Congruence of detection probabilities and co-occurrence of threatened Afromontane damselflies with diverging functional traits (Odonata: Chlorocyphidae, Coenagrionidae)

Congruence of detection probabilities and co-occurrence of threatened Afromontane damselflies with diverging functional traits (Odonata: Chlorocyphidae, Coenagrionidae) | 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 Congruence of detection probabilities and co-occurrence of threatened Afromontane damselflies with diverging functional traits (Odonata: Chlorocyphidae, Coenagrionidae) Anthony Karani, Anne Kairu, Michael Githaiga, Andrew Gichira, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6433576/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 05 Feb, 2026 Read the published version in Journal of Insect Conservation → Version 1 posted 9 You are reading this latest preprint version Abstract The biodiversity crisis is exceptionally severe in the freshwater systems of the highly threatened Afromontane ecosystems. We studied functional traits of ecological significance, estimated abundance, probabilities of occupancy, and detection of adults and nymphs of Kenya Jewel ( Platycypha amboniensis Martin, 1915) and Giant Sprite ( Pseudagrion bicoerulans Martin, 1907) in Mount Kenya Forest. The estimates of abundance were based on replicated counts, while occupancy and detection probabilities were estimated using a single-season, two-species occupancy formulation. The study revealed that detection probabilities of one species were influenced by the detection probabilities of the other, but conversely, occupancy was not. This was supported by morphological traits. The larvae of P. amboniensis is adapted to rocky and fast-flowing lotic streams, while P. bicoerulans is adapted to vegetated, littoral, and slow-moving reaches of the same streams. The study concluded that while these species have different adaptations, their co-occurrence depends on the heterogeneity of the microhabitats and recommended ecosystem restoration approaches that maintain habitat complexity to increase the resilience of co-occurring species to future environmental changes. Implications for conservation : The occurrence of Kenya Jewel was not influenced by occurrence of Giant Sprite and; there was neither competition nor niche overlap. This aligns with the differences in ecological adaptations based on functional morphology of adults and larvae. This evidence is useful for restoration of the ecosystem to ensure that these species are conserved concurrently in their range of co-occurrence. occupancy probability co-occurrence larval morphology functional traits conservation ecosystem restoration Figures Figure 1 Figure 2 Figure 3 Introduction Insects provide a continuum of ecosystem services that are critical for human welfare and survival (Samways et al. 2020 ). These services include food provisioning, nutrient cycling and decomposition of organic matter, pollination, control of pests, and use in ecological monitoring, among others (Samways et al. 2020 ; May 2019 ). Lately, there has been a dramatic decline in insect populations and diversity around the globe, which has inspired multiple conservation actions (Forister et al. 2023 ; Hallmann et al. 2017 ). These actions are urgent because population dynamics of insects have the potential to easily translate to instability of terrestrial ecosystems (May 2019 ). Conservation of threatened insects is shifting from identification of causes of population decline to searching for mechanisms to address effects and understanding the intrinsic responses of the populations to ecosystem restoration (Webster et al. 2023 ; Lambert et al. 2023). This information is required to inform current conservation endeavours and model future population recovery under extreme scenarios and across vast landscapes (Kawahara et al. 2021 ). However, this effort is limited by the paucity of reliable data on the functional traits of species, population abundance and species interactions within microhabitats (Viza et al. 2023 ; Khelifa et al. 2021 ). Exploring this diversity is particularly problematic for aquatic insects because they have complex life history traits that involve metamorphosis and the use of different habitats for their larval and adult stages (Kietzka et al. 2021 ; Stoks and Córdoba-Aguilar 2012 ). The causes of the population decline of aquatic insects is rooted in different developmental stages, resulting in an observed difference in sensitivity and vulnerability of the species (Samways 2008 ). The highest diversity of threatened dragonflies and damselflies ( Odonata ) in Africa is concentrated in the tropical forests and highlands (Clausnitzer et al. 2012 ). Habitat loss in the afromontane radiation due to deforestation impacts both the terrestrial and aquatic phases of odonate species (Samways 2008 ; Clausnitzer et al. 2011 ; Clausnitzer et al. 2012 ; Hořák et al. 2023 ). Damselflies ( Zygoptera ) and dragonflies ( Anisoptera ), being sensitive to different stressors, amphibious in nature, and charismatic players in the food web as top predators of aquatic and terrestrial invertebrates and prey for vertebrates, are potent and powerful ecological indicators (May 2019 ). The knowledge of their ecology and diversity is crucial to conservation biology (Dijkstra et al. 2014 ). Much of the current work on odonate populations in East Africa has focused on biogeography and taxonomy (Njoroge et al. 2017; Clausnitzer et al. 2011 ). These include the characterisation of extinction risk based on trends in population abundance, geographical distribution, and habitat fragmentation (Clausnitzer et al. 2012 ). Species-specific functional traits of larvae and adult damselflies that can contribute to population resilience of threatened species in subtle ecological settings remain unknown. Understanding the abundances of species is important for their conservation, whether threatened or not threatened (Hogreve and Suhling 2022 ; Baker et al. 2019 ). Understanding the detection probabilities of species is important to help scientists and local communities in awareness creation, research, planning, and ecological restoration (Redford et al. 2013 ). Equally, knowledge on species co-occurrence can be used to inform how communities function, such as revealing patterns in competition and niche overlap (Steenweg et al. 2019 ). The aim of this study was to estimate the abundance and probabilities of co-occurrence of Giant Sprite, Pseudagrion bicoerulans Martin, 1907 (Coenagionidae), and Kenya Montane Jewel, Platycypha amboniensis Martin, 1915 (Chlorocyphidae), in Mount Kenya Forest. The study also aimed at elucidating functional traits of ecological significance based on the morphology of the adults and final stadium instar larvae of the two damselfly species. Materials and methods Study species Two threat-defined and range-restricted species of damselflies (Fig. 1 ) were studied. Platycypha amboniensis is endemic to the Central Highlands of Kenya and is highly dependent on montane forest and considered a sub-montane relict species among dancing jewels (Clausnitzer et al. 2011 ). Little is known about its reproductive ecology, behaviour, and life history, even though it is a very rare and highly threatened species (Dijkstra and Clausnitzer 2014 ). The species is classified as Critically Endangered (CR) due to extensive habitat loss as a result of forest clearance within regions it is dependent upon (Clausnitzer et al. 2011 ). The male of this species has bright orange tibiae that are slightly flattened and white interiorly, sky-blue abdominal segments (S5–S10), and large, bulbous eyes (Dijkstra and Clausnitzer 2014 ). Similarly, P. bicoerulans is endemic to streams in the mountains of Kenya, Tanzania, and Uganda. The species is classified as Vulnerable (VU) in the IUCN Red List. It was termed ‘the most alpine of African Odonata’ because it occurs in heather and afroalpine zones above 3000 m above sea level (Clausnitzer et al. 2011 ). The species has high intraspecific genetic diversity and three subspecies: P. b. bicoerulans (Central Kenya), P. b. elgonensi (West Kenya and East Uganda), and P. b. kilimanjaricus (North Tanzania), widespread across high-elevation forests in East Africa (Dijkstra et al. 2007 ). Males of this species are distinguished by a bright yellow to orange labrum and small orange post-ocular spots, a green-striped black thorax, blue or pruinescent abdominal segments (S1–S2, S9–S10), and clasper-like cerci (Dijkstra and Clausnitzer 2014 ). Study area The study was conducted in Kangaita Forest Station (Kirinyaga County) and Irangi Forest Station (Embu County) (Fig. 2 ). These forest stations are part of the Mount Kenya Forest Reserve, which is a protected area in the Central Highlands of Kenya (Nyongesa and Vacik 2019 ). This study area is classified as lower montane humid forest and is renowned for its agro-ecological value and high diversity of woody plant species, birds, and mammals (Onditi et al. 2022 ; Zhou et al. 2018 ). The drainage pattern is dendritic, with all streams forming tributaries of the Tana River. Water is directly abstracted in large quantities from these streams for domestic use, municipal supply, microhydropower stations, and small- and large-scale irrigation schemes. Sampling The sampling period ranged from September 2022 to March 2023 and is considered in this study a 'single season', which runs across two dry periods interspaced with a period of short rains. It is possible that the adults and final instar larvae encountered during this study belong to the same generation. Sampling sites were selected randomly from a list of first-level and second-level streams (five in Kangaita and five in Irangi). At each sampling point, three transects of 100 m in length and covering the width of the stream channel (up to 10 m) were established along each stream following the method of Batista et al. ( 2021 ). Sampling of adults, exuviae, and larvae was undertaken at four sampling occasions (interval: 9–17 days) along each transect at twenty-meter intervals, starting at zero, which translates to five full-width belt transects (FWBT) on each transect (Darshetkar, Patwardhan, and Koporde 2023), equivalent to 200 m 2 . The following water quality parameters were measured using a multi-parameter portable meter (Suzhou Holi, China): Dissolved Oxygen (DO), Electrical Conductivity (EC), Oxidation-reduction potential (ORP), pH, total dissolved solids (TDS), and temperature. Turbidity was measured using a nephelometer (BD PhoenixSpec™, South Africa). Sampling was consistent with the guidelines for handling living insects (Montegomery et al. 2021; Cezário et al. 2021 ). On each sampling occasion, two observers were involved and took turns in data entry and observation. All adult damselflies, exuviae, and nymphs encountered at each interval were counted. Evidence of the presence of adults was confirmed by the presence of damselflies perching on the rocks, twigs overhanging the stream, emergent vegetation, and driftwood. The sampling period was mostly sunny, and the water was shallow. No attempts were made to search for adult damselflies beyond the riverbanks. Exuviae were found by locating suitable emergence sites mostly on emergent vegetation, driftwood, and exposed rocks. Collected exuviae were placed in Falcon tubes and stored in a freezer in order to prevent damage by inadvertently trapped insects and spiders and the growth of mould. After at least 12 hours of freezing, the exuviae were dried and mounted. Larvae were identified by sinking a dip net into the water and scooping debris at the edges of the stream (mostly giant sprite). Movable rocks along the river were also lifted or overturned and checked for clinging or crawling damselfly larvae (mostly montane jewel). These were then returned to position at the point of collection. On account of the rarity of the two species, only a few larvae and adults were collected. Adults were collected with a professional sweep net (38 cm diameter, 86 cm depth, and 100 cm handle). Specimens were placed in specimen jars and washed in acetone or preserved in absolute ethanol. The following parameters were measured for adult damselflies: abdomen length, body length, hind femur length, thorax length, wing area, wing length, wing perimeter, and wing width. The following parameters were measured for nymphs: abdomen length, appendages, body length, gonapophyses, hind femur length, prementum, and wing sheath. Data analyses The pairwise correlation between water properties was calculated using Pearson corrrelation coefficients. The difference in functional traits between males and females was estimated using an independent samples t-test. The replicated count data (pooled to stream level and treated separately for adults and nymphs of both species) were averaged as daily abundance with standard deviation (mean ± sd). A ratio was calculated by dividing the mean daily abundance of nymphs by that of adults. An independent sample t-test was used to determine if there were significant differences in means. These tests were carried out in R versions x64: 4.0.3 (R Core Team 2020 ). Species abundance data denoted as presence-absence at each sampling interval and organised into pairs ( P. amboniensis adult + P. bicoerulans adult; P. amboniensis nymph + P. bicoerulans nymph) to fit the formulation of two-species single-season occupancy models (Richmond et al. 2010 ). This model accepts detection histories of each species at each site across sampling occasions (0 indicating non-detection and 1 indicating detection). The model assumes that one of the species in pairwise interactions is dominant while the other is subordinate (Kleiven et al. 2023 ; Lonsinger 2022 ). Three parameters were used to estimate occupancy: ψA—occupancy probability of species A ( P. amboniensis ); ψBA—occupancy probability of species B ( P. bicoerulans ) in the presence of species A; and ψBa—occupancy probability of species B in the absence of species A. These were used to test the hypothesis that the occupancy probability of P. amboniensis is not dependent on the presence of P.bicoerulans (ψBA ≠ ψBa). Five parameters were estimated for detection probability, namely: pA—detection probability of species A in the absence of species B; pB—detection probability of species B in the absence of species A; rA—detection probability of species A in the presence of species B; rBA—detection probability of species B when species A is present and detected; rBa—detection probability of species B when species A is present but not detected. These were used to test hypotheses that 1) the detection probability of P. amboniensis is not dependent on the presence of P. bicoerulans (pA ≠ rA) and 2) the detection probability of P. bicoerulans is not dependent on the presence and detection of P. amboniensis (pB ≠ rBA). The model also estimates derived parameters of occupancy probability: ψB—probability of occupancy for species B, regardless of occupancy status of species A; ψAB—probability of both species being present. The species interaction factor (SIF or γ) was used to measure co-occurrence, where γ = 1 means that the two species occur independently of each other, γ 1 means that there is overlap in the co-occurrence of the two species (Lonsinger 2022 ) A logit link function was used to model the occupancy and detection of both species in the program MARK (White and Burnham 1999 ). This was conducted separately for pairs of adults and nymphs. A total of 32 predetermined models were developed to include combinations of constant occupancy probability ψ(.), variable occupancy probability ψ(t), constant detection probability p(.), and variable detection probability p(t). All candidate models were ranked from highest to lowest based on their Akaike’s Information Criteria modified for c-hat (QAICc) values and the AIC weights. The top-ranked model has a model likelihood of one (1), the lowest QAICc, and presents the best combination of parameters (ψA, ψBA, ψBa, rA, pA, pB, rBa, rBA). Results The physico-chemical water properties for each stream are shown in Table 1 . There was no significant correlation between dissolved oxygen (DO) and oxidation-reduction potential (ORP) and no