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Patterns of Odonata Assemblages in Lotic and Lentic Systems in the Ankasa Conservation Area, Ghana

Patterns of Odonata Assemblages in Lotic and Lentic Systems in the Ankasa Conservation Area, Ghana Hindawi International Journal of Zoology Volume 2019, Article ID 3094787, 14 pages https://doi.org/10.1155/2019/3094787 Research Article Patterns of Odonata Assemblages in Lotic and Lentic Systems in the Ankasa Conservation Area, Ghana 1 2 1 Issah Seidu , Collins Ayine Nsor , Emmanuel Danquah , 1 1 Paul Tehoda, and Samuel K. Oppong Department of Wildlife and Range Management, Faculty of Renewable Natural Resources, Kwame University of Science and Technology, Kumasi, Ghana Department of Ecotourism and Forest Recreation, Kwame Nkrumah University of Science and Technology, Kumasi, Ghana Correspondence should be addressed to Issah Seidu; antwiseidu88@gmail.com Received 5 November 2018; Accepted 15 January 2019; Published 10 February 2019 Academic Editor: Marco Cucco Copyright © 2019 Issah Seidu et al. is Th is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Our study examined Odonata assemblages distribution pattern and the predictive factors that accounted for this in the lotic and lentic water systems within the Ankasa Conservation Area (Ghana). A total of 23 sites with sampling protocol of 2 researchers per hour per sampling site were used to survey Odonata species over two seasons in the three water bodies (streams, rivers, and ponds). Broken stick model, individual-based rarefaction, and Renyi diversity ordering were employed to quantify community assemblages. Ordination technique was also used to determine the Odonata-environmental relationship. A total of 1403 individuals, belonging to 47 species (22 Zygoptera and 25 Anisoptera) in six families, were recorded. Species richness (Hc =3.414, p =0.169) and diversity (Hc =1.661, p = 0.44) generally did not dieff r among the three water systems. However, from individual sites, ponds appeared mostly diverse (𝛼 -scale = 0.04, Renyi index (r) = 5.86 to 𝛼 = 3.5, r = 3.12), in spite of their lowest species abundance and richness. At the suborder level, ponds equally exhibited the highest Anisoptera species richness (9.90 ± SE 0.640) compared with Zygopterans (0.80± SE 0.291). Overall, Anisopterans (K=16.51, p= 0.00026) and Zygopterans richness (K=16.39, p= 0.00023) differed significantly among the three subsystems, while Odonata composition also differed significantly among the various water bodies (ANOSIM: global R=0.94, p<0.001). Flow rate, water temperature, channel width, and turbidity were the key predictive factors that influence the structure of Odonata species assemblages. The results highlight the need to improve the functional status of the lentic and lotic systems, with the ultimate goal of conserving diverse Odonata fauna and other sympatric freshwater biodiversity. 1. Introduction Freshwater habitats present two major differing water systems, lotic (running) and lentic (standing) waters, which Freshwater habitats cover only 1% of the total earth sur- differ in their environmental and spatiotemporal settings [5]. face and contain 10% of the earth biodiversity [1]. Their They are distinguished by physicochemical parameters of the importance in sustaining biodiversity and human welfare is water such as turbidity, organic matter, pH [6], dissolved oxy- undeniable. Freshwater resources are the major sources of gen [7], nutrients content [8], and flow regimes. These water livelihood to Afrotropical rural and periurban folks [2]. They systems together support heterogeneous environment which provide water supplies for human consumption, industrial provide favorable conditions for both vertebrate and inverte- utilization, and ecosystems support for sher fi ies and other brates communities including the amphibious Odonata taxa. aquatic biodiversity. However, wetlands are considered one Odonata are denizen of freshwater environments such of the most jeopardized ecosystems in the world [3]. Many wetlands worldwide are experiencing dramatic anthropic as rivers, lakes, ponds, wetlands, and, to some extent, phy- totelmata and brackish water