{"paper_id":"41d7d17e-d62a-4f4a-8db7-b4b7ddc5e1db","body_text":"1\n1 Thorax temperature and niche characteristics as predictors of abundance of Amazonian Odonata\n2\n3 Lenize Batista Calvão1,2*, Ana Paula J. Faria1,2, Carina Kaory Sasahara de Paiva2, José Max Barbosa \n4 Oliveira-Junior3, Javier Muzón4, Alex Córdoba-Aguillar*5, Leandro Juen1,2\n5\n6 1Programa de Pós-Graduação em Ecologia, Universidade Federal do Pará, Belém, Pará, Brazil \n7 2Laboratório de Ecologia e Conservação (LABECO), Instituto de Ciências Biológicas (ICB), Universidade \n8 Federal do Pará (UFPA), Belém, Pará, Brazil\n9 3 Instituto de Ciências e Tecnologia das Águas (ICTA), Universidade Federal do Oeste do Pará (UFOPA), \n10 Santarém, Pará, Brazil\n11 4 Laboratorio de Biodiversidad y Genética Ambiental (BioGeA), Universidad Nacional de Avellaneda, \n12 Avellaneda, Buenos Aires, Argentina\n13 5 Departamento de Ecología Evolutiva, Instituto de Ecología, Universidad Nacional Autónoma de México, \n14 , Ciudad Universitaria, Mexico City, Mexico\n15\n16 *Corresponding authors:\n17 E-mail: lenizecalvao@gmail.com  acordoba@iecologia.unam.mx \n.CC-BY 4.0 International licenseperpetuity. It is made available under a \npreprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in \nThe copyright holder for thisthis version posted September 16, 2024. ; https://doi.org/10.1101/2024.09.14.613059doi: bioRxiv preprint \n\n2\n18 Abstract\n19 Environmental architecture and body temperature drive the distribution of ectothermic species, \n20 especially those with specific ecophysiological requirements or narrow ecological niches. In this study, \n21 we evaluated the connection between thorax temperature and niche specialization concerning the \n22 abundance and species contribution to the beta diversity of adult Odonata in Amazonian streams, \n23 employing the Species Contribution to Beta Diversity (SCBD). Our hypotheses were (i) Odonata \n24 species’ thorax temperature is positively correlated with both morphology (thorax width) and air \n25 temperature, and (ii) the thorax temperature of the Odonata assemblage serves as a more influential \n26 predictor than niche specialization in determining species abundance and SCBD. We sampled 46 \n27 streams in an anthropized landscape in the Northeastern and Southeastern regions of Pará state, Brazil. \n28 Notably, niche breadth emerged as the variable influencing the abundance and SCBD of the Odonata \n29 assemblage. Niche position is a predictor for Odonata SCBD and not suborders, and predictor for \n30 abundance, except for Anisoptera. Both suborders exhibited a negative relationship between abundance \n31 and thoracic temperature. In summary, our results underscore the necessity of considering both niche \n32 and ecophysiological predictors to comprehensively assess the Odonata assemblage in Amazonian \n33 streams. This holistic approach has implications for conservation efforts and bioassessment practices, \n34 offering valuable insights into the collective response of Odonata as a group.\n35\n36 Introduction\n37 One aim in ecology is to understand how species assemblages distribute as a function of each \n38 species’ requirements. In this regard, ecological niche breadth and position are key predictors for the local \n39 abundance [1–4]. The specific link that underlies species abundance and such predictors is explained \n40 according to two models. The hypothesis that links niche breadth/tolerance, predicts that populations that \n41 can remain viable in a wide range of environmental conditions have greater abundance. In this context, they \n42 can be characterized as generalists (occur in a wide range of environmental conditions) and specialists \n43 (occur in a more restricted range). Conversely, the hypothesis for the niche position predicts that non-\n44 marginal species, those that occur in a larger availability of habitats, tend to be more abundant(11).