata community, yielding a statistically significant model (F₂, 27 = 1.82; R²adj = 0.054; p < 0.001). The first two RDA axes together explained 11.88% of the total variation in species composition (Fig. 4 ). Streams with lower HII had less detritus and higher conductivity, reflecting more impacted conditions. These reaches were associated with higher abundance of Epipleoneura susanae , a species that showed moderate tolerance to altered conditions. Conversely, sites with higher conductivity showed greater occurrence of Epipleoneura metallica , suggesting that this species may benefit from more altered physicochemical environments (Fig. 4 ). Overall, the RDA results show that water quality and detritus availability are key determinants of Odonata assemblage structure, reflecting the combined effects of environmental degradation and habitat heterogeneity on species composition. Discussion Our results partially supported the proposed hypotheses that conserved igarapés would show greater Zygoptera diversity, whereas degraded igarapés would harbor greater Anisoptera diversity. Thus, conserved igarapés serve to protect Zygoptera. Accordingly, the species composition was strongly associated with the habitat-integrity gradient, with better-preserved streams showing higher relative abundance and richness of Zygoptera (e.g., Epipleoneura susanae , Mnesarete williamsoni , M. aenea , and Heteragrion silvarum ), whereas more degraded sites were dominated by tolerant Anisoptera (e.g., Erythrodiplax basalis , Erythemis sp., and Zenithoptera lanei ). However, contrary to expectations, Anisoptera richness did not consistently increase in streams with lower HII, although their relative abundance was higher in these environments. This pattern indicates that species replacement (turnover) predominated over diversity loss (nestedness), suggesting that environmental degradation promotes a reorganization of Odonata assemblages. The relationship between limnological variables and species composition confirmed that conductivity and detritus load were the main environmental drivers structuring the communities, reinforcing that physical integrity and environmental heterogeneity are key determinants of the differential distribution of Anisoptera and Zygoptera in Amazonian streams. Environmental variables such as temperature, conductivity, pH, and dissolved oxygen were closely associated with habitat integrity. More degraded streams showed higher conductivity and lower dissolved oxygen concentrations, indicating an imbalance in physicochemical conditions. In well-preserved streams (i.e. with high HII), conductivity tends to be lower due to the limited input of dissolved ions, whereas in impacted streams (low HII) there is an increase in ionic load from diffuse sources such as surface runoff and erosion (Barros et al. 2014). An increase in water temperature, for example, reduces oxygen solubility because of greater molecular agitation (Morenco et al. 2014), directly affecting the physiological, metabolic, and behavioral processes of aquatic organisms (Allan and Castillo 2007 ). These results reinforce the importance of habitat integrity as a key predictor of Odonata assemblage structure, corroborating previous studies that highlight the group as an efficient bioindicator of environmental quality (Allan and Castillo 2007 ). However, this effect is strongly modulated by riparian vegetation cover, which regulates light input, thermal stability, and the availability of organic detritus (Morenco et al. 2014; Esteves 1998 ). Water conductivity, by reflecting the accumulation of dissolved ions, is directly related to changes in temperature and thermal stability, influencing both the physiology and behavior of ectothermic species (De Marco et al. 2015; Calvão et al. 2018 ). Thus, the interaction between physicochemical and structural variables determines species distribution patterns, with some taxa dependent on intact habitats and others tolerant of altered environments (Vieira 2022 ; Cortezzi et al. 2009 ). Species such as Mnesarete aenea and Heteragrion silvarum were recorded both at sites with high HII values (> 0.5) and in more degraded reaches (< 0.5), indicating ecological plasticity that allow them to persist under different disturbance levels (Corbet 1999 ; Heckman 2008; Heiser and Schmitt 2010 ). Although sensitive, these species can exploit shaded, structurally complex microhabitats even in partially impacted environments, underscoring the importance of maintaining riparian forest fragments for the conservation of Odonata diversity (Bastos et al. 2021 ; Simaika et al. 2016 ). In turn, Argia indicatrix and Epipleoneura susanae were more abundant in streams with intermediate HII values, suggesting an ability to cope with variable environmental conditions (Juen et al. 2014 ). The RDA revealed that high conductivity and low detritus availability are associated with communities dominated by tolerant species, such as A. ascendens, D. obscura , and representatives of the genera Erythemis and Erythrodiplax . These species display traits of ecological generalists, including short life cycles, high dispersal capacity, and elevated tolerance to modified environments, and they are widely used as indicators of ecological degradation (Oliveira-Junior et al. 2013 ). The loss of riparian vegetation reduces the input of organic detritus (e.g., leaves, branches, and plant material) that is essential as a source of food, shelter, and substrate (Garcia-Junior et al. 2019 ; Brito et al. 2024 ). This reduction leads to changes in physicochemical variables, such as increased temperature and decreased dissolved oxygen, which directly influence species survival and reproduction (Carvalho et al. 2018 ). By contrast, species such as Erythemis and Erythrodiplax exhibit high ecological plasticity, being able to oviposit in environments lacking riparian vegetation and withstand adverse conditions (Trindade and Oliveira 2024 ). Conversely, more intact streams showed higher abundances of small, sensitive species such as Epipleoneura susanae and E. metallica (Miguel et al. 2017 ). The differentiated occurrence of these species across sites with varying levels of environmental integrity may be linked to physiological and behavioral traits, since densely vegetated environments provide greater thermal stability and microhabitats suitable for reproduction and feeding (De Marco et al. 2015; Trindade and Oliveira 2024 ). Overall, habitat integrity directly influences species composition, diversity, and abundance by providing suitable conditions for the coexistence of multiple taxa. Degradation, in turn, leads to structural simplification and reduced environmental heterogeneity, resulting in less diverse communities dominated by generalist species (Calvão et al. 2018 ; Trindade and Oliveira 2024 ). Accordingly, our results underscore riparian integrity as a key mediator linking physico-chemical variables to the structure of Odonata communities in the eastern Amazon. Finally, this study expands knowledge on the effects of environmental degradation in Amazonian streams and reinforces the importance of riparian vegetation as a key element for maintaining Odonata diversity and conservation. We recommend implementing management and restoration strategies that prioritize bank conservation and continuous monitoring of aquatic communities, aiming to restore the ecological integrity of these ecosystems. Taken together, the evidence positions Odonata as a cornerstone group for biomonitoring and for tropical aquatic ecosystem restoration. Declarations Authors declare not to have any conflict of interests related to this work. Author Contribution JLSP, JMRP, FMBS, TAG, LFAM, TSM, LJ, JCGO and ACA conceived the study, took and analyzed the data, and wrote