correlation between turbidity and pH. In contrast, EC was strongly positively correlated with TDS (r = 0.93) and negatively correlated with ORP (r = -0.78). Temperature was negatively correlated with pH (r = -0.72), while OPR was negatively correlated with TDS (-0.79). Several moderate and weak relationships were observed, such as the inverse between temperature and DO. Table 1 Mean and range of water quality parameters measured from forest streams in Kangaita and Irangi Forests, Mount Kenya STREAM DO EC ORP PH TDS TEMPERATURE TURBIDITY Ciaminogia 3.9 (3.9–3.9) 36 (32–42) 183 (159–208) 7.837 (7.68–7.93) 18.33 (17–21) 18 (16.7–18.9) 8.34 (3.43–15.53) Rwamuthambi 4.18 (3.6–4.6) 26.5 (26–27) 270.5 (247–294) 7.875 (7.86–7.89) 13 (13–13) 14.6 (13.9–15.3) 4.22 (2.64–6.52) Mukengeria 3.9 (3.8–4.2) 15.3 (14–16) 334 (153–585) 7.99 (7.75–8.14) 7.33 (7–8) 15.167 (14.1–16.5) 7.39 (2.77–15.63) Ruiru 4.4 (4.2–4.6) 38.6 (34–42) 189 (163–212) 7.32 (7.11–7.56) 15.25 (14–17) 14.52 (13.7–16.3) 11.38 (2.92–16.06) Rundu 4.56 (4.1–4.9) 50.4 (46–68) 215.2 (135–381) 7.67 (6.61–8.26) 25.6 (23–35) 19.02 (17.1–22.1) 4.02 (0.65–8.35) Ena 5.56 (5.2–5.9) 40.75 (23–64) 232.5 (204–273) 7.92 (7.22–8.53) 20.75 (13–32) 13.15 (12.2–14.6) 4.69 (3.1–6.9) Karutune 4.65 (4.6–4.7) 42.25 (36–46) 174.6 (108–248) 7.94 (7.14–8.65) 21.6 (17–24) 15.85 (13.9–18.3) 4.317 (2.27–6.2) Kiye 5.05 (4.4–5.7) 37.8(16–47) 185.8 (163–201) 7.91 (7.21–8.4) 24.8(12–48) 17.92 (12.9–22.8) 5.07 (3.72–6.52) Nyanjara 6.25 (6.2–6.3) 34.5(18–48) 265.75(208–373) 8.11 (7.75–8.37) 21 (17–23) 13.2 (12.5–13.9) 4.55 (3.4–6.5) Thambana 3.95 (3.6–4.3) 38.6 (32–45) 199 (124–257) 7.95 (7.43–8.64) 21.2 (18–26) 12.56 (10.5–14.3) 3.73 (2.39–5.6) Note : Abbreviations; DO: Dissolved Oxygen, EC: Electrical Conductivity, ORP: Oxidation-Reduction Potential, pH: Acidity/Alkalinity, TDS: Total Dissolved Solid Table 1 There was a significant difference between males and females of P. amboniensis for abdomen length, body length, wing area, and wing length. No significant differences were found for hind femur length, thorax length, wing perimeter, and wing width (Table 2 ). There was no evidence that the adult males and females of P. bicoerulans differed significantly. The male and female nymphs of both P. bicoerulans and P. amboniensis did not differ in abdomen length, appendages, body length, hind femur length, prementum, and wing sheath at emergence. There was a significant difference in the size of the gonapophyses of P. bicoerulans and the prementum of P. amboniensis. Table 2 Morphological traits of adults and nymphs of P. amboniensis and P. bicoerulans in Mount Kenya Trait Female Male Ttest df p-value Female Male Ttest df p-value Adult : Platycypha amboniensis Adult : Pseudagrion bicoerulans Abdomen length 17.8 (17.5–18.2) 19.95 (18.9–21.2) -8.09 11.95 > .001* 33.4 (32.2–35.7) 34.36 (32.3–36) -0.88 6.47 0.411 Body length 29.7 (28.2–30.8) 33.32 (32.1–35.1) -5.67 4.70 0.003* 42.7 (40.9–47) 43.54 (41.7–46.2) -0.51 4.80 0.634 Hind femur length 5.58 (4.9–6.6) 5.49 (4.9–6.2) 0.21 3.31 0.847 4.88 (4.3–5.4) 5(4.3–5.3) -0.40 5.63 0.704 Thorax length 8.55 (7.7–9.4) 8.79 (7.9–10.1) -0.52 3.82 0.629 6.85 (6.6–7.3) 6.48 (5.6–7.3) 1.13 6.25 0.299 Wing area 100.94 (87.28-120.32) 84.37 (68.91–93.68) 4.78 11.77 > .001* 98.87 (85.37-105.89) 101.68 (74.81-127.06) -1.37 39.20 0.178 Wing length 27.83 (25.73–30.34) 25.81 (24.64–27.24) 3.47 24.86 0.002* 26.73 (25.7–27.5) 26.36 (25.4–27.4) -0.82 29.00 0.418 Wing perimeter 80.96 (71.96–90.35) 72.27 (62.01-127.43) 1.97 31.46 0.058 82.15 (61.85-139.56) 84.99 (61.34-145.48) -0.35 25.39 0.726 Wing width 5.76 (5.19–6.36) 5.49 (4.69–9.25) 1.18 3.11 0.32 6.11 (5.36–6.65) 6.16 (5.15–8.24) 0.64 6.86 0.541 Nymph : Platycypha amboniensis Nymph : Pseudagrion bicoerulans Abdomen length 10.43 (9.2–11.2) 10.23 (9.6–10.7) 0.35 4.96 0.741 12.27 (12.1–12.5) 12.27 (11.7–12.6) 0.00 2.69 1 Appendages 5.73 (5.3–6.2) 5.77 (5.4–6.2) -0.13 4.69 0.902 6.77 (6.4-7) 6.67 (6.1-7) 0.29 3.44 0.786 Body length 22.63 (22.1–23.5) 22.3 (22.1–22.7) 0.91 4.79 0.407 25.47 (25.1–26) 25.5 (25.1–25.9) -0.09 3.89 0.93 Gonapophyses 1.98 (1.8–2.1) - - - - 1.97 (1.9–2.1) 0.87 (0.8-1) 11.67 4.00 > .001* Hind femur length 5.85 (5.5–6.6) 5.67(5.3-6) 0.57 5.00 0.596 5.3 (5.2–5.5) 5.4 (5.3–5.6) -0.71 4.00 0.519 Prementum 3.63 (3.6–3.7) 3.8 (3.8–3.8) -7.00 3.00 0.006* 3.4 (3.3–3.5) 3.67 (3.3-4) -1.26 2.32 0.318 Wing Sheath 5.5 (5.4–5.7) 5.57 (5.4–5.8) -0.48 3.36 0.662 6.27 (6.1–6.6) 6.4 (6.2–6.7) -0.59 3.97 0.587 The abundance of both adults and nymphs differed between species in all sampling sites (t = -3.99, df = 33.76, p-value < 0.05). The damselflies, P. amboniensis and P. bicoerulans , were encountered in the study area a total of 3605 times. These included 871 adults and 613 nymphs of P. amboniensis and 906 adults and 1215 nymphs of P. bicoerulans . Additionally, the ratio of adults and nymphs at sampling sites differed between species (t = 3.68, df = 40.4, p-value < 0.05). The average number of individuals encountered across sampling occasions was (mean ± sd) 19.85 ± 8.96, 16.70 ± 8.47, 22.00 ± 10.22, and 33.20 ± 13.35 for P. amboniensis adults, P. amboniensis nymphs, P. bicoerulans adults, and P. bicoerulans nymphs, respectively, in Irangi Forest; 23.70 ± 9.43, 13.95 ± 3.55, 23.30 ± 11.81, and 27.55 ± 8.31 for P. amboniensis adults, P. amboniensis nymphs, P. bicoerulans adults, and P. bicoerulans nymphs, respectively, in Kangaita Forest (Fig. 3 ). This translates to a density of 0.218 sqm⁻¹, 0.153 sqm⁻¹, 0.227 sqm⁻¹, and 0.308 sqm⁻¹ of P. amboniensis adults and P. amboniensis nymphs, P. bicoerulans adults, and P. bicoerulans nymphs, respectively. Table 2 The most parsimonious models (Table 3 ) estimated the probability of occupancy for combinations of P. amboniensis adults— P. bicoerulans adults and P. amboniensis nymphs— P. bicoerulans nymphs but showed no evidence that occupancy of either was influenced by occupancy of the other (Table 4 ). The best-supported model for adult occurrence had an AICc value of 645.11, a weight of 0.356, and the model likelihood reduced dramatically to 0.058 in the sixth model (ΔAICc = 5.703). The best-supported model for nymph co-occurrence had an AICc value of 603.84, a weight of 0.333, and the model likelihood reduced to 0.086 in the sixth model (ΔAICc = 4.907). Table 3 Candidate models used to evaluate co-occurrence and detection probability of Platycypha amboniensis Martin, 1915 and Pseudagrion bicoerulans Martin, 1907, using two-species conditional occupancy model Occupancy Detection AICc ΔAICc Weight Likelihood Num. Par Co-occurrence between nymphs of P. amboniensis and P. bicoerulans ψA(.) ψBA(.) ψBa(.) pA(.) pB(.) rA(.) rBA(.) rBa(t) 645.11 0 0.356 1 9 ψA(.) ψBA(.) ψBa(t) pA(.) pB(.) rA(.) rBA(.) rBa(.) 645.61 0.504 0.276 0.777 8 ψA(.) ψBA(t) ψBa(.) pA(.) pB(.) rA(t) rBA(.) rBa(.) 647.44 2.331 0.111 0.312 9 ψA(t) ψBA(.) ψBa(.) pA(.) pB(.) rA(.) rBA(t) rBa(.) 647.67 2.562 0.099 0.278 9 ψA(.) ψBA(t) ψBa(t) pA(.) pB(t) rA(.) rBA(.) rBa(.) 647.75 2.640 0.095 0.267 10 ψA(t) ψBA(t) ψBa(.) pA(t) pB(.) rA(.) rBA(.) rBa(.) 650.81 5.703 0.020 0.058 11 Co-occurrence between adults of P. amboniensis and P. bicoerulans ψA(.) ψBA(.) ψBa(.) pA(.) pB(.) rA(.) rBA(.) rBa(t) 603.84 0 0.333 1 7 ψA(.) ψBA(.) ψBa(t) pA(.) pB(.) rA(.) rBA(.) rBa(.) 604.19 0.357 0.278 0.837 5 ψA(.) ψBA(t) ψBa(.) pA(.) pB(.) rA(t) rBA(.) rBa(.) 606.29 2.455 0.097 0.298 8 ψA(t) ψBA(.) ψBa(.) pA(.) pB(.) rA(.) rBA(t) rBa(.) 606.43 2.592 0.091 0.274 9 ψA(.) ψBA(t) ψBa(t) pA(.) pB(t) rA(.) rBA(.) rBa(.) 608.58 4.739 0.031 0.094 10 ψA(t) ψBA(t) ψBa(.) pA(t) pB(.) rA(.) rBA(.) rBa(.) 608.72 4.907 0.029 0.086 8 Table 4 Estimates of occupancy (ψ), detection probability (p and r), and Species interaction factor (SIF or γ) for Platycypha amboniensis Martin, 1915 and Pseudagrion bicoerulans Martin, 1907, determined using two-species conditional occupancy model with data from Irangi and Kangaita Forests in Mount Kenya Nymphs Adults Co-occurrence Estimate SE Range Estimate SE Range ψA 0.995 0.049 (0.884–1.000) 0.915 0.061 (0.695–0.981) ψBA 0.969 0.060 (0.377–0.999) 0.971 0.032 (0.787–0.997) ψBa 1.000 0.000 (1.000–1.000) 1.000 0.000 (1.000–1.000) pA 0.473 0.390 (0.040–0.951) 0.625 0.122 (0.256–0.744) pB 0.140 0.589 (0.000–1.000) 0.504 0.497 (0.252–0.966) rA 0.334 0.036 (0.268–0.408) 0.338 0.065 (0.225–0.478) rBA 0.429 0.117 (0.227–0.657) 0.328 0.090 (0.181–0.524) rBa 0.401 0.048 (0.311–0.499) 0.308 0.076 (0.183–0.473) γ 0.999 0.002 (0.995–1.002) 0.997 0.002 (0.998–1.001) ψB 0.960 0.042 (0.736–0.995) 0.973 0.018 (0.922–0.993) ψAB 0.926 0.044 (0.781–0.978) 0.888 0.043 (0.853–0.984) Table 3 There was no evidence that occupancy of P. amboniensis was affected by the presence of P. bicoerulans (Table 4 ). The species had high probabilities of occupancy (0.915–1.00) regardless of whether the other species was present or not present (ψA = ψBA = ψBa = ψB ≅ 1.00); the probability of both species being present was 0.926 ± 0.044 for nymphs and 0.888 ± 0.043 for adults. The two species were present and detected independently (γ = 0.999 ± 0.002). There was evidence that the presence of P. amboniensis a ffected the presence and detection probability of P. bicoerulans . The detection probability for nymphs of P. amboniensis was higher when the adults of P. bicoerulans were absent (pA = 0.473 ± 0.39). The detection probability of P. bicoerulans nymphs was lowest when the nymphs of P. amboniensis were present (pB = 0.14 ± 0.589). There was support that the detection probability of P. amboniensis adults was influenced by the detection probability (but not presence) of P. bicoerulans (rA ≠ pA). The detection probability for adults of P. amboniensis was higher when the adults of P. bicoerulans were absent (pA = 0.625 ± 0.122). Table 4 Discussion The study species occur in running water but display clear differences in habitat requirements. The differences and similarities in the morphology of P. amboniensis and P. bicoerulans larvae that are related to ecological adaptations are summarised in Table 5 . These species belong in distinct lineages that have followed different evolutionary trajectories for an extended period (Dijkstra et al. 2014 ). This study contributes towards understanding the patterns of co-occurrence and structure of Afromontane damselfly communities as mediated by species-specific adaptations to varying microhabitats. As expected, P. b. bicoerulans ( sensu stricto ) occur in the same streams occupied by P. amboniensis in the Central Highlands of Kenya. The study showed neither competition nor niche overlap in the occupancy of P. amboniensis adults— P. b. bicoerulans adults as well as P. amboniensis nymphs—and P. bicoerulans nymphs. It is also noteworthy that the method assumes population closure ( sensu Richmond et al. 2010 ), which is limited when associated with open populations (Kleiven et al. 2023 ; Lonsinger 2022 ). Table 5 Similarities and differences in larval morphology of Platycypha amboniensis Martin 1915 and Pseudagrion bicoerulans Martin 1907 and functional traits that account for ecological adaptations Morphology Functional traits Ecological adaptations P. amboniensis General habitus Body is short, stout, cylindrical, and setose; muddy brown to dark brown; legs are long Nymphs sit and wait to ambush prey; their stout body and long legs are suitable for clinging on stones and gravel substrates; setae attract mud or debris for camouflage in dark/brown environments Mouthparts Labium brown without setae, labial palps with long hooks, mandibles setose and heavily sclerotized, hypopharynx well developed Labium colour blends with the environment, hooks for grasping prey, and strong mouthparts for processing food with hard cuticles Caudal lamellae Epiproct rudimentary; paraprocts elongate, triquetral, setose and spinose Long lamellae with spines used for respiration in benthic region with lower dissolved oxygen; the shape enhances balance in current waters and autotomy to escape predators Gonapophyses Well developed in female, marginal in male Fast development of nymphs P. bicoerulans General habitus Body is elongate, cylindrical, pale or dark brown; legs are short Nymphs actively search for prey; their colour blends with litter and vegetation in littoral zone Mouthparts Labium long, pale to translucent, with premental/labial setae; labial palps wide Pale colour of labium for ‘invisibility’ while ambushing prey, wide labial palps for better clutching of larger prey, setae enhance sensitivity Caudal lamellae Epiproct and paraprocts are elongate, leaf-like, flattened, and pigmented Lamella used for swimming, respiration among vegetation, agonistic display among conspecifics, and autotomy to escape predators Gonapophyses Well developed in male and female Quick maturation of adults Table 5 Functional traits determine the distribution and structure of Odonata assemblages in streams (Silva et al. 2021 ). The evolution of morphological traits in Odonata larvae is associated with requirements for larval survival, such as breathing, feeding, and predation refuge. The independent co-occurrence of P. amboniensis and P. bicoerulans can be explained in terms of their feeding ecology as a trade-off for 1) different prey items in littoral zone and fast-flowing sections and 2) specialised morphological features that aid processing of these prey items. The mouthparts of P. amboniensis and P. bicoerulans differ in shape, size, and structure; the significance is that it enhances foraging success in varying microhabitats. The general morphology of larval mouthparts in Odonata is grossly similar, meaning that these small morphological variations are insightful to explain ecological adaptations with regards to food intake in littoral and benthic zones (Büsse et al. 2021 ; Büsse and Gorb 2018 ). The presence of labial palps with long hooks in P. amboniensis is equipped for better grasping of prey, while the prementum with wide labial palps that have setae in P. bicoerulans increases the sensitivity of the prehensile mask. In addition, the presence of a well-developed hypopharynx enhances overall food acquisition in P. amboniensis , as its muscles have been associated with the opening of the labium (Büsse and Gorb 2018 ). The littoral and benthic regions of lotic streams differ in their mechanisms of retaining dissolved oxygen. Stream sections with rapids and stones are cooler and abundant in dissolved solids, while littoral sections are vegetated and warmer. Caudal lamellae with spines allow the larvae to survive through active gaseous exchange as well as predator avoidance. On the other hand, increased sedimentation and substrate homogenisation favour larvae that cling, swim, or climb on substrates (Mendes et al. 2017 ). Predator avoidance can account for disparities in microhabitat selection, resulting in trait compensation in morphological and behavioural defences in damselflies (Pernecker et al. 2020 ). Damselflies have evolved autotomy, where caudal lamellae are self-amputated to escape predators (Black et al. 2019 ). The drab colour and detritus particles attaching to the setae of P. amboniensis make it well camouflaged, while the swimming capabilities of P. bicoerulans make it easy to escape predators. A robust and cylindrical body shape ( P. amboniensis ) is preferred in microhabitats with a higher proportion of stones and gravel substrate (Pires et al. 2020 ); that is also utilised by burrowing predators such as gomphids (mostly Notogomphus spp. and Paragomphus spp.). On the other hand, an elongate body ( P. bicoerulans ) is suited for refuge among vegetation and litter, where climbing and sprawling predators live ( mostly, Pinheyschna, Zygonyx , and Atoconeura ). Siltation of the streams leads to sedimentation that homogenises substrates, excluding larvae that depend on substrates in favour of burrowing species (Mühlenhaupt et al. 2022 ; Dalzochio et al. 2018 ). Larval development, though rarely explored, can augment sexual maturation and influence the phenology of adult reproduction. While it was beyond the scope of this study to examine the development cycle of these species, the observed differences in gonapophyses of P. amboniensis and P. bicoerulans at emergence can be described as a trade-off for 1) faster development of nymphs and 2) quick maturation