\n45 Generalist species, endowed with greater niche breadth, can occur in a wider range of \n46 environmental conditions, leading to a smaller contribution of species for Beta diversity (SCBD) than with \n47 the contribution of specialist species [10]. Accordingly, niche position can influence SCBD, as species in \n48 marginal habitats use more specific environmental conditions than non-marginal species [10]. \n.CC-BY 4.0 International licenseperpetuity. It is made available under a \npreprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in \nThe copyright holder for thisthis version posted September 16, 2024. ; https://doi.org/10.1101/2024.09.14.613059doi: bioRxiv preprint \n\n3\n49 Simultaneously, the relationship of functional traits (e.g., thermoregulatory ability governed by body size) \n50 with SCBD may occur if traits confer adaptations in the species, influencing their abundance and/or \n51 occupancy at the site [2, 4]. \n52 Odonata are aquatic insects whose thermal performance determines their distribution both at the \n53 micro- [12] and macro-scale [13]. In this context, the body size and behavior of adult odonates are \n54 associated with their thermoregulation strategies, categorizing species as thermal conformers, heliothermic \n55 or endothermic [12]. These thermoregulatory strategies are correlated with body size, as heat exchange with \n56 the environment occurs based on the surface/volume ratio [14]. Consequently, small percher species (e.g., \n57 most zygopterans) generally exhibit thermal conformers or heliothermic, relying on air temperature to start \n58 their activities [12]. These species can regulate heat loss by adjusting their body posture in response to light \n59 [15–16] and by selecting habitats that support their thermal requirements [14, 17]. Conversely, larger \n60 species of Odonata can inhabit open areas with reduced canopy cover [12] as they are heliothermic [18], \n61 benefiting from direct sunlight exposure. Finally, larger species (e.g., mostly Anisoptera) are predominantly \n62 endothermic, enabling them to overcome the limitations faced by thermal conformers, as they can internally \n63 control heat loss or production through their thorax muscles [14].\n64 Ecophysiological requirements have frequently been crucial predictors for odonate abundance \n65 or distribution in tropical streams [18, 19, 20, 21]. This correlation is primarily supported by the impact of \n66 changes in vegetation cover and air temperature, both acting as environmental filters influencing odonate \n67 community composition [21]. For example, the absence of vegetation and reduced shade favor most \n68 anisopteran species due to their reliance on increased light input into the stream [12, 18, 20, 21, 22]. \n69 Conversely, these environmental conditions are unsuitable for most Zygoptera or smaller species, as they \n70 typically prefer habitats with a more consistent temperature and may not be able to thermoregulate \n71 effectively in altered streams [20, 21, 22]. \n72 In the present study, we aimed to evaluate the importance of thoracic temperature and niche \n73 specialization on the abundance and contribution of each species to the SCBD of adult Odonata in \n74 Amazonian streams. For this, we had the following predictions: (i) thoracic temperature of Odonata species \n75 differs between suborders and correlates positively with morphology (thorax width) and air temperature. \n76 Thus, larger species (Anisoptera) may exhibit heliothermic behaviors, leading to thoracic temperature \n77 higher than air temperature. Conversely, smaller species (Zygoptera) tend to thermoregulate in response to \n78 air temperature, resulting in thorax temperature similar to the ambient environment; (ii) the thoracic \n.CC-BY 4.0 International licenseperpetuity. It is made available under a \npreprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in \nThe copyright holder for thisthis version posted September 16, 2024. ; https://doi.org/10.1101/2024.09.14.613059doi: bioRxiv preprint \n\n4\n79 temperature of the Odonata assemblage, as well as its suborders (Zygoptera and Anisoptera), serves as a \n80 more important predictor than niche characteristics in predicting the abundance and SCBD of the species \n81 given that ecophysiological traits play a crucial role in Odonata habitat selection (Fig 1).\n82 Fig 1. Schematic drawing of hypotheses and images of Odonata were adapted from Stehr, F.W and \n83 Tennessen, K.J.