resources [9, 10]. They play change, mostly for agricultural purpose [4]. Generally, these changes are associated with abiotic conditions which are significant role in freshwater ecosystem functioning, acting as both prey (fed by vertebrates and other large insects) normally not found in nature with cascading impacts on residence aquatic biota. and top predators (feeding on smaller insects in vertebrate 2 International Journal of Zoology free aquatic environment) [11]. Due to the reliability of both July and September to November, with average annual rainfall larval and adult Odonata to specicfi water conditions for of 1700 to 2000 mm [22]. survival [12], and their sensitivity to habitat disturbances, they are eeff ctively used asindicatorsof water quality [13, .. Description of Sampled Water Types in the Study Area. 14]. Odonata, therefore, serve as an umbrella species in Stream (𝑛 = 7): We located all the sampling sites along biodiversity conservation [15] and represent specific biotic the Asufia stream and a stream which is tributary to the wetland assemblages. Ankasa River (Figure 1). Three sites were laid along the Asuafi Sustainability of Afrotropical freshwater resources and stream, while four were located on the other stream. The sites their associated Odonata fauna requires knowledge of the were characterized by sandy substrate. The channel width contribution of different water bodies in particular ecosys- ranged from 1 m to 1.9 m while the depth was from 0.1 m to tems. Theseinclude knowledgeabout thespecies richness, 0.21 m. The water was o fl wing rapidly through dense canopy diversity, and community structure in different water types, cover, with the trees and shrubs being the dominant bank the variability of water systems across the landscape, and vegetation. the net contribution of these water systems to the catchment River (𝑛 = 6): All the sites were located along two biodiversity [16]. In general, such information is practically major rivers (Ankasa and Bonwere River) in the Ankasa scanty worldwide but particularly in West Africa. This is Conservation Area (Figure 1). Three sites were laid along the result of traditional Odonata research being geared each river, representing the total sampling sites. The Ankasa towards specific water body. For example, especially in and Bonwere Rivers are characterized by rocky and sandy Ghana, most current research on Odonata assemblages has substrates. All sites were associated with rapids and highly virtually focused on rivers (see [17–19]), and streams [19, 20] oxygenated, cold water. The channel width was between 2 m with little or no studies describing other natural and artificial and 15 m while the depth ranged from 0.3 m to 0.75 m. The lentic freshwater systems such as ponds, pools, and lakes, sites were laid adjacent to intact secondary forest vegetation although these water bodies are well known to harbor diverse with the margins mainly composed of trees and shrubs, and Odonata fauna and higher Odonata richness elsewhere [21]. small patches of various grasses (Poaceae). The water bodies In order to contribute to the initial understanding of the also pass through dense canopy with low sun exposure except importance and the influence of dieff rent water types on in sun flecks caused by tree falls. the Ghanaian Odonata biodiversity, differences in Odonata Pond (𝑛 = 10): All the 10 ponds were naturally permanent assemblage structure of lotic and lentic systems were investi- water bodies located in and outside the Ankasa Conservation gated. We hypothesized that adult Odonata composition will Area. Four ponds were located in the forest reserve, while be signicfi antly dieff rent among the water types due to their six were outside the forest adjacent to cultivated rubber, preference for different water bodies [12, 13]. Accordingly, we vegetables, and cocoa plantation which were mostly used for addressed two major questions: (1) are there any signicfi ant irrigation by the local communities (Figure 1). The bottoms differences in Odonata abundance, richness, and community were mainly composed of mud/clay and organic matter. Most composition between the lotic (rivers and streams) and lentic