the main manuscript.LJ gathered financial support for the study. Acknowledgement We thank Alunorte for supporting the project “Avaliação de biota aquática e vegetação ciliar da hidrografia que influencia a bacia do Murucupi e arredores da Hydro Alunorte” and Mineração Paragominas for supporting the project “Aquatic biota monitoring and assessment upstream and downstream of bauxite pipeline Norsk Hydro Paragominas - Barcarena (Pará, Brazil) – an instream and riverscape approach” (process 20/19) via o Biodiversity Research Consortium Brazil-Norway (BRC), which made this work possible. We are deeply grateful to Jair Costa Filho, Cristian Mendoza, Fabio Santos, Viviane Firmino, Juan Bastos, Ingrid Reis, Beatriz Luz, Natalia Santos, Geovani Gomes, Gabrielly Melo, Bethânia Resende, and Tainã Rocha for field assistance and laboratory identifications. The first author thanks the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the doctoral fellowship, the Universidade Federal do Pará (UFPA), the Graduate Program in Ecology (PPGECO), and the Laboratory of Ecology and Conservation (LABECO). LJ and LFAM thank the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for continuous support through productivity fellowships (processes 304710/2019-9 and 302881/2022-0, respectively). Data Availability The data sued for this study can be downloaded for free from https://figshare.com/s/3156634fcfd006efff23 References Allan JD, Castillo MM (2007) Stream ecology: structure and function of running waters, 2nd edn. Springer, Dordrecht. https://doi.org/10.1007/978-1-4020-5583-6 Alves-Martins F, Brasil LS, Juen L, De Marco P Jr, Stropp J, Hortal J (2019) Metacommunity patterns of Amazonian Odonata: the role of environmental gradients and major rivers. 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17:59:53","extension":"xml","order_by":11,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":160405,"visible":true,"origin":"","legend":"","description":"","filename":"d746392d328c4c49ae9ac03ecb1aa27e1structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-8066154/v1/38f1277eaf4d9a643a656aa3.xml"},{"id":98421273,"identity":"1a3fd677-a79f-4c89-b91b-6acc61c3a7e3","added_by":"auto","created_at":"2025-12-17 16:26:15","extension":"html","order_by":12,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":172236,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-8066154/v1/ce3844db3919b13f09225618.html"},{"id":98421002,"identity":"1311f215-167a-49c6-8e8d-480cf4c1ef29","added_by":"auto","created_at":"2025-12-17 16:22:04","extension":"jpeg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":512548,"visible":true,"origin":"","legend":"\u003cp\u003eGeographic distribution of the 30 sampled streams (igarapés) in the Brazilian Amazon, arrayed along a forest-cover gradient.\u003c/p\u003e","description":"","filename":"floatimage1.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8066154/v1/4b1cb078fdd090e5b34167b0.jpeg"},{"id":98421984,"identity":"c0eaf1cd-f1ee-4cb6-a491-dc4b6581ce83","added_by":"auto","created_at":"2025-12-17 16:30:07","extension":"jpeg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":135808,"visible":true,"origin":"","legend":"\u003cp\u003eOrdination of environmental variables from the sampled streams by Principal Components Analysis. DO = dissolved oxygen; pH = hydrogen potential; Temp = temperature; Cond = electrical conductivity; HII = Habitat Integrity Index.\u003c/p\u003e","description":"","filename":"floatimage2.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8066154/v1/be1d27b139137537437a6984.jpeg"},{"id":98422807,"identity":"fed573e6-e301-4f0e-be8e-e4b093863709","added_by":"auto","created_at":"2025-12-17 16:31:32","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":61775,"visible":true,"origin":"","legend":"\u003cp\u003eOrdination of Odonata species composition in Barcarena streams by Principal Coordinates Analysis. Symbol size is proportional to the Habitat Integrity Index. M.wil = \u003cem\u003eMnesarete williamsoni\u003c/em\u003e; Z.lan = \u003cem\u003eZenithoptera lanei\u003c/em\u003e; M.aen = \u003cem\u003eMnesarete aenea\u003c/em\u003e; H.sil = \u003cem\u003eHeteragrion silvarum\u003c/em\u003e; A.ind = \u003cem\u003eArgia indicatrix\u003c/em\u003e; E.sus = \u003cem\u003eEpipleoneura susanae\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-8066154/v1/46543cfc495097732eec3fa7.png"},{"id":98422098,"identity":"d260f9b5-78bc-402e-a828-bbfbc9de6c50","added_by":"auto","created_at":"2025-12-17 16:30:25","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":70773,"visible":true,"origin":"","legend":"\u003cp\u003eOrdination of Odonata composition as a function of conductivity and detritus amount by Redundancy Analysis. Circle sizes are proportional to the Habitat Integrity Index. (A. asce = \u003cem\u003eArgia ascendens\u003c/em\u003e); (D. obsc = \u003cem\u003eDicterias obscura\u003c/em\u003e); (Erythe = \u003cem\u003eErythemis\u003c/em\u003e sp.); (E. basa = \u003cem\u003eErythrodiplax basalis\u003c/em\u003e); (Erythr = \u003cem\u003eErythrodiplax\u003c/em\u003e sp.); (E. meta = \u003cem\u003eEpipleoneura metallica\u003c/em\u003e); (E. susa = \u003cem\u003eE. susanae\u003c/em\u003e); (Z. Lane = \u003cem\u003eZenithoptera lanei\u003c/em\u003e).\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-8066154/v1/e87253c64e16aedda5d9f33f.png"},{"id":98443650,"identity":"e7c7ce37-7e3a-4a80-a11b-c9c7930a9c0e","added_by":"auto","created_at":"2025-12-17 17:13:58","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1715693,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8066154/v1/5cf84534-4588-4e02-9706-167831972b46.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Responses of Dragonflies (Odonata) to Habitat Integrity and Environmental Heterogeneity in Amazonian Streams: lessons for conservation","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe Amazon region is known as one of the most biodiverse on the planet, harboring an extensive network of interconnected aquatic ecosystems (Garda et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Peres et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Latrubesse et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Metzger et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). These ecosystems play a fundamental role in global stability, contributing to carbon sequestration, rainfall formation, and the storage of the world\u0026rsquo;s largest reserve of fresh water (Brasil et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Despite this importance, the Amazon has undergone rapid landscape transformation driven by the expansion of cattle ranching, agriculture, mining, and timber extraction (Fearnside et al. 2005; Faria et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The reduction or simplification of environmental conditions and ecosystem integrity directly affects resource availability, creating habitat patches with low capacity to meet the ecophysiological requirements of many species, including several groups of aquatic insects (Oliveira-Junior et al. 2022; Rocha et al., \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe above environmental changes, especially those affecting riparian vegetation, directly impact aquatic insect communities (Luiza-Andrade et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and promote the replacement of sensitive species by more tolerant ones (Verberk et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Heino \u0026amp; Gr\u0026ouml;nroos \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Such effects tend to be stronger in species with lower dispersal capacity (Lutinski et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), since they depend heavily on local environmental conditions (Juen et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Brasil