\n84 Material and methods\n85 Study area\n86 The study was conducted in 46 streams across four municipalities in the Northeastern region \n87 (Tomé-açu, Ipixuna do Pará, Concórdia do Pará and Acará) and three municipalities in the Southeastern \n88 region (Paragominas, Canaã dos Carajás and Parauapebas) of Pará state, Brazil, covering basins ranging \n89 from the Tocantins-Araguaia River and Capim River (Fig 2, S1 Table). Northeastern Pará is characterized, \n90 according to the Köppen classification, by a tropical rainforest climate (Af) and tropical monsoon climate \n91 (Am) [23], with temperature ranging from 22°C to 34°C (minimum: 22°C to 23°C; maximum: 30°C to \n92 34°C). Southeastern Pará has a Savanna climate, classified as a tropical climate with a dry season (Aw) \n93 [23, 24, 25]. The municipalities of Canaã dos Carajás and Parauapebas, located in Serra dos Carajás area, \n94 are notable for their landscape, mainly due to their elevation, which ranges between 500 and 700 m a.s.l. \n95 [24]. The average monthly temperature in this area varies from approximately 25°C to 26°C, with  an annual \n96 rainfall of 2.033 millimeters [24]. \n97 Fig 2. Study area showing the 46 streams distributed in the Northeastern and Southeastern of Pará, Brazil. \n98 Images of Odonata were adapted from Stehr, F.W and Tennessen, K.J.\n99 Insect sampling\n100 The 46 sampled streams ranged from 1st to 3rd order, with an average width of 2.8 meters and a \n101 depth of 29.9 centimeters, according to Strahler [26] classification. Sampling periods were performed \n102 between July 2017 and October 2018, consistently during the low precipitation period. Adult odonates were \n103 collected only once stream, in 20 segments spaced 5m apart, distributed along a continuous 150-meter \n104 longitudinal stretch in each stream [further details in 20, 27, 28]. Specimens were collected on stream banks \n105 for one hour, always between 11:00 and 12:00 hrs (the peak activity period for adult odonates), at \n106 temperatures above 19°C, using a 40cm diameter and 65cm in length entomological net [21, 29]. \n.CC-BY 4.0 International licenseperpetuity. It is made available under a \npreprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in \nThe copyright holder for thisthis version posted September 16, 2024. ; https://doi.org/10.1101/2024.09.14.613059doi: bioRxiv preprint \n\n5\n107 Standardization of sampling effort and climatic conditions was necessary due to the organisms' \n108 thermoregulatory ability based on solar incidence [30], ensuring the presence of all ecophysiological groups \n109 [21].\n110 The collected specimens were placed in tracing paper envelopes and subsequently fixed in \n111 acetone P.A. (Propanone) for 12 hours for Zygoptera and 48-72 hours for Anisoptera. Species identification \n112 was conducted to the species level using specialized taxonomic keys [31–35]. The biological material was \n113 deposited in the Collection of Aquatic Insects at the Laboratory of Ecology and Conservation, Federal \n114 University of Pará (UFPA), University Campus of Belém, Pará, Brazil [36].\n115 Measurements of physiological traits \n116 The thoracic temperature of males was captured using an entomological net and quickly \n117 measured within a maximum of 10 seconds to avoid physical damage and alteration in body temperature. \n118 We used only male individuals because species identification is possible in this sex. Holding the wings, we \n119 measured the body temperature (°C) of each individual using a Testo-805 infrared thermometer (accuracy \n120 ± 1°C [-2 to +40°C], resolution 0.1°C [-9.9 to +199.9°C], and reaction time <1s). The recording was \n121 performed by pointing the thermometer beam at the center of the right side of the thorax, at a distance of \n122 five centimeters.