of the ponds were surrounded by partial vegetation structure systems (ponds)? and (2) are there signicfi ant differences in with high amount of sun penetration. The dominant plant the abundance and richness of Anisopterans and Zygopterans families in the marginal zones were Cyperaceae and Poaceae. among the water types? In order to address these questions, Pondssurfaceswereassociated with stands ofemergent or we compared adult Odonata assemblages occurring in 7 sites floating vegetation which were utilized by the adult Odonata along two major rivers, 6 sites along three different streams, for perching. and 10 different ponds found in and outside the Ankasa Conservation Area. .. Odonata Sampling Procedures. We sampled adult indi- viduals of all Odonata species at 23 sites with a sampling protocol of 2 researchers per hour per sampling site, along 2. Materials and Methods the three different water types, Rivers, Streams, and Ponds ∘ 󸀠 ∘ .. Study Site. Ankasa Conservation Area (5 17 Nand 2 in the Ankasa Conservation Area. We sampled simulta- 39 W) is a twin Protected Area comprising Nini-Suhien neously, collecting and noting the species occurring, and their abundances in each sampling site until no new species National Park and the Ankasa Resource Reserve [22]. It is about 500 km situated in the Western Region of Ghana, were encountered for approximately one hour for each visit. and the only area in the Wet Evergreen Forest [23]. Ankasa Sampling was done from January, 2017, to March, 2017, for Conservation Area is designated as a Globally Significant thedry season whilethe wetseason sampling took place Biodiversity Area (GSBA) and Important Bird Area (IBA) from May, 2017, to July, 2017. We sampled all adult Odonata [23]. during the day between the hours of 9 am and 5 pm. We Ankasa Conservation Area presents an ideal ecosystem captured all adult Odonata individuals where possible, using forthisstudy, asitboasts ofa signicfi ant number of complex a hand net. We identified each specimen to species level in situ, using Dijkstra, and Clausnitzer, [25] identifiction keys. and diverse freshwater systems including riverine, streams, and ponds. These wetlands and their associated forest envi- Where identification of some species was not possible on ronmentsupport themostbiological diversity of any kind the eld fi , we photographed them and then used the African Dragonflies and Damselflies Online database (ADDO) [26], in Ghana [22]. The climate of the area is characterized by a distinctive bimodal rainfall pattern occurring from April to for subsequent idenitificantion. International Journal of Zoology 3 ∘   ∘   ∘   ∘   ∘   ∘   ∘   2 44 10 W 2 41 40 W 2 39 10 W 2 36 40 W 2 34 10 W 2 31 40 W 2 29 10 W River Stream Roads Towns Ankasa CA Sampling sites Ponds River Kilometers Stream 0 1.25 2.5 5 7.5 10 Figure 1: Map of the study area in and near the Ankasa Conservation Area, Western Region, with the situation of the 23 sampling sites of the three water types. .. Measurement of Biophysical Variables. We recorded abi- Allmultivariate analyses weredoneusing PRIMER 6.1.5 otic variables concurrently during the Odonate sampling, to package [30]. assess their influence on Odonate community structure. Sur- face water temperature ( C), pH, dissolved oxygen (mg/L), ... Species Abundance Distribution (SAD) for Odonata turbidity, conductivity, altitudes, flow rate, channel width Species. The application of species abundance distribution and depth, aquatic vegetation, substrate type, and bankside models in the study of species patterns has been widely vegetation were all measured in all sampling sites following used in community ecology by most scientists [31], as well Seidu et al. [17] procedure. as measuring the impact of disturbance on community structure [32]. In this study, Odonate abundance as a measure .. Data Analysis. We first tested the normality of the abun- of diversity was quantified using rank abundance model [33]. dance data set using Shapiro-Wilk test [27]. The abundance In each site, we listed the number of Odonata species for all of the wet and dry seasons, say 𝑆 data was log(X+1) transformed prior to analysis. Bray-Curtis , represented by one similarity indices and nonparametric multidimensional scal- individual, and the number of species, say 𝑆 , represented ing (NMDS) were used to determine relationships of species by K individuals, where K denotes the abundance of the composition among the sampling sites of the various water most abundant species and 𝑆 + ... + 𝑆 = S [34]. 