et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Similar patterns of shifts in the structure of aquatic insect communities have been observed in Neotropical environments and are mediated by changes in stream limnological characteristics and physical habitat structure (Dias-Silva et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Carvalho et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Monteiro-J\u0026uacute;nior et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eAmong aquatic insects, the order Odonata has been widely used in ecological and biomonitoring studies because of its specific ecophysiological requirements (e.g., Monteiro-J\u0026uacute;nior et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Carvalho et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Brasil et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Members of this order maintain a strong relationship with riparian vegetation structure, with some species restricted to more open-canopy environments, whereas others depend on denser vegetation (Oliveira-Junior et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Carvalho et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Oliveira-Junior \u0026amp; Juen 2019). In Amazonian igarap\u0026eacute;s, Odonata respond rapidly to environmental change owing to the relatively short life cycle of their nymphs, which occupy different aquatic microhabitats (e.g., sandy bottoms, rocky substrates, and submerged vegetation) and exhibit distinct feeding habits (Monteiro-J\u0026uacute;nior et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Carvalho et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Brasil et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Thus, odonates play an important role in food webs and in the population control of other aquatic invertebrates (Corbet \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Carvalho et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). This diversity of niches and ecological functions makes the group highly sensitive to variations in the physicochemical and structural conditions of streams, allowing the detection of subtle changes in the ecosystem (Brasil et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The clear ecological differentiation between the suborders Anisoptera and Zygoptera provides a natural gradient of response to environmental disturbance (Juen and De Marco \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Zygoptera exhibit ecophysiological attributes that favor \u0026ldquo;percher\u0026rdquo; behavior, spending much of their time on perches to defend territories and to carry out mating and oviposition (Heckman 2008; Heiser and Schmitt, \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Carvalho et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). In contrast, most Anisoptera are strong fliers with greater physiological robustness. They require more open habitats with higher solar incidence to regulate body temperature and initiate activity (Corbet \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1999\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eOwing to differences in ecophysiological requirements between the suborders, conserved igarap\u0026eacute;s tend to preserve greater Zygoptera diversity, whereas degraded igarap\u0026eacute;s harbor greater Anisoptera diversity (Monteiro-J\u0026uacute;nior et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2014\u003c/span\u003e, \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Oliveira-Junior et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Oliveira-Junior and Juen 2019). An essential metric for understanding these changes in Odonata communities is beta diversity, which represents the variation in species composition along an environmental gradient (Juen et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Alves-Martins et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Juen and De Marco \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). This metric makes it possible to identify whether species replacement (turnover) or species loss (nestedness) is associated with habitat degradation (Legendre and Condit,2019). In Amazonian igarap\u0026eacute;s, where small variations in limnological and structural conditions can generate large differences in the composition of bioindicator communities (Rivera-P\u0026eacute;rez et al. 2024), beta diversity is a robust tool for understanding how habitat integrity and heterogeneity influence the differential distribution of Anisoptera and Zygoptera along disturbance gradients (Brito et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Bastos et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eAccordingly, we here aimed to: i) describe environmental and structural differences among streams, using limnological and habitat variables to identify which best explain species composition; ii) assess the relationship between variation in beta diversity and environmental variables, testing the hypothesis that streams with greater integrity and physicochemical stability preserve higher Zygoptera richness due to this suborder\u0026rsquo;s stronger dependence on habitat quality and heterogeneity; and iii) evaluate the relationships between habitat integrity and the distribution of Odonata species, testing the hypothesis that species richness of the suborder Anisoptera is higher in environments with lower integrity and greater structural homogeneity.\u003c/p\u003e"},{"header":"Material and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eStudy area\u003c/h2\u003e\u003cp\u003eThe study was conducted in 30 igarap\u0026eacute;s located in the municipalities of Barcarena and Abaetetuba, in the northeastern portion of the state of Par\u0026aacute;, Brazil, during October 2022 (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The regional hydrographic network comprises the Murucupi, Arienga, Arapiranga, Barcarena, and Itaporanga rivers. The climate is humid equatorial, with mean annual precipitation ranging from 1,200 to 1,800 mm and mean annual temperatures between 24\u0026deg;C and 26\u0026deg;C. The predominant vegetation corresponds to sub-evergreen equatorial forest (IBGE 2010).\u003c/p\u003e\u003cp\u003eOriginally, the region was covered by native tropical forest characterized by large trees, as well as secondary forest formations, riparian forest, and floodplain areas influenced by seasonal inundation (Paz et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). From the 1980s onward, extractives were the main local economic activity, that were later replaced by agriculture and livestock, and more recently by urbanization, port activities, and the establishment of industries. These land uses have caused intense reduction and fragmentation of both terrestrial and aquatic habitats.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eBiological sampling\u003c/h3\u003e\n\u003cp\u003eSampling was carried out along a fixed 150 m reach in each stream, subdivided into 10 sections of 15 m, delimited by 11 transects labeled A (downstream) to K (upstream). This subdivision of the water course into standardized sections facilitates sampling and helps to control the spatial distribution of specimens in the field (Shimano et al., \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Brito et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Adult insects were captured by active search along the described transect using an entomological net with a 50 cm diameter. Sampling lasted 1 h 30 min along the 150 m reach and was conducted between 10:00 and 14:00 (Juen et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Captured specimens were placed individually in glassine envelopes and properly labeled with site, section, and collection date.