\n123 Thorax width was measured using a digital caliper (accuracy = 0.02 mm) for males of each species. \n124 For species with multiple individuals, we used the averages, while for those with only one individual, we \n125 recorded the absolute value. \n126 Environmental variables in streams \n127 For each stream, four environmental variables were measured: depth (cm), channel width (m), \n128 Habitat Integrity Index (HII) and air temperature (in ºC). Depth was measured at three points of the stream: \n129 left and right of the bank, and central region. The average of these measurements was used as a measure of \n130 depth in each sampling unit. The HII was composed by 12 items that describe the degree of habitat integrity \n131 in the stream: land use pattern beyond the riparian zone; width of riparian forest; completeness of riparian \n132 forest, vegetation of riparian zone within 10 m of channel,; retention devices and sediment in the channel; \n133 river bank structure; bank undercutting, stream bottom; distribution of riffles and pools; characteristics of \n134 aquatic vegetation and detritus [37]. Each item presented four to six alternatives corresponding to the \n135 observed condition related to habitat integrity. We transformed each item value to produce the HII, which \n.CC-BY 4.0 International licenseperpetuity. It is made available under a \npreprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in \nThe copyright holder for thisthis version posted September 16, 2024. ; https://doi.org/10.1101/2024.09.14.613059doi: bioRxiv preprint \n\n6\n136 ranges from 0 (altered stream) to 1 (preserved stream), according to the habitat integrity conditions found \n137 in each stream [37]. Note that this HII has been widely used in studies that have assessed the environmental \n138 conditions of streams and their relationship with aquatic insect diversity [38–40]. Air temperature was \n139 measured using a Testo-805 infrared thermometer. After measuring the thorax temperature of the \n140 specimens, the thermometer was positioned in the microhabitat where each individual was collected. \n141 Data analysis\n142 To calculate niche breadth and position (i.e., predictor variables) of the species, we used the \n143 Outlying Mean Index (OMI [11]) which is based on the following environmental variables: depth, channel \n144 width, and HII. The OMI analysis calculates the distance between the average environmental conditions \n145 used by the species (centroid) and the average environmental conditions of the sampled sites (hyperspace) \n146 [11,41]. Species abundance data were logarithmized and environmental variables standardized. The \n147 marginality significance (OMI) of the species was evaluated using the Monte-Carlo permutation test with \n148 1000 permutations [11]. Thus, we obtained the breadth (environmental tolerance) and position (marginality) \n149 of the ecological niche for each species. We carried out PCoA for Odonata composition visualization using \n150 Hellinger and Bray-Curtis.\n151 The ecological uniqueness of species (SCBD) was measured from the total variation of the \n152 assemblage (Total Beta Diversity - BD Total) between streams. For this analysis, we submitted the Odonata \n153 composition matrix to the Hellinger transformation, which was used to measure the Sum of Total Squares \n154 (SSTotal). Then, we divided the SS Total by the number of sampled streams (n-1) and obtained the BD Total of \n155 the assemblage that was partitioned into ecological uniqueness of species (SCBD) [9]. For more details on \n156 the calculation of BDTotal and SCBD, see [9]. Finally, species with higher SCBD values had a higher relative \n157 contribution to beta diversity [9].\n158 We performed the analysis of the relationship between the variables from the generalized linear \n159 mixed models (GLMMs) and species as random factors. Thus, to assess (i) the difference in thoracic \n160 temperature of Anisoptera and Zygoptera, the relationship between the interaction of air temperature and \n161 thorax width on Odonata thoracic temperature, and the relationship between the difference of air and \n162 thoracic temperature (Tair-Tth [ºC]) and thorax width for Anisoptera and Zygoptera, with Gaussian \n163 distribution. The analyzed models contained each individual as a sample unit.