1 𝐾 bodies. To test for the significant difference in species com- Accordingly, the sequence of relative frequencies 𝑓 = 𝑆 /S (r 𝑟 𝑟 position among the various water types, we employed one- =1...K) constitutes a frequency distribution for the number way analysis of similarities with 999 permutations (ANOSIM; of individuals per species which is usually referred to as the [28, 29]), with Bray-Curtis similarities as dependent and the species-abundance curve [34]. We then tfi ted the MacArthur three different water types (streams, rivers, and ponds) as broken stick model (BS) [35, 36] in the species abundance independent factor. Similarity percentage analysis (SIMPER) data, using the regression model approach [35] to determine routine in primer [30] was used to determine average dis- the pattern of species communities in each of the freshwater similarity between the water bodies and the various species systems. MacArthur [36] suggested that the niche space could contributing to the most similarity within each water body. be compared to a stick of length 1, where n – 1 points would ∘   ∘   ∘   ∘   ∘   ∘   5 10 50 N 5 13 20 N 5 15 50 N 5 18 20 N 5 20 50 N 5 23 20 N 4 International Journal of Zoology randomly generate n segments of lengths proportional to the suborders (Anisoptera and Zygoptera) for streams, rivers, and number of individuals of each species in the community, ponds. given as Renyi [40] extended the concept of Shannon’s entropy [41], by defining the entropy of order 𝛼 (𝛼≥ 0, 𝛼 ≠ 1) of a probability distribution (p , p ...p ). Diversity prolfi e values 𝑁 1 1 2 s 𝑛 = ∗ ∑ (1) (H-alpha) were calculated from the frequencies of each com- 𝑆 𝑛 𝑖=1 ponent species (proportional abundances pi =abundance of species i/ total abundance) and a scale parameter (𝛼 )ranging (see [36]) Where 𝑛 represents the number of individuals of from zero to infinity as the species i; N represents the total number of individuals; and S represents the total number of species in the commu- 𝑠 (log ∑ 𝑝𝑖 ) 𝑖−1 nity. (3) (𝐻 )= (1− 𝛼 ) Thismodel approach wasused in order to test against the null hypothesis (𝐻 ) that species abundance distribution and (See [42]). Odonate abundance, richness, and diversity richness did not differ in each of the three water systems. ordering were performed using PAST version 3.06 sowa ft re All the species in each of the sampling sites per water type package [43], which provides robust algorithm as shown in were ranked from the most to the least abundant on the rank Krebs et al. [44]. abundant curve [37]. Each species rank is plotted on the x- Due to the nonnormal nature of the data set, a Kruskal- axis, and the abundance is plotted on the y-axis. Wallis test was applied to test for the differences in Odonata With the broken stick model, if a log scale is used for and suborders (Anisoptera and Zygoptera) abundance and abundance, the species exactly fall along a straight line, richness among the 23 sites, using PAST version 3.12 [43]. according to the model equation log𝐴= 𝑏 +𝑏 𝑅 ,where 𝑜 1 Homogeneity of species variance among sample plots was Aisthe speciesabundance, R is the respective rank, and evaluated, using Levene test [45], defined as 𝑊 = ((𝑁−𝑘)/(𝑘− b and b are optimized tfi ting parameters [32]. Analysis of 𝑘 𝑘 2 2 ́ ́ ́ 1))(∑ 𝑁𝑖( Z𝑖− Z𝑖) /∑ ∑ (𝑍𝑗−𝑖 Z𝑖) ) where 𝑗𝑍𝑖 can 𝑖=1 𝑖=1 𝐽=1 covariance (ANCOVA) was applied to test for the significant have one of the following three definitions. difference of the slope of the SADs for the three water types, ́ ́ ́ 2 𝑗=𝑍𝑖 ⌈𝑌 −Y ⌉ where Y is mean of the 𝑖𝑡ℎ subgroup; Y is 𝑖 𝑖 𝑖 while Pearson’s Chi-square test (𝜒 ) was applied to determine the median of the 𝑖𝑡ℎ subgroup, and, finally, 𝑗=𝑍𝑖 ⌈𝑌 − Y 𝑖⌉ , whether an observed distribution along the goodness of tfi statistically differed in the BS model. Among the four notable where Y 𝑖 is the 10% trimmed mean of the 