\u003c/p\u003e\u003cp\u003eThe specimens were then submerged in absolute anhydrous ethanol (PA/ACS grade) for a minimum of 18 h to preserve key diagnostic structures such as genitalia, wing venation, and coloration essential for taxonomic identification. After this step, the material was taken to the laboratory, where specimens were identified to species using specialized taxonomic keys (Garrison \u0026amp; Von Ellenrieder \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Garrison et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Lencioni 2005, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), complemented by consultation with specialists and comparison with the reference collection of the Laboratory of Ecology and Conservation at the Federal University of Par\u0026aacute; (LABECO/UFPA). The identified specimens were subsequently cataloged, accessioned, and incorporated into LABECO\u0026rsquo;s scientific collection.\u003c/p\u003e\n\u003ch3\u003eEnvironmental variables\u003c/h3\u003e\n\u003cp\u003eIn each stream, limnological variables were measured using a multiparameter probe (AKSO-87), including water temperature (\u0026deg;C), pH, electrical conductivity (mS /m⁻\u0026sup1;), and dissolved oxygen (mg/L⁻\u0026sup1;). Measurements were taken at three equidistant transects along the channel (downstream, mid-reach, and upstream), and the mean value for each variable was then calculated for each stream.\u003c/p\u003e\u003cp\u003eHabitat integrity was assessed using the Habitat Integrity Index (HII) (Nessimian et al. 2008), which considers physical characteristics and the degree of impact from human activities across 12 parameters related to the structure and physical condition of the stream, its banks, and substrate: land use beyond the riparian zone, riparian forest width, riparian forest preservation state, vegetation condition within a 10 m buffer, retention devices, sediment deposition in the channel, bank structure, undercutting of banks, condition of the streambed, presence of riffles, pools, and meanders, aquatic vegetation, and presence of detritus. The index ranges from 0 (least intact) to 1 (most intact), representing a gradient of environmental degradation. In a recent review, HII was identified as one of the most robust metrics for explaining variation in the composition of aquatic insect communities in the Amazon (Brasil et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e\u003ch2\u003eData analysis\u003c/h2\u003e\u003cp\u003eInitially, each sampled stream was treated as a sampling unit, totaling 30 observations. To reduce the dimensionality of the environmental variables, we applied a Principal Component Analysis (PCA) (Legendre and Legendre, \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). This analysis was used to visualize relationships among pH, temperature, conductivity, and dissolved oxygen. Variables were standardized using z-scores to remove scale effects (Legendre \u0026amp; Legendre \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). The broken-stick criterion was adopted for principal component retention (Jackson \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e1993\u003c/span\u003e), and variables with loadings greater than |0.6| were considered relevant.\u003c/p\u003e\u003cp\u003eThe structure of the Odonata community was visualized using Principal Coordinates Analysis (PCoA) based on Bray\u0026ndash;Curtis distance applied to Hellinger-transformed abundance data, to minimize the effects of rare and abundant species (Legendre \u0026amp; De C\u0026aacute;ceres, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). The Hellinger transformation also makes the composition matrix suitable for PCoA, preventing negative eigenvalues (Legendre \u0026amp; Legendre \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). The HII index was overlaid onto the ordination to visually assess the influence of habitat integrity on assemblage composition.\u003c/p\u003e\u003cp\u003eTo quantify how much of the variation in the Odonata community is explained by environmental variables, we applied Redundancy Analysis (RDA) (Legendre \u0026amp; Legendre \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). The response matrix consisted of Hellinger-transformed abundances, and predictor variables included physicochemical and structural attributes, such as conductivity and amount of detritus. All variables were standardized by z-scores. Variable selection was performed by forward selection with 9,999 permutations. Multicollinearity was assessed using the Variance Inflation Factor (VIF), adopting a threshold of VIF\u0026thinsp;\u0026ge;\u0026thinsp;3 as indicative of collinearity (Zuur et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2010\u003c/span\u003e); all included variables had VIF\u0026thinsp;\u0026lt;\u0026thinsp;3. The significance of the final model was tested with 9,999 permutations. Additionally, we applied Pearson\u0026rsquo;s correlation test to evaluate the relationship between the HII and two community metrics: species richness and the total abundance of Odonata per stream.\u003c/p\u003e\u003cp\u003eAll statistical analyses were conducted in R (R Core Team, 2020) using the vegan package (Oksanen et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), with a 5% significance level.\u003c/p\u003e\u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\u003ch2\u003eCommunity Description\u003c/h2\u003e\u003cp\u003eA total of 519 individuals were collected, representing 57 species distributed across eight families (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The suborder Zygoptera was the most abundant and diverse, with 409 individuals (78.8% of the total) distributed among eight family and 32 species. Within this suborder, the family Protoneuridae was the most numerous, with \u003cem\u003eEpipleoneura metallica\u003c/em\u003e (R\u0026aacute;cenis, 1955) as the most abundant species (58 individuals), followed by \u003cem\u003eMnesarete aenea\u003c/em\u003e (Selys, 1853) with 42 individuals and \u003cem\u003eMnesarete williamsoni\u003c/em\u003e (Garrison, 2006) with 36 individuals. The suborder Anisoptera contributed 110 individuals (21.2% of the total) distributed among 25 species. The family Libellulidae was the most representative, with \u003cem\u003eErythrodiplax basalis\u003c/em\u003e (Kirby, 1897) as the most abundant species (13 individuals), followed by \u003cem\u003eErythrodiplax fusca\u003c/em\u003e (Rambur, 1842) with 11 individuals.\u003c/p\u003e\u003cp\u003eIn addition, five species stood out as potential indicators of the ecological conditions of the sampled streams: \u003cem\u003eAeolagrion dorsale\u003c/em\u003e (Burmeister, 1839), \u003cem\u003ePerithemis tenera\u003c/em\u003e (Say, 1840), \u003cem\u003eAcanthagrion kennedii\u003c/em\u003e (Williamson, 1916), \u003cem\u003eChalcopteryx rutilans\u003c/em\u003e (Rambur, 1842), and \u003cem\u003eErythrodiplax castanea\u003c/em\u003e (Burmeister, 1839). These species exhibit distinct preferences regarding habitat structure and environmental quality. Whereas \u003cem\u003eC. rutilans\u003c/em\u003e and \u003cem\u003eA. dorsale\u003c/em\u003e are associated with more preserved, shaded reaches, \u003cem\u003eE. castanea\u003c/em\u003e and \u003cem\u003eP. tenera\u003c/em\u003e occur more frequently in open, human-altered environments. \u003cem\u003eA. kennedii\u003c/em\u003e, in turn, tends to occupy transitional areas, reflecting intermediate conditions. Taken together, these species provide a valuable ecological interpretation of the environmental status of the studied igarap\u0026eacute;s (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\u003eList of Odonata species collected, and their abundance.