of the emergent adults. These species have endophytic oviposition, where females deposit their eggs within plant tissues by means of a well-developed ovipositor. It is also noteworthy that larval behaviour and adult behaviour in these damselflies are intertwined, where, for instance, the site of oviposition and habitat for early larval development is chosen by the adult. This compensates for higher activity, which results in a higher growth rate but increases predation risk in damselfly larvae (Mühlenhaupt et al. 2022 ) and predicts developmental plasticity in changing environments such as drying pools (Pernecker et al. 2020 ). This can be used to explain why adult detection probability differed from that of nymphs within a shared habitat. The co-occurrence of P. amboniensis and P. bicoerulans can be explained by differences in microhabitat use and behaviour as predicted by functional larval morphological traits. Co-occurring species exhibit a wealth of differing traits that allow them to partition the resources and reduce competition. Alteration of habitat has consequences on abundance (Dalzochio, Périco, Dametto, and Sahlén 2020 ), and size structures of multiple co-occurring species (Oliviera-Junior, Teodósio, and Juen 2021). This affects species simultaneously and sometimes shifts in favour of species that have evolved in heterogeneous environments. The naiads of P. bicoerulans were detected mostly on sites with slower water flows and higher abundances of emergent vegetation characterised by warm waters and low abundances of total dissolved solids. This microhabitat specialisation is probably the reason why P. bicoerulans is easily displaced by competitive lowland species (Clausnitzer et al. 2011 ). Nonetheless, the overall detection probability of naiads was lower than that of adults in the presence of P. amboniensis . The adults of P. amboniensis were high in areas with driftwood, while naiads were detected mostly where the stream was rocky and fast-flowing. This study also sheds light on stage-specific differences in the occupancy and detection probabilities of adults and larvae of co-occurring species. The sensitivity of Afromontane forests is due to their distinct vegetation types that are ecologically meaningful for the terrestrial adult Odonata (Clausnitzer et al. 2011 ; Dijkstra and Clausnitzer 2006). However, owing to the complex life history traits that involve metamorphosis and use of different habitats for larvae and adults (Hogreve and Suhling 2022 ; Steenweg et al. 2019 ), stability of these populations can be regulated at different developmental stages (Lambert et al. 2023). Determining the demographic structure of such a population leads to understanding how each stage influences the other. Adults contribute to the recruitment of nymphs through laying eggs and selection of oviposition sites that enhance survival, while nymphs transform into adults through emergence. The presence of many adults on a site of oviposition means that a large number of nymphs will be recruited, while the emergence of a large number of larvae translates into adults. There is a difference in the means of dispersal between life stages. The adults fly to better microhabitats suitable for oviposition, while naiads have the ability to swim or drift away, especially downstream. Implications for conservation It has been established that East African Montane Forests, Ethiopian Highlands, Eastern Arc, and Coastal Forest Mosaics are regions of conservation value due to the large number of threatened species in these regions (Clausnitzer et al. 2011 ; Clausnitzer 2003 ). The sensitivity and diversity of threatened species also coincide with the endemism patterns linked with habitat heterogeneity in the highlands (Dijkstra et al. 2007 ). The highly susceptible species are habitat specialists that occur in a small geographical range and have a limited area of occupancy. These include those that are confined to mountain peaks and those living in highly fragmented landscapes. Although the occupancy of P. amboniensis and P. bicoerulans in Mount Kenya is promising, these populations are separated from the closest populations located in the Aberdare Ranges the risk of continued habitat isolation means that even when a population is seemingly stable in a restricted range, it is still vulnerable to stochastic environmental events like extreme weather and genetic effects such as bottlenecks and inbreeding depression that hasten biotic attrition (Webster et al., 2023 ). Dragonflies and damselflies are agile organisms because they require more than one ecosystem to become mature adults. Changes incurred at the larval stage are reflected in the adult stage and vice versa. The exposé that these species have invested in the evolution of functional traits in larvae is timely. Moving forward, it will be necessary to define stage-specific and species-specific threats and evaluate extinction probabilities. For instance, in reality, habitat loss is a broad term in the context of odonates and other freshwater invertebrates (Forister et al. 2023 ) and fails to account for all life stages. It has been shown that even when protected areas are rarely designed with conservation of insects in mind (Chowdhury et al. 2023 ), conservation of threatened species can be achieved through conservation of targeted non-threatened species (Baker et al. 2019 ). Species in the Afromontane streams are adapted to living in clear waters with little natural silting while species that occur downstream are more tolerant of human disturbance and natural silting from storms (Deacon and Samways 2021 ). The nymphs are critical for conservation because a great proportion of Odonata larvae do not emerge or develop to adulthood, and this number may rise with increased human pressure on these habitats caused by multiple factors (Kietzka et al. 2021 ). These include soil erosion and siltation caused by deforestation and livestock grazing along riverbanks and invasive alien trees that affect freshwater systems through synergistic effects. When water conditions are good, removal of alien invasive trees can lead to the rapid recovery of dragonfly populations, as long as a source population still exists in the area (Samways 2008 ). The first- and second-level streams in Mount Kenya are being tapped heavily throughout the year for community water supply. Water abstraction affects abundances of many species groups, including fish (Merciai et al. 2017 ) and dragonflies. Dynamics of water abstraction from small montane streams may prevent threatened species (such as P. amboniensis and P. bicoerulans ) from becoming abundant and encourage generalist species. It could also result in fine partitioning of narrow-range endemic species in small Afromontane streams influenced by damming and siltation. Damming of waterways can locally impoverish riverine fauna, affect water flow, create lentic conditions in lotic systems, and incur long-term changes in odonate assemblages (Samways 2008 ). These newly created spaces appear to favour occupancy by P. bicoerulans but fail to guarantee that these microhabitats are suitable for the species long-term survival. For instance, when a dam is left to follow natural succession and becomes fully vegetated, it is likely to attract different generalist species. This reduces the abundance of specialists in favour of widespread generalists. Damming encourages colonisation of invasive predatory species such as clawed frogs ( Xenopus borealis) , which has cascading effects on various biotas and neighbouring ecosystems through impact on dragonfly larvae. Similarly, the silt that accumulates on the ponds is often released into the stream and washed downstream. The effects of these recurrent small-scale events, though barely detected in the short term, affect local species co-occurrence and can alter community structure in the long term. Conclusion This study states a situation where two threatened species co-occur but utilise different microhabitats, which is underpinned in their functional morphological traits. These two distantly related damselfly species are a good case for comparative studies because their distinct morphological differences are evident and adequate to infer the evolutionary significance of life history and behaviour. Functional traits, when taken in the context of sympatric species, are an important tool for understanding differentiation or potential interaction between species. While this study is not adequate to decipher the historical pathways that influenced these evolutionary traits, it reveals adaptations to different microhabitats and the co-occurrence of damselflies in montane streams. The intimation of this conservation dilemma is ensuring habitat heterogeneity as a common ground for ecosystem restoration when multiple threatened species are involved. Declarations Acknowledgements We wish to thank Prof. Eucharia Kenya for her constructive comments and suggestions that substantially improved the quality of this research. Special gratitude goes to Samuel Thuita and Maria Dzine for their assistance. We wish to thank Dr. Peter Mosongo for the suggestions that greatly contributed to the quality of the original manuscript. Author contributions Conceptualisation: AMK, AWK, and MNG; methodology: AMK, IWN, and AWK; formal analysis: AMK, data collection: AMK, AWK, and MNG; data curation: AMK; writing—original draft preparation: AMK and IWN; writing—review and editing: AWK, MNG, and AWG; visualisation: AMK; supervision: AWK and MNG; funding acquisition: AMK, IWN, AWK, AWG, and MNG. All authors have read and agreed to the published version of the manuscript. Funding AMK was funded by the Mohamed bin Zayed Species Conservation Fund under the project ‘220529140: Research and Conservation of the Critically Endangered Kenya Montane Dancing Jewel ( Platycypha amboniensis ) in Mount Kenya.’ AG was supported by the Centre for Ecosystem Restoration - Kenya. Data availability The following supporting information: abundance, occupancy data, and AICc model results are available at (Zonedo). doi: 10.5281/zenodo.8101703 ORCIDs A Karani https://orcid.org/0000-0001-8547-9847 A Kairu https://orcid.org/0000-0001-6915-1948 A Gichira https://orcid.org/0000-0002-0530-7605 M Githaiga https://orcid.org/0000-0001-7890-1819 I Ng’iru https://orcid.org/0000-0002-4766-6251 Disclosure statement The authors declare no conflict of interest. References Baker DJ, Garnett ST, O’Connor J et al (2019) Conserving the abundance of nonthreatened species. 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Ecol Ind 133:108367. https://doi.org/10.1016/j.ecolind.2021.108367 Steenweg R, Hebblewhite M, Whittington J, McKelvey K (2019) Species-specific differences in detection and occupancy probabilities help drive ability to detect trends in occupancy. Ecosphere 10:e02639. https://doi.org/10.1002/ecs2.2639 Stoks R, Córdoba-Aguilar A (2012) Evolutionary ecology of odonata: a complex life cycle perspective. Annu Rev Entomol 57:249–265. https://doi.org/10.1146/annurev-ento-120710-100557 Viza A, Garcia-Raventós A, Ll. Riera J et al (2023) Species‐specific functional traits rather than phylogenetic relatedness better predict future range‐shift responses of odonates. Insect Conserv Divers 16:574–587. https://doi.org/10.1111/icad.12647 Webster MT, Beaurepaire A, Neumann P, Stolle E (2023) Population genomics for insect conservation. Annu Rev Anim Biosci 11:115–140. https://doi.org/10.1146/annurev-animal-122221-075025 White GC, Burnham KP (1999) Program MARK: survival estimation from populations of marked animals. Bird Study 46:S120–S139. https://doi.org/10.1080/00063659909477239 Zhou Y, Chen S, Hu G et al (2018) Species richness and phylogenetic diversity of seed plants across vegetation zones of Mount Kenya, East Africa. Ecol Evol 8:8930–8939. https://doi.org/10.1002/ece3.4428 Additional Declarations No competing interests reported. Cite Share Download PDF Status: Published Journal Publication published 05 Feb, 2026 Read the published version in Journal of Insect Conservation → Version 1 posted Editorial decision: Revision requested 22 Oct, 2025 Reviews received at journal 15 Oct, 2025 Reviews received at journal 24 Jun, 2025 Reviewers agreed at journal 16 May, 2025 Reviewers agreed at journal 11 May, 2025 Reviewers invited by journal 11 May, 2025 Editor assigned by journal 12 Apr, 2025 Submission checks completed at journal 12 Apr, 2025 First submitted to journal 12 Apr, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6433576","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":442050662,"identity":"b289dcf0-88a0-4aa7-9e2f-e8168cfc603d","order_by":0,"name":"Anthony Karani","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA6klEQVRIiWNgGAWjYDACHjApwcDAzmD4AMTnI14LM4OxAYjPRqQWBpAWMwkQTVALf8/hZx9+7rCw62dm3lb5NcdOho2B+eGjG3i0SJxtM57Ze0YieWYzW9lt2W3JQIexGRvn4LPmPIMxA2+bRLLBYR6z25LbmIFaeNik8WmRP8/+mfEvVEux5LZ6wloMzvYYMwNtsQNpYfy47TBhLYZnzhQzy7ZJJEg2sxVLM247zsPGTMAvcmfSNzO+bauz52dv3vjx57ZqEOPhY7zeh4LEBiDBDI4kZiKUg4A9iGD8QaTqUTAKRsEoGFkAAMZ/P0xfjyrwAAAAAElFTkSuQmCC","orcid":"","institution":"University of Embu","correspondingAuthor":true,"prefix":"","firstName":"Anthony","middleName":"","lastName":"Karani","suffix":""},{"id":442050663,"identity":"ff1f90fa-28cf-42d5-bfc5-af8750754cbd","order_by":1,"name":"Anne Kairu","email":"","orcid":"","institution":"University of Embu","correspondingAuthor":false,"prefix":"","firstName":"Anne","middleName":"","lastName":"Kairu","suffix":""},{"id":442050664,"identity":"d488db15-0425-469e-8a9a-79fcdc70e669","order_by":2,"name":"Michael Githaiga","email":"","orcid":"","institution":"University of Embu","correspondingAuthor":false,"prefix":"","firstName":"Michael","middleName":"","lastName":"Githaiga","suffix":""},{"id":442050665,"identity":"41c683bf-4953-4bd2-a49e-5bc8fa150083","order_by":3,"name":"Andrew Gichira","email":"","orcid":"","institution":"Centre for Ecosystem Restoration Kenya","correspondingAuthor":false,"prefix":"","firstName":"Andrew","middleName":"","lastName":"Gichira","suffix":""},{"id":442050667,"identity":"a94e771f-7c0c-431a-bb45-9b0796892a32","order_by":4,"name":"Ivy Ng’iru","email":"","orcid":"","institution":"Mpala Research Centre","correspondingAuthor":false,"prefix":"","firstName":"Ivy","middleName":"","lastName":"Ng’iru","suffix":""}],"badges":[],"createdAt":"2025-04-12 09:23:33","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6433576/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6433576/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10841-026-00750-7","type":"published","date":"2026-02-05T15:58:15+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":81503737,"identity":"e8e48d52-ba92-4144-a397-192947cd8471","added_by":"auto","created_at":"2025-04-28 04:57:04","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":624835,"visible":true,"origin":"","legend":"\u003cp\u003eThreatened Afromontane damselflies in Kenya: (a) \u003cem\u003ePlatycypha amboniensis \u003c/em\u003e(Martin, 1915), male; (b) \u003cem\u003eP. amboniensis \u003c/em\u003e(Martin, 1915)\u003cem\u003e, \u003c/em\u003ecopulation; (c) \u003cem\u003ePseudagrion bicoerulans \u003c/em\u003e(Martin, 1907), copulation; (d\u003cem\u003e) P. bicoerulans \u003c/em\u003e(Martin, 1907), male\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-6433576/v1/d87b332f9c9f22f705aff08c.png"},{"id":81503735,"identity":"56ac6b53-c1b9-46ab-b6e9-c1461fa9ed4a","added_by":"auto","created_at":"2025-04-28 04:57:03","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":234755,"visible":true,"origin":"","legend":"\u003cp\u003e(a) Map showing the distribution of \u003cem\u003eP. bicoerulans\u003c/em\u003eMartin, 1907 (shaded green) and the occurrence of \u003cem\u003eP. bicoerulans\u003c/em\u003e Martin, 1907 (black dots) and \u003cem\u003eP. amboniensis\u003c/em\u003e Martin, 1915 (blue triangles) along the elevation gradient in Afromontane forests of East Africa; (1) Mt. Kenya, (2) Aberdare Ranges, (3) Mau Forest Complex, (4) Nandi Hills, (5) Mount Elgon, and (6) Mt. Kilimanjaro and Mt. Meru; (b) map of Kirinyaga and Embu Counties (shaded blue) showing the location of Mount Kenya and sampling sites in Kangaita Forest (Red Crosses) and Irangi Forest (Yellow Squares).