𝑖𝑡ℎ subgroup. Z𝑖 SAD models (i.e., geometric, log series, log normal, and BS), are the group means of the 𝑗𝑍𝑖 and Z is the overall mean of the BS model is the only one that fundamentally describes the the 𝑗𝑍𝑖 . process of niche partitioning in a community where species exhibit continuous nonoverlapping niches [33]. .. Environmental Predictors of Odonata Distribution. We Individual-based rarefaction techniques [38] were used determined the relationships between the abiotic variables to compare Odonate richness across the three water sys- recorded and the species occurrence in the various water tems (rarefaction curves). Rarefaction curves are created by bodies using a canonical correspondence analysis (CCA, randomly resampling the pool of N samples multiple times [45]). We used the Environmental Community Analysis and then plotting the average number of species found in (ECOM.exe) version 1.4 packages [46] to perform the CCA each sample (1,2 .. . 𝑁) [24]. us, Th rarefaction generates analysis. The significance of the rfi st two axes generated the expected number of species in a small collection of n in the analysis was validated through the Monte Carlo test individuals (or n samples) drawn at random from the large (using 5000 iterations) [47]. Environmental variables utilized pool of N samples. The rarefaction curve 𝑓 is defined as in the CCA were water temperature, dissolved oxygen, pH, turbidity, conductivity, o fl w rate, and channel width and −1 𝑁 𝑁− 𝑁𝑖 depth. CCA is a direct method of ordination with the (2) 𝑓 =𝐸[𝑋 ]=𝐾 − ( ) ∑ ( ) 𝑛 𝑛 resulting outcome being the variability of the environmental 𝑛 𝑛 𝑖=1 data, aswell asthe variability ofspecies data [48]. (See [38]). Where 𝑋 = the number of groups still present in the subsample of “n”lessthan 𝐾 whenever 3. Results at least one group is missing from this subsample, 𝑁= total number of items, 𝐾= total number of groups, 𝑁𝑖 = .. General Pattern of Odonata Composition and Abundance total number of items in group 𝑖 (𝑖 = 1,... 𝑘) [24, 39]. u Th s, Distribution across the Streams, Rivers, and Ponds. Atotal the linear model for the BS was tfi ted for each rarefied rank of 1403 adult Odonata specimens belonging to 47 species, in order to build the 95% cond fi ence limits for the slopes of and six families, were registered in streams, rivers, and all sampling sites. ponds in the study area (Tables 1(a) and 1(b)). Of the 47 Rarefaction methods, both sample based and individual species recorded, 22 Zygoptera species belonging to four based, allow for meaningful standardization and compari- families (Calopterygidae, Chlorocyphidae, Coenagrionidae, son of datasets [24]. We compared the estimated Odonata and Platycnemididae) and 25 Anisopterans from two families abundance and species richness, as well as the estimated (Aeshnidae and Libellulidae) were recorded (Tables 1(a) and abundance and number of species belonging to the respective 1(b)). Libellulidae was the dominant family with 13 species, 𝐼𝐽 𝐼𝐽 𝑁𝐼 International Journal of Zoology 5 Table 1 (a) Checklist and abundance of Zygoptera (damselflies) species recorded in streams, rivers, and ponds in the Ankasa Conservation Area. Species that o ccurred exclusively in streams are represented by (∗), exclusively in rivers (#), and exclusively in ponds (!). Species shared between streams and rivers are represented by (∗#), between streams and ponds (∗!), and between rivers and ponds (#!) Family Zygopterans Stream River Ponds Total Calopterygidae Phaon camerunensis Sjosted ¨ t, 1900∗ 19 0 0 19 Phaon iridipennis (Burmeister, 1839)∗#6 16 0 22 Sapho bicolor Selys, 1853∗ 20 0 2 Sapho ciliata (Fabricius, 1781)∗#39 10 0 49 Umma cincta (Hagen in Selys, 1853)∗ 20 0 0 20 Chlorocyphidae Chlorocypha luminosa (Karsch, 1893)∗#33 19 0 52 Chlorocypha radix Longfild, 1959 ∗#16 8 0 24 Chlorocypha selysi Karsch, 1899∗#10 29 0 39 Coenagrionidae Agriocnemis exilis Selys, 1872! 0 0 4 4 Agriocnemis zerafica Le Roi, 1915! 0 0 13 13 Ceriagrion corallinum Campion, 1914∗!8 0 13 21 Ceriagrion glabrum (Burmeister, 1839)! 