\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=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003eTAXON\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e\u003cp\u003eZYGOPTERA / Family\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eGenus\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eSpecies / author\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eAbundance\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eCalopterygidae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eHetaerina\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eH. laesa\u003c/em\u003e Hagen in Selys, 1853\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eH. sangu\u0026iacute;nea\u003c/em\u003e Selys, 1853\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eHetaerina\u003c/em\u003e sp. Hagen in Selys, 1853\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eH. westfalli\u003c/em\u003e R\u0026aacute;cenis, 1968\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eMnesarete\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. aenea\u003c/em\u003e Selys, 1853\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e42\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eMnesarete\u003c/em\u003e sp. Cowley, 1934\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e11\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. williamsoni\u003c/em\u003e Garrison, 2006\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e36\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eCoenagrionidae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eAcanthagrion\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eA. adustum\u003c/em\u003e Williamson, 1916\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eAcanthagrion\u003c/em\u003e sp. Selys, 1876\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eA. ascendens\u003c/em\u003e Calvert, 1909\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e12\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eA. kennedii\u003c/em\u003e Williamson, 1916\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eAcanthallagma\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eA. luteum\u003c/em\u003e Willianson \u0026amp; Willianson, 1924\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eAeolagrion\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eA. dorsale\u003c/em\u003e Burmeister, 1839\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eArgia\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eA. collata\u003c/em\u003e Selys, 1865\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e30\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eA. indicatrix\u003c/em\u003e Calvert, 1909\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eA. infumata\u003c/em\u003e Selys, 1865\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eA. oculata\u003c/em\u003e Hagen in Selys, 1865\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eArgia\u003c/em\u003e sp. Rambur, 1842\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eA. tinctipennis\u003c/em\u003e Selys, 1865\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e20\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eDicteriadidae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eDicterias\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eD. atrosanguinea\u003c/em\u003e Selys, 1862\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eHeteragrionidae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eHeteragrion\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eH. silvarum\u003c/em\u003e Sj\u0026ouml;stedt, 1918\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e30\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eHeteragrion\u003c/em\u003e sp. Selys, 1862\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eOxystigma\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eO. petiolatum\u003c/em\u003e Selys, 1862\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e14\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eOxystigma\u003c/em\u003e sp. Selys, 1862\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003ePolythoridae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eChalcopteryx\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eC. rutilans\u003c/em\u003e Rambur, 1842\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eProtoneuridae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eEpipleoneura\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eEpipleoneura\u003c/em\u003e sp.\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e11\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eE. metallica\u003c/em\u003e R\u0026aacute;cenis, 1955\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e58\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eE. susanae\u003c/em\u003e Pessacq, 2014\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e27\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eNeoneora\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eN. luzmarina\u003c/em\u003e De Marmels, 1989\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003ePhasmoneura\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eP. exigua\u003c/em\u003e Selys, 1886\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eProtoneura\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eP. tenuis\u003c/em\u003e Selys, 1860\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e16\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003ePsaironeura\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eP. tenuissima\u003c/em\u003e Selys. 1886\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eANISOPTERA\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eGomphidae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eZonophora\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eZ. calippus klugi\u003c/em\u003e Schmidt, 1941\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eLibellulidae\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eArgyrothemis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eA. argentea\u003c/em\u003e Ris, 1909\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eBrachymesia\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eB. herbida\u003c/em\u003e Gundlach, 1889\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eDiastatops\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eD. obscura\u003c/em\u003e Fabricius, 1775\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eErythemis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eE. peruviana\u003c/em\u003e Rambur, 1842\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eErythemis\u003c/em\u003e sp. Hagen, 1861\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eErythrodiplax\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eE. basalis\u003c/em\u003e Kirby, 1897\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e13\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eE. cast\u0026acirc;nea\u003c/em\u003e Burmeister, 1839\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eE. fusca\u003c/em\u003e Rambur, 1842\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e11\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eErythrodiplax\u003c/em\u003e sp. Brauer, 1868\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eFylgia\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eF. amazonica amaz\u0026ocirc;nica\u003c/em\u003e Kirby, 1889\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eGynothemis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eG. pumila\u003c/em\u003e Karsch, 1890\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eGynothemis\u003c/em\u003e sp. Calvert, 1909\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eMacrothemis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. hahneli\u003c/em\u003e Ris, 