\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-6433576/v1/68d159fa04bda717c07f0fa7.png"},{"id":81503185,"identity":"b7994d04-d3f1-4fc8-b481-6f30a228baa8","added_by":"auto","created_at":"2025-04-28 04:49:03","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":33549,"visible":true,"origin":"","legend":"\u003cp\u003ePoint range diagram of daily abundance (mean ± sd) for both adult and nymphs of montane damselflies \u003cem\u003eP. amboniensis\u003c/em\u003e Martin, 1915 (Kenya Jewel), and \u003cem\u003eP. bicoerulans\u003c/em\u003eMartin, 1907 (Giant Sprite): (a) Irangi forest (dots) and (b) Kangaita forest (triangles).\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-6433576/v1/703080c63ebf90121393d85e.png"},{"id":102234159,"identity":"e1435d5e-200a-40cd-b61f-abc39ff5368d","added_by":"auto","created_at":"2026-02-09 16:06:47","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2027041,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6433576/v1/87206dac-5afc-4eb6-9205-d24390890394.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Congruence of detection probabilities and co-occurrence of threatened Afromontane damselflies with diverging functional traits (Odonata: Chlorocyphidae, Coenagrionidae)","fulltext":[{"header":"Introduction","content":"\u003cp\u003eInsects provide a continuum of ecosystem services that are critical for human welfare and survival (Samways et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). These services include food provisioning, nutrient cycling and decomposition of organic matter, pollination, control of pests, and use in ecological monitoring, among others (Samways et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; May \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Lately, there has been a dramatic decline in insect populations and diversity around the globe, which has inspired multiple conservation actions (Forister et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Hallmann et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). These actions are urgent because population dynamics of insects have the potential to easily translate to instability of terrestrial ecosystems (May \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eConservation of threatened insects is shifting from identification of causes of population decline to searching for mechanisms to address effects and understanding the intrinsic responses of the populations to ecosystem restoration (Webster et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Lambert et al. 2023). This information is required to inform current conservation endeavours and model future population recovery under extreme scenarios and across vast landscapes (Kawahara et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). However, this effort is limited by the paucity of reliable data on the functional traits of species, population abundance and species interactions within microhabitats (Viza et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Khelifa et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Exploring this diversity is particularly problematic for aquatic insects because they have complex life history traits that involve metamorphosis and the use of different habitats for their larval and adult stages (Kietzka et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Stoks and C\u0026oacute;rdoba-Aguilar \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). The causes of the population decline of aquatic insects is rooted in different developmental stages, resulting in an observed difference in sensitivity and vulnerability of the species (Samways \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2008\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe highest diversity of threatened dragonflies and damselflies (\u003cem\u003eOdonata\u003c/em\u003e) in Africa is concentrated in the tropical forests and highlands (Clausnitzer et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Habitat loss in the afromontane radiation due to deforestation impacts both the terrestrial and aquatic phases of odonate species (Samways \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Clausnitzer et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Clausnitzer et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Hoř\u0026aacute;k et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Damselflies (\u003cem\u003eZygoptera\u003c/em\u003e) and dragonflies (\u003cem\u003eAnisoptera\u003c/em\u003e), being sensitive to different stressors, amphibious in nature, and charismatic players in the food web as top predators of aquatic and terrestrial invertebrates and prey for vertebrates, are potent and powerful ecological indicators (May \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The knowledge of their ecology and diversity is crucial to conservation biology (Dijkstra et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Much of the current work on odonate populations in East Africa has focused on biogeography and taxonomy (Njoroge et al. 2017; Clausnitzer et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). These include the characterisation of extinction risk based on trends in population abundance, geographical distribution, and habitat fragmentation (Clausnitzer et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSpecies-specific functional traits of larvae and adult damselflies that can contribute to population resilience of threatened species in subtle ecological settings remain unknown. Understanding the abundances of species is important for their conservation, whether threatened or not threatened (Hogreve and Suhling \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Baker et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Understanding the detection probabilities of species is important to help scientists and local communities in awareness creation, research, planning, and ecological restoration (Redford et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Equally, knowledge on species co-occurrence can be used to inform how communities function, such as revealing patterns in competition and niche overlap (Steenweg et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The aim of this study was to estimate the abundance and probabilities of co-occurrence of Giant Sprite, \u003cem\u003ePseudagrion bicoerulans\u003c/em\u003e Martin, 1907 (Coenagionidae), and Kenya Montane Jewel, \u003cem\u003ePlatycypha amboniensis\u003c/em\u003e Martin, 1915 (Chlorocyphidae), in Mount Kenya Forest. The study also aimed at elucidating functional traits of ecological significance based on the morphology of the adults and final stadium instar larvae of the two damselfly species.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy species\u003c/h2\u003e \u003cp\u003eTwo threat-defined and range-restricted species of damselflies (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) were studied. \u003cem\u003ePlatycypha amboniensis\u003c/em\u003e is endemic to the Central Highlands of Kenya and is highly dependent on montane forest and considered a sub-montane relict species among dancing jewels (Clausnitzer et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Little is known about its reproductive ecology, behaviour, and life history, even though it is a very rare and highly threatened species (Dijkstra and Clausnitzer \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). The species is classified as Critically Endangered (CR) due to extensive habitat loss as a result of forest clearance within regions it is dependent upon (Clausnitzer et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). The male of this species has bright orange tibiae that are slightly flattened and white interiorly, sky-blue abdominal segments (S5\u0026ndash;S10), and large, bulbous eyes (Dijkstra and Clausnitzer \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Similarly, \u003cem\u003eP. bicoerulans\u003c/em\u003e is endemic to streams in the mountains of Kenya, Tanzania, and Uganda. The species is classified as Vulnerable (VU) in the IUCN Red List. It was termed \u0026lsquo;the most alpine of African Odonata\u0026rsquo; because it occurs in heather and afroalpine zones above 3000 m above sea level (Clausnitzer et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). The species has high intraspecific genetic diversity and three subspecies: \u003cem\u003eP. b. bicoerulans\u003c/em\u003e (Central Kenya), \u003cem\u003eP. b. elgonensi\u003c/em\u003e (West Kenya and East Uganda), and \u003cem\u003eP. b. kilimanjaricus\u003c/em\u003e (North Tanzania), widespread across high-elevation forests in East Africa (Dijkstra et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Males of this species are distinguished by a bright yellow to orange labrum and small orange post-ocular spots, a green-striped black thorax, blue or pruinescent abdominal segments (S1\u0026ndash;S2, S9\u0026ndash;S10), and clasper-like cerci (Dijkstra and Clausnitzer \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eStudy area\u003c/h3\u003e\n\u003cp\u003eThe study was conducted in Kangaita Forest Station (Kirinyaga County) and Irangi Forest Station (Embu County) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). These forest stations are part of the Mount Kenya Forest Reserve, which is a protected area in the Central Highlands of Kenya (Nyongesa and Vacik \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). This study area is classified as lower montane humid forest and is renowned for its agro-ecological value and high diversity of woody plant species, birds, and mammals (Onditi et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Zhou et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The drainage pattern is dendritic, with all streams forming tributaries of the Tana River. Water is directly abstracted in large quantities from these streams for domestic use, municipal supply, microhydropower stations, and small- and large-scale irrigation schemes.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eSampling\u003c/h3\u003e\n\u003cp\u003eThe sampling period ranged from September 2022 to March 2023 and is considered in this study a 'single season', which runs across two dry periods interspaced with a period of short rains. It is possible that the adults and final instar larvae encountered during this study belong to the same generation. Sampling sites were selected randomly from a list of first-level and second-level streams (five in Kangaita and five in Irangi). At each sampling point, three transects of 100 m in length and covering the width of the stream channel (up to 10 m) were established along each stream following the method of Batista et al. (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Sampling of adults, exuviae, and larvae was undertaken at four sampling occasions (interval: 9\u0026ndash;17 days) along each transect at twenty-meter intervals, starting at zero, which translates to five full-width belt transects (FWBT) on each transect (Darshetkar, Patwardhan, and Koporde 2023), equivalent to 200 m\u003csup\u003e2\u003c/sup\u003e. The following water quality parameters were measured using a multi-parameter portable meter (Suzhou Holi, China): Dissolved Oxygen (DO), Electrical Conductivity (EC), Oxidation-reduction potential (ORP), pH, total dissolved solids (TDS), and temperature. Turbidity was measured using a nephelometer (BD PhoenixSpec\u0026trade;, South Africa).\u003c/p\u003e \u003cp\u003eSampling was consistent with the guidelines for handling living insects (Montegomery et al. 2021; Cez\u0026aacute;rio et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). On each sampling occasion, two observers were involved and took turns in data entry and observation. All adult damselflies, exuviae, and nymphs encountered at each interval were counted. Evidence of the presence of adults was confirmed by the presence of damselflies perching on the rocks, twigs overhanging the stream, emergent vegetation, and driftwood. The sampling period was mostly sunny, and the water was shallow. No attempts were made to search for adult damselflies beyond the riverbanks. Exuviae were found by locating suitable emergence sites mostly on emergent vegetation, driftwood, and exposed rocks. Collected exuviae were placed in Falcon tubes and stored in a freezer in order to prevent damage by inadvertently trapped insects and spiders and the growth of mould. After at least 12 hours of freezing, the exuviae were dried and mounted.\u003c/p\u003e \u003cp\u003eLarvae were identified by sinking a dip net into the water and scooping debris at the edges of the stream (mostly giant sprite). Movable rocks along the river were also lifted or overturned and checked for clinging or crawling damselfly larvae (mostly montane jewel). These were then returned to position at the point of collection. On account of the rarity of the two species, only a few larvae and adults were collected. Adults were collected with a professional sweep net (38 cm diameter, 86 cm depth, and 100 cm handle). Specimens were placed in specimen jars and washed in acetone or preserved in absolute ethanol. The following parameters were measured for adult damselflies: abdomen length, body length, hind femur length, thorax length, wing area, wing length, wing perimeter, and wing width. The following parameters were measured for nymphs: abdomen length, appendages, body length, gonapophyses, hind femur length, prementum, and wing sheath.\u003c/p\u003e\n\u003ch3\u003eData analyses\u003c/h3\u003e\n\u003cp\u003eThe pairwise correlation between water properties was calculated using Pearson corrrelation coefficients. The difference in functional traits between males and females was estimated using an independent samples t-test. The replicated count data (pooled to stream level and treated separately for adults and nymphs of both species) were averaged as daily abundance with standard deviation (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;sd). A ratio was calculated by dividing the mean daily abundance of nymphs by that of adults. An independent sample t-test was used to determine if there were significant differences in means. These tests were carried out in R versions x64: 4.0.3 (R Core Team \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSpecies abundance data denoted as presence-absence at each sampling interval and organised into pairs (\u003cem\u003eP. amboniensis\u003c/em\u003e adult\u0026thinsp;+\u0026thinsp;\u003cem\u003eP. bicoerulans\u003c/em\u003e adult; \u003cem\u003eP. amboniensis\u003c/em\u003e nymph\u0026thinsp;+\u0026thinsp;\u003cem\u003eP. bicoerulans\u003c/em\u003e nymph) to fit the formulation of two-species single-season occupancy models (Richmond et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). This model accepts detection histories of each species at each site across sampling occasions (0 indicating non-detection and 1 indicating detection). The model assumes that one of the species in pairwise interactions is dominant while the other is subordinate (Kleiven et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Lonsinger \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Three parameters were used to estimate occupancy: ψA\u0026mdash;occupancy probability of species A (\u003cem\u003eP. amboniensis\u003c/em\u003e); ψBA\u0026mdash;occupancy probability of species B (\u003cem\u003eP. bicoerulans\u003c/em\u003e) in the presence of species A; and ψBa\u0026mdash;occupancy probability of species B in the absence of species A. These were used to test the hypothesis that the occupancy probability of \u003cem\u003eP. amboniensis\u003c/em\u003e is not dependent on the presence of \u003cem\u003eP.bicoerulans\u003c/em\u003e (ψBA\u0026thinsp;\u0026ne;\u0026thinsp;ψBa).