\n164 To evaluate (ii) the effect of thoracic temperature, niche breadth and position (predictor variables \n165 standardized) on the abundance (response variable) of Odonata species, we carried out a GLMM and \n.CC-BY 4.0 International licenseperpetuity. It is made available under a \npreprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in \nThe copyright holder for thisthis version posted September 16, 2024. ; https://doi.org/10.1101/2024.09.14.613059doi: bioRxiv preprint \n\n7\n166 suborder as a random factor, using the negative binomial distribution due to data overdispersion [42]. For \n167 this model we used the Log linkage function. The analyzed models contained species as a sample unit. For \n168 Suborders, we carried out a GLM (Negative Binomial). We performed the visual validation of the models \n169 using the simulated envelope [43]. To assess the effect of thoracic temperature, niche breadth and position \n170 (predictor variables standardized) on Odonata and suborders SCBD (response variable), we used a Beta \n171 Regression [44]. This analysis is more suitable for the response variable (SCBD) distributed between values \n172 from 0 to 1 [44]. The binding function used in this model was logit.\n173 All analyses were performed using the R software [45] using MASS [46], hnp [43], betareg [44], \n174 and vegan packages [47].\n175 Results\n176 Odonata assemblage\n177 859 specimens were collected, belonging to 56 species: 22 anisopterans and 34 zygopterans (S2 \n178 Table).\n179 Thoracic temperature and size, and their relation with ambient temperature \n180  Anisopterans showed greater temperature and thorax width than zygopterans. On average, \n181 Anisoptera species have 5°C more than Zygoptera. Air temperature was higher 2°C in the places where \n182 Anisoptera species were sampled (Table 1, Table 2 and Fig 3). Both thoracic width and air temperature \n183 affect thoracic temperature, and (Table 2). \n184 Table 1. Mean and standard deviation (SD) values of thorax temperature, air temperature, and thorax width \n185 for Zygoptera and Anisoptera sampled in Brazilian Amazon streams.\nAnisoptera\nThorax width (cm) Thorax temperature (ºC) Air temperature (ºC)\nMean 3.439 33.867 31.636\nSD 1.465 3.677 2.013\nZygoptera\nMean 1.663 29.510 30.113\nSD 0.446 1.603 1.327\n186\n187 Table 2. Results of the GLMM (Gaussian distribution) (species random effect) evaluating the relationship \n188 between thorax temperature of Odonata (response variable) and suborders and interaction of air temperature \n189 and thorax width.\n.CC-BY 4.0 International licenseperpetuity. It is made available under a \npreprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in \nThe copyright holder for thisthis version posted September 16, 2024. ; https://doi.org/10.1101/2024.09.14.613059doi: bioRxiv preprint \n\n8\n Value Std.Error DF t-value p-value\nIntercept 29.800 0.257 521 116.042 <0.001\nSuborders 1.761 0.501 54 3.517 <0.001\nThorax width (cm) 0.820 0.171 521 4.809 <0.001\nAir temperature (ºC) 1.159 0.096 521 12.020 <0.001\nThorax width *Air temperature 0.288 0.086 521 3.361 <0.001\n190 Significant p values appear in bold.\n191\n192 Fig 3. Thoracic temperature of Anisoptera (Dark orange) and Zygoptera (Cyan). Black dots are outliers.\n193 Anisopterans presented an average difference of air and thoracic temperature up to 8ºC above air \n194 temperature and 6ºC below air temperature, showing a negative relationship with thorax width (Table 3 and \n195 Fig 4). For Zygoptera, the same pattern was observed with an average difference of 3ºC above and  4ºC  \n196 below air temperature (Table 3 and Fig 4).Table 3. GLMM (Gaussian distribution) (species random effect) \n197 results evaluating the difference between air and thorax temperature (Tair-Tth – variable response) with \n198 thorax width of Anisoptera and Zygoptera (predictor).\nAnisoptera\nValue Std.Error DF t-value p-value\nIntercept 0.753 1.034 192 0.728 0.468\nThorax width (cm) -0.908 0.290 192 -3.132 0.002\nZygoptera\nValue Std.Error DF t-value p-value\nIntercept 2.237 0.578 330 3.867 0.000\nThorax width (cm) -0.983 0.341 330 -2.886 0.004\n199  Significant p value appears in bold.\n200 Fig 4. Relation of the difference between air and thorax temperature (Tair-Tth (ºC) response variable) and \n201 thorax width (cm), for Anisoptera (Dark orange) dots bellow dotted line are species that have the thorax \n202 temperature above air temperature. Zygoptera (Cyan). Images of Odonata from Stehr, F.W and Tennessen, \n203 K.J.