1913\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eMicrathyria\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eM. artemis\u003c/em\u003e Ris, 1911\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eMicrathyria\u003c/em\u003e sp. Kirby, 1889\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eOligoclada\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eOligoclada\u003c/em\u003e sp. Karsch, 1890\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eOrthemis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eOrthemis biolleyi\u003c/em\u003e Calvert, 1906\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eO. cultriformis\u003c/em\u003e Calvert, 1899\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003ePerithemis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eP. lais\u003c/em\u003e Perty, 1834\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eP. tenera\u003c/em\u003e Say, 1840\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eP. thais\u003c/em\u003e Kirby, 1889\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eUracis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eU. imbuta\u003c/em\u003e Burmeister, 1839\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eZenithoptera\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eZ. lanei\u003c/em\u003e Santos, 1941\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eZenithoptera_\u003c/em\u003esp. Selys, 1869\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eRichness and abundance\u003c/h3\u003e\n\u003cp\u003eIn the sampled streams, species richness ranged from 4 to 17 (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD\u0026thinsp;=\u0026thinsp;8.07\u0026thinsp;\u0026plusmn;\u0026thinsp;2.95). Despite this variation, no significant correlation was observed between the HII and total richness (r\u0026thinsp;=\u0026thinsp;\u0026minus;\u0026thinsp;0.146; p\u0026thinsp;=\u0026thinsp;0.440), indicating that habitat integrity did not consistently influence the number of recorded species. Abundance showed a wider range, from 4 to 41 individuals (10.82 mean\u0026thinsp;\u0026plusmn;\u0026thinsp;17.30 SD). As with richness, the correlation between HII and abundance was not significant (r\u0026thinsp;=\u0026thinsp;0.216; p\u0026thinsp;=\u0026thinsp;0.252), suggesting that the habitat integrity gradient did not determine differences in the number of individuals among streams.\u003c/p\u003e\n\u003ch3\u003eEnvironmental variables of streams\u003c/h3\u003e\n\u003cp\u003eThe PCA explained 72.52% of the total variation of the set of environmental variables in the first two axes (Component 1\u0026thinsp;=\u0026thinsp;46.44%; Component 2\u0026thinsp;=\u0026thinsp;25.98%). Dissolved oxygen (DO) showed the highest positive correlation with the first axis, while conductivity exhibited the highest negative correlation (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e; Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Visually, streams with higher HII values tended to cluster on the positive side of axis 1, associated with higher DO, whereas streams with lower HII values clustered on the negative side, associated with higher conductivity (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). This pattern indicates that the first axis represents a habitat-integrity gradient, in which more intact environments are characterized by higher oxygenation and lower conductivity (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). On the second axis, pH and temperature showed the strongest positive correlations, contributing similarly to the formation of this component.\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\u003eLoadings of environmental variables on the first two axes of the PCA conducted with the set of streams limnological variables. Variables with loadings \u0026gt;|0.6| are in bold.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"3\"\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=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eVariables\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003ePCA 1\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePCA 2\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003epH\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.321\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e0.794\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eConductivity\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.903\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-0.129\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDissolved oxygen\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e0.872\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.164\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTemperature\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.372\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e0.706\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eEigenvalue\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e2.322\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e1.299\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eBroken-stick\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e2.283\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e1.283\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003e% Explanation\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e46.44\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e25.98\u003c/b\u003e\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\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\u003ch2\u003eSpecies composition\u003c/h2\u003e\u003cp\u003eThe ordination obtained by Principal Coordinates Analysis (PCoA), based on Bray\u0026ndash;Curtis distance, explained 25.82% of the total variation in species composition across the first two axes (PCoA1\u0026thinsp;=\u0026thinsp;13.85%; PCoA2\u0026thinsp;=\u0026thinsp;11.97%). Axis 1 indicated a partial separation of Odonata assemblages along the habitat-integrity gradient.\u003c/p\u003e\u003cp\u003eReaches with higher HII values tended to cluster on the left side of the ordination, associated with species characteristic of more shaded, structurally heterogeneous environments, such as \u003cem\u003eEpipleoneura susanae\u003c/em\u003e, \u003cem\u003eMnesarete williamsoni\u003c/em\u003e, \u003cem\u003eM. aenea\u003c/em\u003e, and \u003cem\u003eHeteragrion silvarum\u003c/em\u003e. Conversely, reaches with lower HII values were positioned to the right, showing greater representation of heliophilous species tolerant of open, modified environments, such as \u003cem\u003eArgia indicatrix\u003c/em\u003e and \u003cem\u003eZenithoptera lanei\u003c/em\u003e. The second axis reflected an additional organization of the Odonata community along the environmental gradient, albeit with lower explanatory power. Overall, the ordination indicated that variation in species composition is driven primarily by habitat integrity and the structural heterogeneity of the streams (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e\u003ch2\u003eRelationship between environmental variables and species composition\u003c/h2\u003e\u003cp\u003eRDA indicated that higher conductivity and lower amounts of detritus were the environmental variables that best explained variation in the structure of the Odonata community, yielding a statistically significant model (F₂,\u003csub\u003e27\u003c/sub\u003e = 1.82; R\u0026sup2;adj\u0026thinsp;=\u0026thinsp;0.054; p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). The first two RDA axes together explained 11.88% of the total variation in species composition (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Streams with lower HII had less detritus and higher conductivity, reflecting more impacted conditions. These reaches were associated with higher abundance of \u003cem\u003eEpipleoneura susanae\u003c/em\u003e, a species that showed moderate tolerance to altered conditions. Conversely, sites with higher conductivity showed greater occurrence of \u003cem\u003eEpipleoneura metallica\u003c/em\u003e, suggesting that this species may benefit from more altered physicochemical environments (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Overall, the RDA results show that water quality and detritus availability are key determinants of Odonata assemblage structure, reflecting the combined effects of environmental degradation and habitat heterogeneity on species composition.