0 0 26 26 Ceriagrion rubellocerinum Fraser, 1947∗#7 6 0 13 Pseudagrion hamoni Fraser, 1955∗#3 9 0 12 Pseudagrion isidromorai Sart, 1967∗ 15 0 6 Pseudagrion kersteni (Gerstack ¨ er, 1869)∗ 50 0 5 Pseudagrion melanicterum Selys, 1876∗#23 24 0 47 Pseudagrion hamoni Fraser, 1955∗ 26 0 8 Pseudagrion sjoestedti F¨orster, 1906 02 0 2 Platycnemididae Mesocnemis singularis Karsch, 1891# 0 26 0 26 Elattoneura balli Kimmins, 1938∗#38 16 0 54 Elattoneura villiersi (Fraser,1948)∗ 34 3 0 37 Total number of individuals Total number of species (b) Checklist and abundance of Anisoptera (dragonflies) species recorded in streams, rivers, and ponds in the Ankasa Conservation Area. Species that occurred exclusively in streams are represented by (∗), exclusively in river (#), and exclusively in pond (!). Species shared between streams and rivers are represented by (∗#), between streams and ponds (∗!), and between rivers and ponds (#!) Family Anisopterans Stream River Ponds Total Aeshnidae Gynacantha bullata Karsch, 1891∗ 50 0 5 Gynacantha cylindrata Karsch, 1891∗ 10 0 1 Libellulidae Acisoma inflatum Selys, 1882! 00 147 147 Aethriamanta rezia Kirby, 1889! 0 0 33 33 Chalcostephia flavifrons Kirby, 1889! 0 0 90 90 Cyanothemis simpsoni Ris, 1915# 0 9 0 9 Eleuthemis buettikoferi Ris, 1910# 0 9 0 9 Neodythemis klingi (Karsch, 1890)∗ 14 0 0 14 Micromacromia zygoptera (Ris, 1909)∗ 1000 10 Olpogastra lugubris (Karsch, 1895)#! 0 3 22 25 Orthetrum austeni (Kirby, 1900)! 0 0 19 19 Orthetrum julia Kirby, 1900∗!7 0 13 20 Orthetrum microstigma Ris, 1911! 0 0 6 6 Orthetrum stemmale (Burmeister, 1839)∗!6 0 7 13 Orthetrum trinacria (Selys, 1841)! 00 6 6 Palpopleura lucia (Drury, 1773)! 0 0 81 81 Palpopleura portia (Drury,1773)! 0 0 71 71 Pantala flavescens (Fabricius, 1798)! 0 0 5 5 6 International Journal of Zoology (b) Continued. Family Anisopterans Stream River Ponds Total Rhyothemis notata (Fabricius, 1781)#! 0 3 100 103 Rhyothemis semihyalina (Desjardins, 1832)! 0 0 38 38 Trithemis aconita Lieinc ft k, 1969∗!3 0 17 20 Trithemis arteriosa (Burmeister, 1839)#! 0 9 86 95 Trithemis bifida Pinhey, 1970#! 0 3 2 5 Trithemis dichroa Karsch, 1893#! 0 3 43 46 Urothemis edwardsii (Selys, 1849)#! 0 3 28 31 Total number of individuals Total number of species Table 2: Results of the broken stick model for the abundance rank distribution of Odonata species, calculated for each of the three water types. Sample Intercept ±S.E. Slope ±S.E. R Prob. Streams 4.05 ± 1.75 0.55 ± 0.19 0.37 0.009 Rivers 2.96 ± 1.19 0.26 ± 0.09 0.38 0.008 Ponds 25.27 ± 5.41 -1.02 ± 0.43 -0.33 0.02 Slope of SAD: 𝐹 = 6.22, p (regr): 0.002 2,138 (ANCOVA interactions x species rank) Monte-Carlo Permutation (n = 99999): p<0.0014 Levene test for homogeneity of variance:p<0.0015 However, from three water types, we observed Odonate followed by Coenagrionidae (n =12) andCalopterygidae (n = 4), in rivers and ponds. Community assemblages across abundance in ponds to be the highest (n =870), but their the three sites were ranked from the most abundant to the spatial distribution did not differ significantly along the least abundant (Figure 2). Their abundance distribution fitted slopes of the curve (𝜒 P = 25.07, P = 0.24). Similar abundance well in the broken stick distribution (BS) model and generally and distribution trends were observed in streams (n =312, showed significant difference in the slopes of the three water 𝜒 P =7.12, P = 0.99) and rivers (n = 221, 𝜒 P = 4.11, P = systems (𝐹 = 6.22, p(regr) = 0.002, ANCOVA interactions 0.99) (Table 2, Figure 2). Individuals per sample site, in ponds 2,138 xspecies rank)(Table 2,Figure 2). Further Monte Carlotest (87.00 ± SE 8.83), streams (44.6± SE 4.4), and rivers (36.8 ± SE (n = 99999) revealed signicfi ant dieff rence in SAD slopes ( p 4.23), equally followed similar trend (Hc =16.72, P = 0.0002, = 0.001). Kruskal-Wallis test)(Figure 3). Pairwise comparisontest showed a significant difference between ponds and streams At the suborder level, streams had the greatest mean (P = 0.003) and ponds and rivers (P = 0.004). However, Zygopterans abundance (38.0± SE 4.29) (e.g., E. balli =54, C. there was no significant difference in Odonata abundance luminosa =52, and S. ciliata =49),compared with Anisopter- between rivers and streams (P > 0.05). Comparison of the ans (6.57± SE 2.05). Conversely, the ponds exhibited the SADs for the three