\n204 Stream structure and odonate assemblage\n205 There was a relation between environmental variables (depth, channel width and HII) and \n206 Odonata species (p = 0.010) (OMI analysis global test). The variable that contributed the most to the first \n207 sorting axis was HII (0.88), followed by width (-0.22) and depth (0.03). This variable is essential to assess \n208 environmental integrity and demonstrates that Odonata species change composition with more intact \n209 streams (HII above 0.7) and with multiple anthropic activities (Fig 5). Streams with greater habitat integrity \n210 have a greater number of species of Anisoptera and Zygoptera that are only collected in these environments \n211 (S3 Table).\n212 Fig 5. Ordination (PCoA) of Odonata species and habitat integrity index (HII): green dots are HII above \n213 0.7 and Coral dots are streams with multiple anthropic activities (HII bellow 0.7).\n214 Niche and abundance of Odonata \n.CC-BY 4.0 International licenseperpetuity. It is made available under a \npreprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in \nThe copyright holder for thisthis version posted September 16, 2024. ; https://doi.org/10.1101/2024.09.14.613059doi: bioRxiv preprint \n\n9\n215 The most abundant species of Zygoptera were C. rutilans, M. aenea and E. metallica. For \n216 Anisoptera were F. amazonica, E. basalis and P. lais. The average the niche breadth was 0.221 (standard \n217 deviation ± 0.289) and the position 2.219 (±1.94). Species with greater niche breadth were M. cupraea, H. \n218 silvarum and M. aenea (Zygoptera) and E. fusca, O. walkeri and O abbreviata (Anisoptera). For niche \n219 positions were H. auripennis, A. luteum and H. icterops (Zygoptera) and D. obscura, B. herbida and E. \n220 cannacrioides (Anisoptera).\n221  Niche breadth and position were the most important predictors of Odonata abundance and \n222 suborders (Table 4 and Fig 6). Except niche position for Anisoptera. Only when evaluating suborders \n223 separately, thorax temperature emerges as important predictor for them.\n224 Table 4. GLMM results (negative binomial distribution) for Odonata (Suborder as a random effect), and \n225 GLM for suborders evaluating the relationship of species abundance with niche breadth, position and thorax \n226 temperature.\nOdonata\n Value Std.Error DF t-value p-value\nIntercept 2.232 0.188 51 11.878 <0.001\nNiche position (NP) -0.503 0.241 51 -2.085 0.042\nNiche breadth (NB) 0.856 0.189 51 4.529 <0.001\nThorax temperature (ºC) -0.201 0.213 51 -0.942 0.351\nAnisoptera\nEstimate Std. Error\nz \nValue Pr(>|z|)\nIntercept 2.434 0.257 9.482 <0.001\nNiche position (NP) -0.257 0.231 -1.110 0.267\nNiche breadth (NB) 0.850 0.302 2.817 0.005\nThorax temperature (ºC) -0.612 0.219 -2.798 0.005\nZygoptera\nEstimate Std. Error\nz \nValue Pr(>|z|)\nIntercept 2.569 0.285 9.006 <0.001\nNiche position (NP) -0.886 0.310 -2.854 0.004\nNiche breadth (NB) 0.866 0.175 4.963 <0.001\nThorax temperature (ºC) 0.868 0.421 2.064 0.039\n227\n228 Significant p values appear in bold.\n229 On average, SCBD was 0.017 (standard deviation ± 0.026). Species with greater SCBD were \n230 C. rutilans, H. indeprensa and M. aenea for the suborder Zygoptera and E. basalis, O. abbreviata and F. \n231 amazonia for the suborder Anisoptera. When evaluating predictor variables for Odonata's SCBD, niche \n232 breadth and position emerged as important predictor. When evaluating predictor variables for suborders \n233 SCBD, niche breadth only emerged as predictor. (Table 5 and Fig 6).\n.CC-BY 4.0 International licenseperpetuity. It is made available under a \npreprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in \nThe copyright holder for thisthis version posted September 16, 2024. ; https://doi.org/10.1101/2024.09.14.613059doi: bioRxiv preprint \n\n10\n234 Table 5. Beta regression results between SCBD of Odonata (suborders shown separately) and niche \n235 breadth, position, and thorax temperature.