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eOur results partially supported the proposed hypotheses that conserved igarap\u0026eacute;s would show greater Zygoptera diversity, whereas degraded igarap\u0026eacute;s would harbor greater Anisoptera diversity. Thus, conserved igarap\u0026eacute;s serve to protect Zygoptera. Accordingly, the species composition was strongly associated with the habitat-integrity gradient, with better-preserved streams showing higher relative abundance and richness of Zygoptera (e.g., \u003cem\u003eEpipleoneura susanae\u003c/em\u003e, \u003cem\u003eMnesarete williamsoni\u003c/em\u003e, \u003cem\u003eM. aenea\u003c/em\u003e, and \u003cem\u003eHeteragrion silvarum\u003c/em\u003e), whereas more degraded sites were dominated by tolerant Anisoptera (e.g., \u003cem\u003eErythrodiplax basalis\u003c/em\u003e, \u003cem\u003eErythemis\u003c/em\u003e sp., and \u003cem\u003eZenithoptera lanei\u003c/em\u003e). However, contrary to expectations, Anisoptera richness did not consistently increase in streams with lower HII, although their relative abundance was higher in these environments. This pattern indicates that species replacement (turnover) predominated over diversity loss (nestedness), suggesting that environmental degradation promotes a reorganization of Odonata assemblages.\u003c/p\u003e\u003cp\u003eThe relationship between limnological variables and species composition confirmed that conductivity and detritus load were the main environmental drivers structuring the communities, reinforcing that physical integrity and environmental heterogeneity are key determinants of the differential distribution of Anisoptera and Zygoptera in Amazonian streams. Environmental variables such as temperature, conductivity, pH, and dissolved oxygen were closely associated with habitat integrity. More degraded streams showed higher conductivity and lower dissolved oxygen concentrations, indicating an imbalance in physicochemical conditions. In well-preserved streams (i.e. with high HII), conductivity tends to be lower due to the limited input of dissolved ions, whereas in impacted streams (low HII) there is an increase in ionic load from diffuse sources such as surface runoff and erosion (Barros et al. 2014). An increase in water temperature, for example, reduces oxygen solubility because of greater molecular agitation (Morenco et al. 2014), directly affecting the physiological, metabolic, and behavioral processes of aquatic organisms (Allan and Castillo \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). These results reinforce the importance of habitat integrity as a key predictor of Odonata assemblage structure, corroborating previous studies that highlight the group as an efficient bioindicator of environmental quality (Allan and Castillo \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eHowever, this effect is strongly modulated by riparian vegetation cover, which regulates light input, thermal stability, and the availability of organic detritus (Morenco et al. 2014; Esteves \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). Water conductivity, by reflecting the accumulation of dissolved ions, is directly related to changes in temperature and thermal stability, influencing both the physiology and behavior of ectothermic species (De Marco et al. 2015; Calv\u0026atilde;o et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Thus, the interaction between physicochemical and structural variables determines species distribution patterns, with some taxa dependent on intact habitats and others tolerant of altered environments (Vieira \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Cortezzi et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2009\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eSpecies such as \u003cem\u003eMnesarete aenea\u003c/em\u003e and \u003cem\u003eHeteragrion silvarum\u003c/em\u003e were recorded both at sites with high HII values (\u0026gt;\u0026thinsp;0.5) and in more degraded reaches (\u0026lt;\u0026thinsp;0.5), indicating ecological plasticity that allow them to persist under different disturbance levels (Corbet \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Heckman 2008; Heiser and Schmitt \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Although sensitive, these species can exploit shaded, structurally complex microhabitats even in partially impacted environments, underscoring the importance of maintaining riparian forest fragments for the conservation of Odonata diversity (Bastos et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Simaika et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). In turn, \u003cem\u003eArgia indicatrix\u003c/em\u003e and \u003cem\u003eEpipleoneura susanae\u003c/em\u003e were more abundant in streams with intermediate HII values, suggesting an ability to cope with variable environmental conditions (Juen et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe RDA revealed that high conductivity and low detritus availability are associated with communities dominated by tolerant species, such as \u003cem\u003eA. ascendens, D. obscura\u003c/em\u003e, and representatives of the genera \u003cem\u003eErythemis\u003c/em\u003e and \u003cem\u003eErythrodiplax\u003c/em\u003e. These species display traits of ecological generalists, including short life cycles, high dispersal capacity, and elevated tolerance to modified environments, and they are widely used as indicators of ecological degradation (Oliveira-Junior et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe loss of riparian vegetation reduces the input of organic detritus (e.g., leaves, branches, and plant material) that is essential as a source of food, shelter, and substrate (Garcia-Junior et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Brito et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). This reduction leads to changes in physicochemical variables, such as increased temperature and decreased dissolved oxygen, which directly influence species survival and reproduction (Carvalho et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). By contrast, species such as \u003cem\u003eErythemis\u003c/em\u003e and \u003cem\u003eErythrodiplax\u003c/em\u003e exhibit high ecological plasticity, being able to oviposit in environments lacking riparian vegetation and withstand adverse conditions (Trindade and Oliveira \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eConversely, more intact streams showed higher abundances of small, sensitive species such as \u003cem\u003eEpipleoneura susanae\u003c/em\u003e and \u003cem\u003eE. metallica\u003c/em\u003e (Miguel et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The differentiated occurrence of these species across sites with varying levels of environmental integrity may be linked to physiological and behavioral traits, since densely vegetated environments provide greater thermal stability and microhabitats suitable for reproduction and feeding (De Marco et al. 2015; Trindade and Oliveira \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Overall, habitat integrity directly influences species composition, diversity, and abundance by providing suitable conditions for the coexistence of multiple taxa. Degradation, in turn, leads to structural simplification and reduced environmental heterogeneity, resulting in less diverse communities dominated by generalist species (Calv\u0026atilde;o et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Trindade and Oliveira \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Accordingly, our results underscore riparian integrity as a key mediator linking physico-chemical variables to the structure of Odonata communities in the eastern Amazon.