water systems helps to distinguish a greatest Anisoptera abundance (81.40± SE 8.264) (e.g., A. specific habitat quality, in relation to its influence on Odonate inflatum =147, R. notata =103, and T. arteriosa =95) while abundance, while the shape of the rank abundance curve zygopterans were the least abundant (5.60± SE 1.96) (Table 3 generally revealed differences in Odonate dominance and and Figure 4). Sapho bicolor and P. sjoestedti represented by evenness from individual habitats, and which reflects in their double individuals (doubleton) and Gynacantha cylindrata, relative tolerance to disturbances. single individual (singleton), T. bifida (𝑛 = 5) and G. bullata (n = 5), were the least dominant Zygopterans and Anisopterans, respectively, in the study area (Tables 1(a) .. Comparison between Zygopterans and Anisopterans Spe- and 1(b), Figure 2). There was a significant difference in cies Richness among the Water Types. Ponds exhibited the the abundance of Zygopterans (K =16.5, p = 0.00025) and highest Anisoptera species richness (9.90± SE 0.640) but Anisopterans (K = 16.28, p= 0.0003) among the three sites. the lowest number of Zygopterans (0.80± SE 0.291) (Fig- Zygopteran abundance in ponds differed signicfi antly in ure 5). The streams had the highest Zygopteran richness pairwise comparison with streams (p= 0.0007) and rivers (p= (7.57± SE 0.481) but exhibited almost similar Anisoptera 0.0018) but showed no difference between rivers and streams species richness (2.0± SE 0.577) with rivers (1.8± SE 1.014) (p= 0.174). Similarly, the Anisoptera abundance in ponds (Figure 5). Kruskal-Wallis test showed a significant differ- varied significantly in the pairwise comparison with streams ence in Zygoptera species richness (K=16.39, p= 0.0002) (p= 0.0007) and rivers (p= 0.001) but no significant difference and Anisoptera richness (K=16.51, p= 0.0003) among the occurred between the rivers and streams (p = 0.825). water types. Pairwise comparison test showed a significant International Journal of Zoology 7 2.0 2.0 3.0 1.5 1.5 2.5 1.0 1.0 2.0 0.5 0.5 1.5 Rivers 0.0 0.0 1.0 Streams Ponds −0.5 −0.5 0.5 −1.0 −1.0 0.0 Species rank order Species rank order Species rank order Figure 2: Broken stick model for Odonata rank abundance distribution across the three water types in Ankasa Conservation Area. Abundance is based on cumulative values per species test sites. Notice that SADs are ordered in decreasing magnitude and plotted against the corresponding rank order. Table 3: Canonical coefficients and the correlations with the first three axes of the environmental variables of the canonical correspondence analysis (CCA) for the three water types. Interset correlations were significant ( p <0.05∗) for the three axes. Axis I Axis II Axis III Correlation 0.573∗ 0.098 -0.156 Turbidity 0.549∗ 0.081 0.114 Flow rate Width -0.706∗ -0.076 0.164 -0.260 -0.009 0.114 Depth -0.421 0.054 -0.092 Conductivity 0.203 0.079 0.145 Do Temperature -0.748∗ -0.060 0.070 0.154 -0.137 0.091 pH Canonical Eigen value 0.651 0.445 0.185 22.3 15.29 6.338 % variance explained 22.3 37.6 43.96 Cumulative % variance Pearson correlation 0.979 0.708 0.781 species/environment scores Kendal rank correlation of 0.684 0.463 0.597 species/environment scores difference in Zygopteran richness between ponds and streams However, mean species richness per sample site was rather (p= 0.0006), and between rivers and ponds (p= 0.001), the highest in ponds (10.7 ± SE 0.56), while rivers had the least but no difference existed between streams and rivers ( p= number (8.7 ± SE 0.92) (Figure 7). Homogeneity of species 0.56). Similarly, Anisoptera species richness in ponds differed variance among the three water systems differed significantly significantly with streams ( p= 0.00071) and rivers (p= 0.001), (p<0.0002, Levene test)(Table 2). but there was no significant difference between streams and Observed trends in Odonate structural assemblages (i.e., rivers (p= 0.82). abundance, evenness, and richness) reflected in the Renyi diversity ordering (from higher to lower indices; along an .. Trends in Odonata Richness and Diversity in the ree increasing alpha scale values) (Figure 8). Overall, Odonate Water