\u003c/p\u003e \u003cp\u003eFive parameters were estimated for detection probability, namely: pA\u0026mdash;detection probability of species A in the absence of species B; pB\u0026mdash;detection probability of species B in the absence of species A; rA\u0026mdash;detection probability of species A in the presence of species B; rBA\u0026mdash;detection probability of species B when species A is present and detected; rBa\u0026mdash;detection probability of species B when species A is present but not detected. These were used to test hypotheses that 1) the detection probability of \u003cem\u003eP. amboniensis\u003c/em\u003e is not dependent on the presence of \u003cem\u003eP. bicoerulans\u003c/em\u003e (pA\u0026thinsp;\u0026ne;\u0026thinsp;rA) and 2) the detection probability of \u003cem\u003eP. bicoerulans\u003c/em\u003e is not dependent on the presence and detection of \u003cem\u003eP. amboniensis\u003c/em\u003e (pB\u0026thinsp;\u0026ne;\u0026thinsp;rBA). The model also estimates derived parameters of occupancy probability: ψB\u0026mdash;probability of occupancy for species B, regardless of occupancy status of species A; ψAB\u0026mdash;probability of both species being present. The species interaction factor (SIF or γ) was used to measure co-occurrence, where γ\u0026thinsp;=\u0026thinsp;1 means that the two species occur independently of each other, γ\u0026thinsp;\u0026lt;\u0026thinsp;1 means that the two species avoid each other and γ\u0026thinsp;\u0026gt;\u0026thinsp;1 means that there is overlap in the co-occurrence of the two species (Lonsinger \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2022\u003c/span\u003e)\u003c/p\u003e \u003cp\u003eA logit link function was used to model the occupancy and detection of both species in the program MARK (White and Burnham \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). This was conducted separately for pairs of adults and nymphs. A total of 32 predetermined models were developed to include combinations of constant occupancy probability ψ(.), variable occupancy probability ψ(t), constant detection probability p(.), and variable detection probability p(t). All candidate models were ranked from highest to lowest based on their Akaike\u0026rsquo;s Information Criteria modified for c-hat (QAICc) values and the AIC weights. The top-ranked model has a model likelihood of one (1), the lowest QAICc, and presents the best combination of parameters (ψA, ψBA, ψBa, rA, pA, pB, rBa, rBA).\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eThe physico-chemical water properties for each stream are shown in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. There was no significant correlation between dissolved oxygen (DO) and oxidation-reduction potential (ORP) and no correlation between turbidity and pH. In contrast, EC was strongly positively correlated with TDS (r\u0026thinsp;=\u0026thinsp;0.93) and negatively correlated with ORP (r = -0.78). Temperature was negatively correlated with pH (r = -0.72), while OPR was negatively correlated with TDS (-0.79). Several moderate and weak relationships were observed, such as the inverse between temperature and DO.\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\u003eMean and range of water quality parameters measured from forest streams in Kangaita and Irangi Forests, Mount Kenya\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"8\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"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=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSTREAM\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDO\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEC\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eORP\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePH\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTDS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eTEMPERATURE\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eTURBIDITY\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCiaminogia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3.9 (3.9\u0026ndash;3.9)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e36 (32\u0026ndash;42)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e183 (159\u0026ndash;208)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7.837 (7.68\u0026ndash;7.93)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e18.33 (17\u0026ndash;21)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e18 (16.7\u0026ndash;18.9)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e8.34 (3.43\u0026ndash;15.53)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRwamuthambi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e4.18 (3.6\u0026ndash;4.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e26.5 (26\u0026ndash;27)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e270.5 (247\u0026ndash;294)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7.875 (7.86\u0026ndash;7.89)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e13 (13\u0026ndash;13)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e14.6 (13.9\u0026ndash;15.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e4.22 (2.64\u0026ndash;6.52)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMukengeria\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3.9 (3.8\u0026ndash;4.2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e15.3 (14\u0026ndash;16)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e334 (153\u0026ndash;585)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7.99 (7.75\u0026ndash;8.14)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7.33 (7\u0026ndash;8)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e15.167 (14.1\u0026ndash;16.5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e7.39 (2.77\u0026ndash;15.63)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRuiru\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e4.4 (4.2\u0026ndash;4.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e38.6 (34\u0026ndash;42)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e189 (163\u0026ndash;212)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7.32 (7.11\u0026ndash;7.56)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e15.25 (14\u0026ndash;17)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e14.52 (13.7\u0026ndash;16.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e11.38 (2.92\u0026ndash;16.06)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRundu\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e4.56 (4.1\u0026ndash;4.9)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e50.4 (46\u0026ndash;68)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e215.2 (135\u0026ndash;381)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7.67 (6.61\u0026ndash;8.26)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e25.6 (23\u0026ndash;35)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e19.02 (17.1\u0026ndash;22.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e4.02 (0.65\u0026ndash;8.35)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEna\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e5.56 (5.2\u0026ndash;5.9)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e40.75 (23\u0026ndash;64)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e232.5 (204\u0026ndash;273)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7.92 (7.22\u0026ndash;8.53)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e20.75 (13\u0026ndash;32)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e13.15 (12.2\u0026ndash;14.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e4.69 (3.1\u0026ndash;6.9)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKarutune\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e4.65 (4.6\u0026ndash;4.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e42.25 (36\u0026ndash;46)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e174.6 (108\u0026ndash;248)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7.94 (7.14\u0026ndash;8.65)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e21.6 (17\u0026ndash;24)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e15.85 (13.9\u0026ndash;18.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e4.317 (2.27\u0026ndash;6.2)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKiye\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e5.05 (4.4\u0026ndash;5.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e37.8(16\u0026ndash;47)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e185.8 (163\u0026ndash;201)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7.91 (7.21\u0026ndash;8.4)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e24.8(12\u0026ndash;48)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e17.92 (12.9\u0026ndash;22.8)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e5.07 (3.72\u0026ndash;6.52)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNyanjara\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e6.25 (6.2\u0026ndash;6.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e34.5(18\u0026ndash;48)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e265.75(208\u0026ndash;373)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8.11 (7.75\u0026ndash;8.37)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e21 (17\u0026ndash;23)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e13.2 (12.5\u0026ndash;13.9)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e4.55 (3.4\u0026ndash;6.5)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eThambana\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3.95 (3.6\u0026ndash;4.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e38.6 (32\u0026ndash;45)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e199 (124\u0026ndash;257)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7.95 (7.43\u0026ndash;8.64)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e21.2 (18\u0026ndash;26)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e12.56 (10.5\u0026ndash;14.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e3.73 (2.39\u0026ndash;5.6)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"8\"\u003e\u003cb\u003eNote\u003c/b\u003e: Abbreviations; DO: Dissolved Oxygen, EC: Electrical Conductivity, ORP: Oxidation-Reduction Potential, pH: Acidity/Alkalinity, TDS: Total Dissolved Solid\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eTable\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e\u003c/p\u003e \u003cp\u003eThere was a significant difference between males and females of \u003cem\u003eP. amboniensis\u003c/em\u003e for abdomen length, body length, wing area, and wing length. No significant differences were found for hind femur length, thorax length, wing perimeter, and wing width (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). There was no evidence that the adult males and females of \u003cem\u003eP. bicoerulans\u003c/em\u003e differed significantly. The male and female nymphs of both \u003cem\u003eP. bicoerulans\u003c/em\u003e and \u003cem\u003eP. amboniensis\u003c/em\u003e did not differ in abdomen length, appendages, body length, hind femur length, prementum, and wing sheath at emergence. There was a significant difference in the size of the gonapophyses of \u003cem\u003eP. bicoerulans\u003c/em\u003e and the prementum of \u003cem\u003eP. amboniensis.\u003c/em\u003e\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\u003eMorphological traits of adults and nymphs of \u003cem\u003eP. amboniensis\u003c/em\u003e and \u003cem\u003eP. bicoerulans\u003c/em\u003e in Mount Kenya\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"11\"\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 \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTrait\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFemale\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMale\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTtest\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003edf\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ep-value\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eFemale\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eMale\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eTtest\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003edf\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e \u003cp\u003ep-value\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c6\" namest=\"c2\"\u003e \u003cp\u003e\u003cb\u003eAdult\u003c/b\u003e: \u003cb\u003ePlatycypha amboniensis\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c11\" namest=\"c7\"\u003e \u003cp\u003e\u003cb\u003eAdult\u003c/b\u003e: \u003cb\u003ePseudagrion bicoerulans\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAbdomen length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e17.8 (17.5\u0026ndash;18.2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e19.95 (18.9\u0026ndash;21.2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-8.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e11.95\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026gt;\u0026thinsp;.001*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e33.4 (32.2\u0026ndash;35.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e34.36 (32.3\u0026ndash;36)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e6.47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.411\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBody length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e29.7 (28.2\u0026ndash;30.8)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e33.32 (32.1\u0026ndash;35.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-5.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.003*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e42.7 (40.9\u0026ndash;47)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e43.54 (41.7\u0026ndash;46.2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e4.80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.634\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHind femur length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5.58 (4.9\u0026ndash;6.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.49 (4.9\u0026ndash;6.2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.847\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e4.88 (4.3\u0026ndash;5.4)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5(4.3\u0026ndash;5.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e5.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.704\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eThorax length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e8.55 (7.7\u0026ndash;9.4)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e8.79 (7.9\u0026ndash;10.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.629\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e6.85 (6.6\u0026ndash;7.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6.48 (5.6\u0026ndash;7.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e6.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.299\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWing area\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e100.94 (87.28-120.32)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e84.37 (68.91\u0026ndash;93.68)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e11.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026gt;\u0026thinsp;.001*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e98.87 (85.37-105.89)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e101.68 (74.81-127.06)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-1.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e39.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.178\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWing length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e27.83 (25.73\u0026ndash;30.34)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e25.81 (24.64\u0026ndash;27.24)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e24.86\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.002*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e26.73 (25.7\u0026ndash;27.5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e26.36 (25.4\u0026ndash;27.4)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e29.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.418\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWing perimeter\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e80.96 (71.96\u0026ndash;90.35)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e72.27 (62.01-127.43)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e31.