\u003c/p\u003e\u003cp\u003eFinally, this study expands knowledge on the effects of environmental degradation in Amazonian streams and reinforces the importance of riparian vegetation as a key element for maintaining Odonata diversity and conservation. We recommend implementing management and restoration strategies that prioritize bank conservation and continuous monitoring of aquatic communities, aiming to restore the ecological integrity of these ecosystems. Taken together, the evidence positions Odonata as a cornerstone group for biomonitoring and for tropical aquatic ecosystem restoration.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eAuthors declare not to have any conflict of interests related to this work.\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eJLSP, JMRP, FMBS, TAG, LFAM, TSM, LJ, JCGO and ACA conceived the study, took and analyzed the data, and wrote the main manuscript.LJ gathered financial support for the study.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eWe thank Alunorte for supporting the project \u0026ldquo;Avalia\u0026ccedil;\u0026atilde;o de biota aqu\u0026aacute;tica e vegeta\u0026ccedil;\u0026atilde;o ciliar da hidrografia que influencia a bacia do Murucupi e arredores da Hydro Alunorte\u0026rdquo; and Minera\u0026ccedil;\u0026atilde;o Paragominas for supporting the project \u0026ldquo;Aquatic biota monitoring and assessment upstream and downstream of bauxite pipeline Norsk Hydro Paragominas - Barcarena (Par\u0026aacute;, Brazil) \u0026ndash; an instream and riverscape approach\u0026rdquo; (process 20/19) via o Biodiversity Research Consortium Brazil-Norway (BRC), which made this work possible. We are deeply grateful to Jair Costa Filho, Cristian Mendoza, Fabio Santos, Viviane Firmino, Juan Bastos, Ingrid Reis, Beatriz Luz, Natalia Santos, Geovani Gomes, Gabrielly Melo, Beth\u0026acirc;nia Resende, and Tain\u0026atilde; Rocha for field assistance and laboratory identifications. The first author thanks the Coordena\u0026ccedil;\u0026atilde;o de Aperfei\u0026ccedil;oamento de Pessoal de N\u0026iacute;vel Superior (CAPES) for the doctoral fellowship, the Universidade Federal do Par\u0026aacute; (UFPA), the Graduate Program in Ecology (PPGECO), and the Laboratory of Ecology and Conservation (LABECO). LJ and LFAM thank the Conselho Nacional de Desenvolvimento Cient\u0026iacute;fico e Tecnol\u0026oacute;gico (CNPq) for continuous support through productivity fellowships (processes 304710/2019-9 and 302881/2022-0, respectively).\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eThe data sued for this study can be downloaded for free from https://figshare.com/s/3156634fcfd006efff23\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAllan JD, Castillo MM (2007) Stream ecology: structure and function of running waters, 2nd edn. 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Methods Ecol Evol 1(1):3\u0026ndash;14. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.2041-210X.2009.00001.x\u003c/span\u003e\u003cspan address=\"10.1111/j.2041-210X.2009.00001.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"journal-of-insect-conservation","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jico","sideBox":"Learn more about [Journal of Insect Conservation](http://link.springer.com/journal/10841)","snPcode":"10841","submissionUrl":"https://submission.nature.com/new-submission/10841/3","title":"Journal of Insect Conservation","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Odonata, stream ecology, land uses impact, bioindicators, conservation","lastPublishedDoi":"10.21203/rs.3.rs-8066154/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8066154/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eRapid landscape transformation in the Amazon, driven by human activities such as deforestation and intensive land use, has deeply affected the conservation of their aquatic ecosystems, especially small forest streams (i.e. \u0026ldquo;igarap\u0026eacute;s\u0026rdquo;). These environments rely heavily on riparian vegetation to maintain the physical and chemical conditions that support biodiversity. Among the many threatened aquatic organisms, insects of the order Odonata stand out as sensitive bioindicators of habitat alteration. We here evaluated the effects of environmental variables on the distribution of adult Odonata species in streams of the Eastern Amazon. We expected more intact streams to harbor higher Zygoptera richness, whereas less intact streams would show higher Anisoptera richness. We sampled 30 streams along a gradient of environmental alteration. We found that better-preserved streams were also better at preserving Zygoptera diversity while more degraded environments were dominated by tolerant Anisoptera as predicted. However, Anisoptera richness did not significantly increase in low-HII streams, although their relative abundance was higher there, suggesting a more pronounced species turnover than diversity loss. In addition, limnological variables, such as electrical conductivity and coarse particulate detritus, played central roles in structuring communities. These results demonstrate the efficiency of adult Odonata as bioindicators of environmental change and reinforce their use as an effective tool for their conservation and assessing environmental quality in Amazonian streams.\u003c/p\u003e","manuscriptTitle":"Responses of Dragonflies (Odonata) to Habitat Integrity and Environmental Heterogeneity in Amazonian Streams: lessons for conservation","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-12-10 17:59:48","doi":"10.21203/rs.3.rs-8066154/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-02-11T07:40:04+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-02-03T14:11:13+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-01-07T19:56:36+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"199598138627567432158551762392296791036","date":"2025-12-15T14:19:51+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"302740286873970873546701287095903360026","date":"2025-12-08T12:36:51+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-12-07T20:58:11+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-11-10T07:21:12+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-11-10T07:20:45+00:00","index":"","fulltext":""},{"type":"submitted","content":"Journal of Insect Conservation","date":"2025-11-08T21:26:57+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"journal-of-insect-conservation","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jico","sideBox":"Learn more about [Journal of Insect Conservation](http://link.springer.com/journal/10841)","snPcode":"10841","submissionUrl":"https://submission.nature.com/new-submission/10841/3","title":"Journal of Insect Conservation","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"b4655f63-d2eb-4a6c-9217-3dde3ff6aece","owner":[],"postedDate":"December 10th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-03-25T01:53:23+00:00","versionOfRecord":[],"versionCreatedAt":"2025-12-10 17:59:48","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8066154","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8066154","identity":"rs-8066154","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}