\nOdonata\nEstimate\nStd. \nError\nz \nValue Pr(>|z|)\nIntercept -4.174 0.150\n-\n27.860 <0.001\nNiche position (NP) -0.290 0.136 -2.128 0.033\nNiche breadth (NB) 0.404 0.090 4.481 <0.001\nThorax temperature (ºC) -0.034 0.127 -0.268 0.789\nAnisoptera\nEstimate\nStd. \nError\nz \nValue Pr(>|z|)\nIntercept -4.393 0.232\n-\n18.962 <0.001\nNiche position (NP) -0.250 0.173 -1.444 0.149\nNiche breadth (NB) 0.535 0.225 2.375 0.018\nThorax temperature (ºC) -0.111 0.172 -0.643 0.520\nZygoptera\nEstimate\nStd. \nError\nz \nValue Pr(>|z|)\nIntercept -4.053 0.260\n-\n15.606 <0.001\nNiche position (NP) -0.371 0.213 -1.743 0.081\nNiche breadth (NB) 0.384 0.108 3.547 <0.001\nThorax temperature (ºC) 0.058 0.328 0.178 0.859\n236 Significant p values appear in bold.\n237 Fig 6. Significant relationships of Abundance and SCBD of Odonata and the suborders and niche breadth, \n238 position and thorax temperature. Odonata (Black dots), Anisoptera (Dark orange) and Zygoptera (Cyan). \n239 Images of Odonata from Stehr, F.W.\n240 Discussion\n241 Niche breadth was an important predictor for abundance and SCBD for Odonata and its suborders. \n242 Niche position also is important for Odonata abundance and SCBD, and Zygoptera abundance. Contrary \n243 our expected hypothesis, the thoracic temperature was a good predictor only for abundance and when \n244 evaluated suborders separately. Research shows that species with greater niche breadth are more abundant \n245 and tend to present greater regional occupation [6]. Furthermore, these species, often display a more \n246 generalist or tolerant behavior towards diverse environmental conditions, which may contribute to their \n247 reduced vulnerability. [50]. In Odonata, this macroecological pattern may be due to the generalist species \n248 being able to persist in degraded environments [51], resulting in a broader distribution range [52]. In our \n249 study the families Libellulidae and Coenagrionidae, are more abundant and both are diverse families at a \n250 continental scale which, which may explain their wide distribution [53–54]. Some species of \n251 Coenagrionidae are associated to degraded environments, within this family E. metallica (associated with \n252 streams with environmental change Faria et al. 2021) and N. luzmarinas uch as E. fusca, and O. walkeri \n.CC-BY 4.0 International licenseperpetuity. It is made available under a \npreprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in \nThe copyright holder for thisthis version posted September 16, 2024. ; https://doi.org/10.1101/2024.09.14.613059doi: bioRxiv preprint \n\n11\n253 [55–56].  In addition, other families, M. cupraea, H. silvarum and M. aenea also presented greater niche \n254 breadth. Species with lower abundance and smaller niche breadth tend to be sensitive to environmental \n255 changes, which make them less likely to persist when the physical characteristics of streams change [29]. \n256 The analysis shows that Odonata and the suborders separately, species with greater niche breadth \n257 had a greater contribution to beta diversity, which is opposite to expected [6,50]. Possibly more habitat \n258 generalist species and adults that are active dispersers and can survive in a wide range of environmental \n259 conditions play an important role in governing SCBD. Previous study show that the pattern of beta diversity \n260 of an assemblage is also adequately described by the common species, according to [58]. Only for Odonata \n261 and Zygoptera we found niche position negatively associated with abundance e and com Odonata SCBD, \n262 indicating that species, when very abundant, occur in non-marginal environments and occurs in more \n263 common habitats in the region. However, we highlight here that species such as C. rutilans and M. aenea \n264 (both contribute to beta diversity and have high niche breadth) in Amazonian streams seem to have limits \n265 to occur in different intensities of multiple impacts, as these species can disappear with impacts that lead to \n266 drastic changes in the streams (Faria et al. 2021). Contrary to our hypothesis, niche characteristics appear \n267 more important for contribution of these species to beta diversity than body temperature. On the other hand, \n268 for species abundance, thorax temperature is an important predictor, only when evaluated suborders \n269 separately. In fact, much previous work has demonstrated how environmental filters affect Odonata as a \n270 whole as microhabitat structure (Bank angle, woo