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.058\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e82.15 (61.85-139.56)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e84.99 (61.34-145.48)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e25.39\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.726\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWing width\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5.76 (5.19\u0026ndash;6.36)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.49 (4.69\u0026ndash;9.25)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e6.11 (5.36\u0026ndash;6.65)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6.16 (5.15\u0026ndash;8.24)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.64\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e6.86\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.541\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=\"5\" nameend=\"c6\" namest=\"c2\"\u003e \u003cp\u003e\u003cb\u003eNymph\u003c/b\u003e: \u003cb\u003ePlatycypha amboniensis\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"5\" nameend=\"c11\" namest=\"c7\"\u003e \u003cp\u003e\u003cb\u003eNymph\u003c/b\u003e: \u003cb\u003ePseudagrion bicoerulans\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAbdomen length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e10.43 (9.2\u0026ndash;11.2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10.23 (9.6\u0026ndash;10.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.96\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.741\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e12.27 (12.1\u0026ndash;12.5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e12.27 (11.7\u0026ndash;12.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e2.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAppendages\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5.73 (5.3\u0026ndash;6.2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.77 (5.4\u0026ndash;6.2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.902\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e6.77 (6.4-7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6.67 (6.1-7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e3.44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.786\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBody length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e22.63 (22.1\u0026ndash;23.5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e22.3 (22.1\u0026ndash;22.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.91\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.407\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e25.47 (25.1\u0026ndash;26)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e25.5 (25.1\u0026ndash;25.9)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e3.89\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.93\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGonapophyses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.98 (1.8\u0026ndash;2.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.97 (1.9\u0026ndash;2.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.87 (0.8-1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e11.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e4.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e\u0026gt;\u0026thinsp;.001*\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHind femur length\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5.85 (5.5\u0026ndash;6.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.67(5.3-6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.57\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.596\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e5.3 (5.2\u0026ndash;5.5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.4 (5.3\u0026ndash;5.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e4.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.519\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePrementum\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3.63 (3.6\u0026ndash;3.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.8 (3.8\u0026ndash;3.8)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-7.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.006*\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3.4 (3.3\u0026ndash;3.5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e3.67 (3.3-4)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-1.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e2.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.318\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWing Sheath\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5.5 (5.4\u0026ndash;5.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.57 (5.4\u0026ndash;5.8)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.662\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e6.27 (6.1\u0026ndash;6.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6.4 (6.2\u0026ndash;6.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e3.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.587\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe abundance of both adults and nymphs differed between species in all sampling sites (t = -3.99, df\u0026thinsp;=\u0026thinsp;33.76, p-value\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The damselflies, \u003cem\u003eP. amboniensis\u003c/em\u003e and \u003cem\u003eP. bicoerulans\u003c/em\u003e, were encountered in the study area a total of 3605 times. These included 871 adults and 613 nymphs of \u003cem\u003eP. amboniensis\u003c/em\u003e and 906 adults and 1215 nymphs of \u003cem\u003eP. bicoerulans\u003c/em\u003e. Additionally, the ratio of adults and nymphs at sampling sites differed between species (t\u0026thinsp;=\u0026thinsp;3.68, df\u0026thinsp;=\u0026thinsp;40.4, p-value\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The average number of individuals encountered across sampling occasions was (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;sd) 19.85\u0026thinsp;\u0026plusmn;\u0026thinsp;8.96, 16.70\u0026thinsp;\u0026plusmn;\u0026thinsp;8.47, 22.00\u0026thinsp;\u0026plusmn;\u0026thinsp;10.22, and 33.20\u0026thinsp;\u0026plusmn;\u0026thinsp;13.35 for \u003cem\u003eP. amboniensis\u003c/em\u003e adults, \u003cem\u003eP. amboniensis\u003c/em\u003e nymphs, \u003cem\u003eP. bicoerulans\u003c/em\u003e adults, and \u003cem\u003eP. bicoerulans\u003c/em\u003e nymphs, respectively, in Irangi Forest; 23.70\u0026thinsp;\u0026plusmn;\u0026thinsp;9.43, 13.95\u0026thinsp;\u0026plusmn;\u0026thinsp;3.55, 23.30\u0026thinsp;\u0026plusmn;\u0026thinsp;11.81, and 27.55\u0026thinsp;\u0026plusmn;\u0026thinsp;8.31 for \u003cem\u003eP. amboniensis\u003c/em\u003e adults, \u003cem\u003eP. amboniensis\u003c/em\u003e nymphs, \u003cem\u003eP. bicoerulans\u003c/em\u003e adults, and \u003cem\u003eP. bicoerulans\u003c/em\u003e nymphs, respectively, in Kangaita Forest (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). This translates to a density of 0.218 sqm⁻\u0026sup1;, 0.153 sqm⁻\u0026sup1;, 0.227 sqm⁻\u0026sup1;, and 0.308 sqm⁻\u0026sup1; of \u003cem\u003eP. amboniensis\u003c/em\u003e adults and \u003cem\u003eP. amboniensis\u003c/em\u003e nymphs, \u003cem\u003eP. bicoerulans\u003c/em\u003e adults, and \u003cem\u003eP. bicoerulans\u003c/em\u003e nymphs, respectively.\u003c/p\u003e \u003cp\u003eTable\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e\u003c/p\u003e \u003cp\u003eThe most parsimonious models (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e) estimated the probability of occupancy for combinations of \u003cem\u003eP. amboniensis\u003c/em\u003e adults\u0026mdash;\u003cem\u003eP. bicoerulans\u003c/em\u003e adults and \u003cem\u003eP. amboniensis\u003c/em\u003e nymphs\u0026mdash;\u003cem\u003eP. bicoerulans\u003c/em\u003e nymphs but showed no evidence that occupancy of either was influenced by occupancy of the other (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The best-supported model for adult occurrence had an AICc value of 645.11, a weight of 0.356, and the model likelihood reduced dramatically to 0.058 in the sixth model (ΔAICc\u0026thinsp;=\u0026thinsp;5.703). The best-supported model for nymph co-occurrence had an AICc value of 603.84, a weight of 0.333, and the model likelihood reduced to 0.086 in the sixth model (ΔAICc\u0026thinsp;=\u0026thinsp;4.907).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eCandidate models used to evaluate co-occurrence and detection probability of \u003cem\u003ePlatycypha amboniensis\u003c/em\u003e Martin, 1915 and \u003cem\u003ePseudagrion bicoerulans\u003c/em\u003e Martin, 1907, using two-species conditional occupancy model\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\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 \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOccupancy\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDetection\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAICc\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eΔAICc\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eWeight\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eLikelihood\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNum. Par\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"7\" nameend=\"c7\" namest=\"c1\"\u003e \u003cp\u003eCo-occurrence between nymphs of \u003cem\u003eP. amboniensis\u003c/em\u003e and \u003cem\u003eP. bicoerulans\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA(.) ψBA(.) ψBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epA(.) pB(.) rA(.) rBA(.) rBa(t)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e645.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.356\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA(.) ψBA(.) ψBa(t)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epA(.) pB(.) rA(.) rBA(.) rBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e645.61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.504\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.276\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.777\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA(.) ψBA(t) ψBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epA(.) pB(.) rA(t) rBA(.) rBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e647.44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.331\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.111\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.312\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA(t) ψBA(.) ψBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epA(.) pB(.) rA(.) rBA(t) rBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e647.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.562\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.099\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.278\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA(.) ψBA(t) ψBa(t)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epA(.) pB(t) rA(.) rBA(.) rBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e647.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.640\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.095\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.267\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA(t) ψBA(t) ψBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epA(t) pB(.) rA(.) rBA(.) rBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e650.81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.703\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.020\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.058\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"7\" nameend=\"c7\" namest=\"c1\"\u003e \u003cp\u003eCo-occurrence between adults of \u003cem\u003eP. amboniensis\u003c/em\u003e and \u003cem\u003eP. bicoerulans\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA(.) ψBA(.) ψBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epA(.) pB(.) rA(.) rBA(.) rBa(t)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e603.84\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.333\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA(.) ψBA(.) ψBa(t)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epA(.) pB(.) rA(.) rBA(.) rBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e604.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.357\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.278\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.837\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA(.) ψBA(t) ψBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epA(.) pB(.) rA(t) rBA(.) rBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e606.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.455\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.097\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.298\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA(t) ψBA(.) ψBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epA(.) pB(.) rA(.) rBA(t) rBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e606.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.592\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.091\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.274\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA(.) ψBA(t) ψBa(t)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epA(.) pB(t) rA(.) rBA(.) rBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e608.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.739\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.031\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.094\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA(t) ψBA(t) ψBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003epA(t) pB(.) rA(.) rBA(.) rBa(.)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e608.72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.907\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.029\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.086\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e8\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 \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eEstimates of occupancy (ψ), detection probability (p and r), and Species interaction factor (SIF or γ) for \u003cem\u003ePlatycypha amboniensis\u003c/em\u003e Martin, 1915 and \u003cem\u003ePseudagrion bicoerulans\u003c/em\u003e Martin, 1907, determined using two-species conditional occupancy model with data from Irangi and Kangaita Forests in Mount Kenya\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\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 \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c4\" namest=\"c2\"\u003e \u003cp\u003eNymphs\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c7\" namest=\"c5\"\u003e \u003cp\u003eAdults\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCo-occurrence\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEstimate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSE\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eRange\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEstimate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSE\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eRange\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.995\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.049\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(0.884\u0026ndash;1.000)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.915\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.061\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(0.695\u0026ndash;0.981)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψBA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.969\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.060\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(0.377\u0026ndash;0.999)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.971\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.032\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(0.787\u0026ndash;0.997)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψBa\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.000\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.000\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(1.000\u0026ndash;1.000)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.000\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.000\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(1.000\u0026ndash;1.000)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.473\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.390\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(0.040\u0026ndash;0.951)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.625\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.122\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(0.256\u0026ndash;0.744)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003epB\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.140\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.589\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(0.000\u0026ndash;1.000)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.504\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.497\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(0.252\u0026ndash;0.966)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003erA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.334\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.036\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(0.268\u0026ndash;0.408)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.338\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.065\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(0.225\u0026ndash;0.478)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003erBA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.429\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.117\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(0.227\u0026ndash;0.657)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.328\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.090\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(0.181\u0026ndash;0.524)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003erBa\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.401\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.048\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(0.311\u0026ndash;0.499)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.308\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.076\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(0.183\u0026ndash;0.473)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eγ\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.999\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(0.995\u0026ndash;1.002)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.997\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(0.998\u0026ndash;1.001)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψB\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.960\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.042\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(0.736\u0026ndash;0.995)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.973\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.018\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(0.922\u0026ndash;0.993)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eψAB\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.926\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.044\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e(0.781\u0026ndash;0.978)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.888\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.043\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(0.853\u0026ndash;0.984)\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\u003eTable\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e\u003c/p\u003e \u003cp\u003eThere was no evidence that occupancy of \u003cem\u003eP. amboniensis\u003c/em\u003e was affected by the presence of \u003cem\u003eP. bicoerulans\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The species had high probabilities of occupancy (0.915\u0026ndash;1.00) regardless of whether the other species was present or not present (ψA\u0026thinsp;=\u0026thinsp;ψBA\u0026thinsp;=\u0026thinsp;ψBa\u0026thinsp;=\u0026thinsp;ψB\u0026thinsp;\u0026cong;\u0026thinsp;1.00); the probability of both species being present was 0.926\u0026thinsp;\u0026plusmn;\u0026thinsp;0.044 for nymphs and 0.888\u0026thinsp;\u0026plusmn;\u0026thinsp;0.043 for adults. The two species were present and detected independently (γ\u0026thinsp;=\u0026thinsp;0.999\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002). There was evidence that the presence of \u003cem\u003eP. amboniensis a\u003c/em\u003effected the presence and detection probability of \u003cem\u003eP. bicoerulans\u003c/em\u003e. The detection probability for nymphs of \u003cem\u003eP. amboniensis\u003c/em\u003e was higher when the adults of \u003cem\u003eP. bicoerulans\u003c/em\u003e were absent (pA\u0026thinsp;=\u0026thinsp;0.473\u0026thinsp;\u0026plusmn;\u0026thinsp;0.39). The detection probability of \u003cem\u003eP. bicoerulans\u003c/em\u003e nymphs was lowest when the nymphs of \u003cem\u003eP. amboniensis\u003c/em\u003e were present (pB\u0026thinsp;=\u0026thinsp;0.14\u0026thinsp;\u0026plusmn;\u0026thinsp;0.589). There was support that the detection probability of \u003cem\u003eP. amboniensis\u003c/em\u003e adults was influenced by the detection probability (but not presence) of \u003cem\u003eP. bicoerulans\u003c/em\u003e (rA\u0026thinsp;\u0026ne;\u0026thinsp;pA). The detection probability for adults of \u003cem\u003eP. amboniensis\u003c/em\u003e was higher when the adults of \u003cem\u003eP. bicoerulans\u003c/em\u003e were absent (pA\u0026thinsp;=\u0026thinsp;0.625\u0026thinsp;\u0026plusmn;\u0026thinsp;0.122).\u003c/p\u003e \u003cp\u003eTable\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe study species occur in running water but display clear differences in habitat requirements. The differences and similarities in the morphology of \u003cem\u003eP. amboniensis\u003c/em\u003e and \u003cem\u003eP. bicoerulans\u003c/em\u003e larvae that are related to ecological adaptations are summarised in Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e. These species belong in distinct lineages that have followed different evolutionary trajectories for an extended period (Dijkstra et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). This study contributes towards understanding the patterns of co-occurrence and structure of Afromontane damselfly communities as mediated by species-specific adaptations to varying microhabitats. As expected, \u003cem\u003eP. b. bicoerulans\u003c/em\u003e (\u003cem\u003esensu stricto\u003c/em\u003e) occur in the same streams occupied by \u003cem\u003eP. amboniensis\u003c/em\u003e in the Central Highlands of Kenya. The study showed neither competition nor niche overlap in the occupancy of \u003cem\u003eP. amboniensis\u003c/em\u003e adults\u0026mdash;\u003cem\u003eP. b. bicoerulans\u003c/em\u003e adults as well as \u003cem\u003eP. amboniensis\u003c/em\u003e nymphs\u0026mdash;and \u003cem\u003eP. bicoerulans\u003c/em\u003e nymphs. It is also noteworthy that the method assumes population closure (\u003cem\u003esensu\u003c/em\u003e Richmond et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), which is limited when associated with open populations (Kleiven et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Lonsinger \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSimilarities and differences in larval morphology of \u003cem\u003ePlatycypha amboniensis\u003c/em\u003e Martin 1915 and \u003cem\u003ePseudagrion bicoerulans\u003c/em\u003e Martin 1907 and functional traits that account for ecological adaptations\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\u003eMorphology\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFunctional traits\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEcological adaptations\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eP. amboniensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGeneral habitus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBody is short, stout, cylindrical, and setose; muddy brown to dark brown; legs are long\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNymphs sit and wait to ambush prey; their stout body and long legs are suitable for clinging on stones and gravel substrates; setae attract mud or debris for camouflage in dark/brown environments\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMouthparts\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLabium brown without setae, labial palps with long hooks, mandibles setose and heavily sclerotized, hypopharynx well developed\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLabium colour blends with the environment, hooks for grasping prey, and strong mouthparts for processing food with hard cuticles\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCaudal lamellae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEpiproct rudimentary; paraprocts elongate, triquetral, setose and spinose\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLong lamellae with spines used for respiration in benthic region with lower dissolved oxygen; the shape enhances balance in current waters and autotomy to escape predators\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGonapophyses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eWell developed in female, marginal in male\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFast development of nymphs\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eP. bicoerulans\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGeneral habitus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBody is elongate, cylindrical, pale or dark brown; legs are short\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNymphs actively search for prey; their colour blends with litter and vegetation in littoral zone\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMouthparts\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLabium long, pale to translucent, with premental/labial setae; labial palps wide\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePale colour of labium for \u0026lsquo;invisibility\u0026rsquo; while ambushing prey, wide labial palps for better clutching of larger prey, setae enhance sensitivity\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCaudal lamellae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEpiproct and paraprocts are elongate, leaf-like, flattened, and pigmented\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLamella used for swimming, respiration among vegetation, agonistic display among conspecifics, and autotomy to escape predators\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGonapophyses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eWell developed in male and female\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eQuick maturation of adults\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\u003eTable\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e\u003c/p\u003e \u003cp\u003eFunctional traits determine the distribution and structure of \u003cem\u003eOdonata\u003c/em\u003e assemblages in streams (Silva et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The evolution of morphological traits in \u003cem\u003eOdonata\u003c/em\u003e larvae is associated with requirements for larval survival, such as breathing, feeding, and predation refuge. The independent co-occurrence of \u003cem\u003eP. amboniensis\u003c/em\u003e and \u003cem\u003eP. bicoerulans\u003c/em\u003e can be explained in terms of their feeding ecology as a trade-off for 1) different prey items in littoral zone and fast-flowing sections and 2) specialised morphological features that aid processing of these prey items. The mouthparts of \u003cem\u003eP. amboniensis\u003c/em\u003e and \u003cem\u003eP. bicoerulans\u003c/em\u003e differ in shape, size, and structure; the significance is that it enhances foraging success in varying microhabitats. The general morphology of larval mouthparts in \u003cem\u003eOdonata\u003c/em\u003e is grossly similar, meaning that these small morphological variations are insightful to explain ecological adaptations with regards to food intake in littoral and benthic zones (B\u0026uuml;sse et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; B\u0026uuml;sse and Gorb \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The presence of labial palps with long hooks in \u003cem\u003eP. amboniensis\u003c/em\u003e is equipped for better grasping of prey, while the prementum with wide labial palps that have setae in \u003cem\u003eP. bicoerulans\u003c/em\u003e increases the sensitivity of the prehensile mask. In addition, the presence of a well-developed hypopharynx enhances overall food acquisition in \u003cem\u003eP. amboniensis\u003c/em\u003e, as its muscles have been associated with the opening of the labium (B\u0026uuml;sse and Gorb \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe littoral and benthic regions of lotic streams differ in their mechanisms of retaining dissolved oxygen. Stream sections with rapids and stones are cooler and abundant in dissolved solids, while littoral sections are vegetated and warmer. Caudal lamellae with spines allow the larvae to survive through active gaseous exchange as well as predator avoidance. On the other hand, increased sedimentation and substrate homogenisation favour larvae that cling, swim, or climb on substrates (Mendes et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Predator avoidance can account for disparities in microhabitat selection, resulting in trait compensation in morphological and behavioural defences in damselflies (Pernecker et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Damselflies have evolved autotomy, where caudal lamellae are self-amputated to escape predators (Black et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The drab colour and detritus particles attaching to the setae of \u003cem\u003eP. amboniensis\u003c/em\u003e make it well camouflaged, while the swimming capabilities of \u003cem\u003eP. bicoerulans\u003c/em\u003e make it easy to escape predators. A robust and cylindrical body shape (\u003cem\u003eP. amboniensis\u003c/em\u003e) is preferred in microhabitats with a higher proportion of stones and gravel substrate (Pires et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2020\u003c/span\u003e); that is also utilised by burrowing predators such as gomphids (mostly \u003cem\u003eNotogomphus\u003c/em\u003e spp. and \u003cem\u003eParagomphus\u003c/em\u003e spp.). On the other hand, an elongate body (\u003cem\u003eP. bicoerulans\u003c/em\u003e) is suited for refuge among vegetation and litter, where climbing and sprawling predators live (\u003cem\u003emostly, Pinheyschna, Zygonyx\u003c/em\u003e, and \u003cem\u003eAtoconeura\u003c/em\u003e). Siltation of the streams leads to sedimentation that homogenises substrates, excluding larvae that depend on substrates in favour of burrowing species (M\u0026uuml;hlenhaupt et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Dalzochio et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eLarval development, though rarely explored, can augment sexual maturation and influence the phenology of adult reproduction. While it was beyond the scope of this study to examine the development cycle of these species, the observed differences in gonapophyses of \u003cem\u003eP. amboniensis\u003c/em\u003e and \u003cem\u003eP. bicoerulans\u003c/em\u003e at emergence can be described as a trade-off for 1) faster development of nymphs and 2) quick maturation of the emergent adults. These species have endophytic oviposition, where females deposit their eggs within plant tissues by means of a well-developed ovipositor. It is also noteworthy that larval behaviour and adult behaviour in these damselflies are intertwined, where, for instance, the site of oviposition and habitat for early larval development is chosen by the adult. This compensates for higher activity, which results in a higher growth rate but increases predation risk in damselfly larvae (M\u0026uuml;hlenhaupt et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and predicts developmental plasticity in changing environments such as drying pools (Pernecker et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This can be used to explain why adult detection probability differed from that of nymphs within a shared habitat.