in the stream bed), physicals and chemical, and canopy \n271 cover dossel (Brito et al., 2021) and regional variables surroundings streams (Luke et al. 2017). \n272  Ecofisiological traits as body size and thorax temperature, therefore, provides additional insights into the \n273 patterns of suborders abundance and contribute to metacommunity dynamics due to its thermoregulatory \n274 restrictions [18]. There is negative relationship between thoracic temperature and abundance. In general, \n275 Anisoptera can heat their bodies through the heliothermic or even endothermic ability. These \n276 thermoregulatory abilities are crucial for Odonata distribution [18]. Furthermore, heliothermic anisopterans \n277 can benefit from habitats with reduced riparian vegetation and greater sunlight input and can be quite \n278 abundant in these areas [21]. Conversely, larger species of Anisoptera (e.g., Gomphidae) with higher body \n279 temperature may be endothermic. These species live inside forests and are often found in streams only when \n280 they arrive for mating and oviposition [59]. These ecological and behavioral traits, make this species with \n281 higher thoracic temperatures are more difficult to collect, at the same time more sensitive to loss of riparian \n.CC-BY 4.0 International licenseperpetuity. It is made available under a \npreprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in \nThe copyright holder for thisthis version posted September 16, 2024. ; https://doi.org/10.1101/2024.09.14.613059doi: bioRxiv preprint \n\n12\n282 vegetation. Zygoptera with higher temperatures are large individuals such as H. auripennis and A. dives \n283 and had lower abundance.\n284 Thoracic temperature differs between suborders and is related to thorax width and air \n285 temperature. Anisopterans presented on average thoracic temperature up to 8ºC above air temperature and \n286 6ºC bellow, showing a negative relationship with thorax width. For Zygoptera, the thorax temperature is \n287 maintained 3ºC above and  4ºC  below air temperature. Differences in the behavior of these groups may \n288 help explain this pattern. Anisoptera tend to be heliothermic or endothermic fliers and maintain the thoracic \n289 temperature above that of the air. One of the ways to control heat loss is to alter the circulation of \n290 hemolymph between the thorax and abdomen [15], posture adjustment (Corbet, 1999). Zygoptera can \n291 maintain the temperature of the thorax closer to the air, previous studies demonstrate that this difference \n292 can be up to 1ºC (Shelly, 1982). They can be conformers or in some cases and the largest being heliothermic, \n293 or even a continuum that may exist between these groups (Corbet and May, 2018) and future studies can \n294 better investigate these categories. \n295 Finally, niche characteristics may be important for the distribution of Odonata. ecophysiological \n296 traits also was important for Anisoptera and Zygpoptera abundance. May [15] suggests also that climate, \n297 body size and behavior are essential for maintaining the body temperature of Odonata. Changes in streams \n298 due of anthropic activities alter microclimatic patterns such as air temperature, fundamental for the \n299 physiological processes of Odonata species, leading to a change in the composition of species in these \n300 environments [38]. We have demonstrated that adult odonate species composition varies in relation to \n301 habitat integrity. We therefore suggest that their monitoring would provide a good indicator of riparian zone \n302 quality considering niche characteristics and their thermorregulation habilities.\n303\n304 Acknowledgments\n305 We thank Ana Luisa Fares, Ana Luiza Andrade, Erlane José Cunha, Fernando Geraldo de Carvalho, for \n306 helping us with the biological sampling. \n307 References\n308 1. Siqueira T, Bini LM, Cianciaruso MV, Roque FO, Trivinho-Strixino S. The role of niche measures in \n309 explaining the abundance–distribution relationship in tropical lotic chironomids. Hydrobiologia. 2009; 636: \n310 163. https://doi.org/10.1007/s10750-009-9945-z\n.CC-BY 4.0 International licenseperpetuity. 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