\u003c/p\u003e \u003cp\u003eThe co-occurrence of \u003cem\u003eP. amboniensis\u003c/em\u003e and \u003cem\u003eP. bicoerulans\u003c/em\u003e can be explained by differences in microhabitat use and behaviour as predicted by functional larval morphological traits. Co-occurring species exhibit a wealth of differing traits that allow them to partition the resources and reduce competition. Alteration of habitat has consequences on abundance (Dalzochio, P\u0026eacute;rico, Dametto, and Sahl\u0026eacute;n \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), and size structures of multiple co-occurring species (Oliviera-Junior, Teod\u0026oacute;sio, and Juen 2021). This affects species simultaneously and sometimes shifts in favour of species that have evolved in heterogeneous environments. The naiads of \u003cem\u003eP. bicoerulans\u003c/em\u003e were detected mostly on sites with slower water flows and higher abundances of emergent vegetation characterised by warm waters and low abundances of total dissolved solids. This microhabitat specialisation is probably the reason why \u003cem\u003eP. bicoerulans\u003c/em\u003e is easily displaced by competitive lowland species (Clausnitzer et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Nonetheless, the overall detection probability of naiads was lower than that of adults in the presence of \u003cem\u003eP. amboniensis\u003c/em\u003e. The adults of \u003cem\u003eP. amboniensis\u003c/em\u003e were high in areas with driftwood, while naiads were detected mostly where the stream was rocky and fast-flowing.\u003c/p\u003e \u003cp\u003eThis study also sheds light on stage-specific differences in the occupancy and detection probabilities of adults and larvae of co-occurring species. The sensitivity of Afromontane forests is due to their distinct vegetation types that are ecologically meaningful for the terrestrial adult Odonata (Clausnitzer et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Dijkstra and Clausnitzer 2006). However, owing to the complex life history traits that involve metamorphosis and use of different habitats for larvae and adults (Hogreve and Suhling \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Steenweg et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), stability of these populations can be regulated at different developmental stages (Lambert et al. 2023). Determining the demographic structure of such a population leads to understanding how each stage influences the other. Adults contribute to the recruitment of nymphs through laying eggs and selection of oviposition sites that enhance survival, while nymphs transform into adults through emergence. The presence of many adults on a site of oviposition means that a large number of nymphs will be recruited, while the emergence of a large number of larvae translates into adults. There is a difference in the means of dispersal between life stages. The adults fly to better microhabitats suitable for oviposition, while naiads have the ability to swim or drift away, especially downstream.\u003c/p\u003e\n\u003ch3\u003eImplications for conservation\u003c/h3\u003e\n\u003cp\u003eIt has been established that East African Montane Forests, Ethiopian Highlands, Eastern Arc, and Coastal Forest Mosaics are regions of conservation value due to the large number of threatened species in these regions (Clausnitzer et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Clausnitzer \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). The sensitivity and diversity of threatened species also coincide with the endemism patterns linked with habitat heterogeneity in the highlands (Dijkstra et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). The highly susceptible species are habitat specialists that occur in a small geographical range and have a limited area of occupancy. These include those that are confined to mountain peaks and those living in highly fragmented landscapes. Although the occupancy of \u003cem\u003eP. amboniensis\u003c/em\u003e and \u003cem\u003eP. bicoerulans\u003c/em\u003e in Mount Kenya is promising, these populations are separated from the closest populations located in the Aberdare Ranges the risk of continued habitat isolation means that even when a population is seemingly stable in a restricted range, it is still vulnerable to stochastic environmental events like extreme weather and genetic effects such as bottlenecks and inbreeding depression that hasten biotic attrition (Webster et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDragonflies and damselflies are agile organisms because they require more than one ecosystem to become mature adults. Changes incurred at the larval stage are reflected in the adult stage and vice versa. The expos\u0026eacute; that these species have invested in the evolution of functional traits in larvae is timely. Moving forward, it will be necessary to define stage-specific and species-specific threats and evaluate extinction probabilities. For instance, in reality, habitat loss is a broad term in the context of odonates and other freshwater invertebrates (Forister et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and fails to account for all life stages. It has been shown that even when protected areas are rarely designed with conservation of insects in mind (Chowdhury et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), conservation of threatened species can be achieved through conservation of targeted non-threatened species (Baker et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSpecies in the Afromontane streams are adapted to living in clear waters with little natural silting while species that occur downstream are more tolerant of human disturbance and natural silting from storms (Deacon and Samways \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The nymphs are critical for conservation because a great proportion of Odonata larvae do not emerge or develop to adulthood, and this number may rise with increased human pressure on these habitats caused by multiple factors (Kietzka et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These include soil erosion and siltation caused by deforestation and livestock grazing along riverbanks and invasive alien trees that affect freshwater systems through synergistic effects. When water conditions are good, removal of alien invasive trees can lead to the rapid recovery of dragonfly populations, as long as a source population still exists in the area (Samways \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2008\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe first- and second-level streams in Mount Kenya are being tapped heavily throughout the year for community water supply. Water abstraction affects abundances of many species groups, including fish (Merciai et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and dragonflies. Dynamics of water abstraction from small montane streams may prevent threatened species (such as \u003cem\u003eP. amboniensis\u003c/em\u003e and \u003cem\u003eP. bicoerulans\u003c/em\u003e) from becoming abundant and encourage generalist species. It could also result in fine partitioning of narrow-range endemic species in small Afromontane streams influenced by damming and siltation. Damming of waterways can locally impoverish riverine fauna, affect water flow, create lentic conditions in lotic systems, and incur long-term changes in odonate assemblages (Samways \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). These newly created spaces appear to favour occupancy by \u003cem\u003eP. bicoerulans\u003c/em\u003e but fail to guarantee that these microhabitats are suitable for the species long-term survival. For instance, when a dam is left to follow natural succession and becomes fully vegetated, it is likely to attract different generalist species. This reduces the abundance of specialists in favour of widespread generalists. Damming encourages colonisation of invasive predatory species such as clawed frogs (\u003cem\u003eXenopus borealis)\u003c/em\u003e, which has cascading effects on various biotas and neighbouring ecosystems through impact on dragonfly larvae. Similarly, the silt that accumulates on the ponds is often released into the stream and washed downstream. The effects of these recurrent small-scale events, though barely detected in the short term, affect local species co-occurrence and can alter community structure in the long term.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eThis study states a situation where two threatened species co-occur but utilise different microhabitats, which is underpinned in their functional morphological traits. These two distantly related damselfly species are a good case for comparative studies because their distinct morphological differences are evident and adequate to infer the evolutionary significance of life history and behaviour. Functional traits, when taken in the context of sympatric species, are an important tool for understanding differentiation or potential interaction between species. While this study is not adequate to decipher the historical pathways that influenced these evolutionary traits, it reveals adaptations to different microhabitats and the co-occurrence of damselflies in montane streams. The intimation of this conservation dilemma is ensuring habitat heterogeneity as a common ground for ecosystem restoration when multiple threatened species are involved.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch3\u003eAcknowledgements\u003c/h3\u003e\n\u003cp\u003eWe wish to thank Prof. Eucharia Kenya for her constructive comments and suggestions that substantially improved the quality of this research. Special gratitude goes to Samuel Thuita and Maria Dzine for their assistance. We wish to thank Dr. Peter Mosongo for the suggestions that greatly contributed to the quality of the original manuscript.\u0026nbsp;\u003c/p\u003e\n\u003ch3\u003eAuthor contributions\u003c/h3\u003e\n\u003cp\u003eConceptualisation: AMK, AWK, and MNG; methodology: AMK, IWN, and AWK; formal analysis: AMK, data collection: AMK, AWK, and MNG; data curation: AMK; writing\u0026mdash;original draft preparation: AMK and IWN; writing\u0026mdash;review and editing: AWK, MNG, and AWG; visualisation: AMK; supervision: AWK and MNG; funding acquisition: AMK, IWN, AWK, AWG, and MNG. All authors have read and agreed to the published version of the manuscript.\u0026nbsp;\u003c/p\u003e\n\u003ch3\u003eFunding\u0026nbsp;\u003c/h3\u003e\n\u003cp\u003eAMK was funded by the Mohamed bin Zayed Species Conservation Fund under the project \u0026lsquo;220529140: Research and Conservation of the Critically Endangered Kenya Montane Dancing Jewel (\u003cem\u003ePlatycypha amboniensis\u003c/em\u003e) in Mount Kenya.\u0026rsquo; AG was supported by the Centre for Ecosystem Restoration - Kenya.\u0026nbsp;\u003c/p\u003e\n\u003ch3\u003eData availability\u003c/h3\u003e\n\u003cp\u003eThe following supporting information: abundance, occupancy data, and AICc model results are available at (Zonedo). doi: 10.5281/zenodo.8101703\u0026nbsp;\u003c/p\u003e\n\u003ch3\u003eORCIDs\u003c/h3\u003e\n\u003cp\u003eA Karani \u003cimg width=\"15\" height=\"15\" 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alt=\"https://orcid.org/0000-0002-4766-6251 \"\u003e\u0026nbsp; https://orcid.org/0000-0002-4766-6251\u003c/p\u003e\n\u003ch3\u003eDisclosure statement\u003c/h3\u003e\n\u003cp\u003eThe authors declare no conflict of interest.\u0026nbsp;\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eBaker DJ, Garnett ST, O\u0026rsquo;Connor J et al (2019) Conserving the abundance of nonthreatened species. 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Ecol Evol 8:8930\u0026ndash;8939. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ece3.4428\u003c/span\u003e\u003cspan address=\"10.1002/ece3.4428\" 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":true,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"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":"occupancy probability, co-occurrence, larval morphology, functional traits, conservation, ecosystem restoration","lastPublishedDoi":"10.21203/rs.3.rs-6433576/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6433576/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe biodiversity crisis is exceptionally severe in the freshwater systems of the highly threatened Afromontane ecosystems. We studied functional traits of ecological significance, estimated abundance, probabilities of occupancy, and detection of adults and nymphs of Kenya Jewel (\u003cem\u003ePlatycypha amboniensis\u003c/em\u003e Martin, 1915) and Giant Sprite (\u003cem\u003ePseudagrion bicoerulans\u003c/em\u003e Martin, 1907) in Mount Kenya Forest. The estimates of abundance were based on replicated counts, while occupancy and detection probabilities were estimated using a single-season, two-species occupancy formulation. The study revealed that detection probabilities of one species were influenced by the detection probabilities of the other, but conversely, occupancy was not. This was supported by morphological traits. The larvae of \u003cem\u003eP. amboniensis\u003c/em\u003e is adapted to rocky and fast-flowing lotic streams, while \u003cem\u003eP. bicoerulans\u003c/em\u003e is adapted to vegetated, littoral, and slow-moving reaches of the same streams. The study concluded that while these species have different adaptations, their co-occurrence depends on the heterogeneity of the microhabitats and recommended ecosystem restoration approaches that maintain habitat complexity to increase the resilience of co-occurring species to future environmental changes.\u003c/p\u003e \u003cp\u003e \u003cb\u003eImplications for conservation\u003c/b\u003e: The occurrence of Kenya Jewel was not influenced by occurrence of Giant Sprite and; there was neither competition nor niche overlap. This aligns with the differences in ecological adaptations based on functional morphology of adults and larvae. This evidence is useful for restoration of the ecosystem to ensure that these species are conserved concurrently in their range of co-occurrence.\u003c/p\u003e","manuscriptTitle":"Congruence of detection probabilities and co-occurrence of threatened Afromontane damselflies with diverging functional traits (Odonata: Chlorocyphidae, Coenagrionidae)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-04-28 04:48:59","doi":"10.21203/rs.3.rs-6433576/v1","editorialEvents":[{"type":"communityComments","content":2},{"type":"decision","content":"Revision requested","date":"2025-10-22T12:05:55+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-15T09:45:25+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-06-24T12:43:33+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"82051994772461842719738615725781161741","date":"2025-05-16T19:52:04+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"245922593647105353247735349502762878499","date":"2025-05-11T08:41:09+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-05-11T08:12:28+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-04-12T13:02:18+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-04-12T13:00:20+00:00","index":"","fulltext":""},{"type":"submitted","content":"Journal of Insect Conservation","date":"2025-04-12T09:19:18+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":"31343603-52f9-477f-9261-248bf45ae457","owner":[],"postedDate":"April 28th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-02-09T16:04:45+00:00","versionOfRecord":{"articleIdentity":"rs-6433576","link":"https://doi.org/10.1007/s10841-026-00750-7","journal":{"identity":"journal-of-insect-conservation","isVorOnly":false,"title":"Journal of Insect Conservation"},"publishedOn":"2026-02-05 15:58:15","publishedOnDateReadable":"February 5th, 2026"},"versionCreatedAt":"2025-04-28 04:48:59","video":"","vorDoi":"10.1007/s10841-026-00750-7","vorDoiUrl":"https://doi.org/10.1007/s10841-026-00750-7","workflowStages":[]},"version":"v1","identity":"rs-6433576","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6433576","identity":"rs-6433576","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}