Systems. Interpolating the SADs across the streams, diversity did not differ significantly ( Hc =1.661, p = 0.44) rivers, and ponds, with sample-based rarefaction, revealed across the three water types. However, from individual sites, that Odonate richness among the three systems was not we observed that Odonates from ponds appeared mostly diverse (𝛼 -scale = 0.04, Renyi index (r)= 5.86 to 𝛼 =3.5, r = signicfi antly dieff rent ( Hc =3.414, p =0.169, Kruskal-Wallis test) (Figure 6) and did not follow similar pattern observed 3.12), in spite of their lowest species abundance and richness in individual abundance. Chao-1 estimated species richness (Figure 5). This was linked to the shallower SAD curve for the three sites showed streams to be the highest (n = observed in Figure 2. u Th s, species abundance distributions, 24.33), followed by ponds (n =23) andrivers (n = 22). with shallower curve, tended to be highest in diversity, while log Abundance 0.0 2.5 5.0 7.5 10.0 12.5 15.0 17.5 20.0 22.5 25.0 log Abundance 0.0 2.5 5.0 7.5 10.0 12.5 15.0 17.5 20.0 22.5 log Abundance 0.0 2.5 5.0 7.5 10.0 12.5 15.0 17.5 20.0 22.5 8 International Journal of Zoology Hc = 16.72, P = 0.0002, Kruskal Wallis test Stream River Pond Water type Figure 3: Mean Odonata species abundance among the various water types. Stream River Pond Water type Zygoptera Anisoptera Figure 4: Comparison of mean species abundance of Zygopterans and Anisopterans among the water types. those with steeper curves were less diverse (Figure 6). Species luminosa (16%). Cholorocypha selysi (26%), P. melanicterum from the riverine systems were the least diverse and ranged (19%), and M. singularis (13%) contributed most to similarity from 𝛼 = 0.04, r =5.83to 𝛼 =3.5, r = 3.08 and were found at the in river community, whereas T. arteriosa (16%), A. inflatum bottom of the Renyi index curve (Figure 6). Odonate diversity (15%), and P. lucia (14%) were greatest contributing species in in streams (𝛼 = 0.04, r =5.84 to 𝛼 =3.5, r = 3.07) could barely pond communities. be distinguished from those in the riverine systems, as their The species composition of Odonata dieff red signicfi antly curves were spatially similar. between the various water bodies (ANOSIM: global R= 0.94, p<0.001). Pairwise comparison test showed a significant difference in species composition between rivers and ponds .. Similarity in Odonata Composition among Streams, (R= 0.98, p= 0.002). Also, streams revealed weak significant Rivers, and Ponds. The Nonparametric Hierarchical Cluster dieff rence with rivers ( R= 0.52, p= 0.02) but higher significant analysis of species occurrence showed vfi e different clusters difference with ponds ( R=0.99, p= 0.001). (P8, P5, P1, P2, P3, P7, P9, P6, P4, and P10), (R5 and R6), (S4, R2, R1, and S6), (S1 and S2), and (S3, S5, R4, R3, and R7) at 40% similarity index (Figure 9). The species .. Environmental Predictors of Odonata Structural Dis- tribution and Diversity. Canonical correspondence analysis occurrence in ponds showed a strong significant separation from streams and rivers communities. However, the sampling (CCA) showed the overall relationships between species sites of stream and river were ecologically less distinct and distribution and the biophysical variables recorded (Table 3, showed a higher species overlap with each other (Figure 9). Figure 10). Among the eight biophysical variables initially The Similarity Percentage (SIMPER) analysis revealed a included in the analysis, only four biophysical variables, similar trend, suggesting that streams and ponds (98.72%) namely, o fl w rate, water temperature, channel width, and and rivers and ponds (93.87%) exhibited greatest average turbidity, were shown to strongly inu fl ence the structure dissimilarity in species composition to one another. Streams of species assemblages. Species assemblages along the rfi st and rivers (67.31%) were relatively similar to each other axis correlated significantly with water temperature (r =- in Odonata species composition. SIMPER also revealed an 0.74, p<0.05), channel width (r = -0.70), o fl w rate ( r = 0.54, average similarity within the streams (49%), rivers (43%), and p<0.05), and turbidity (r = 0.57, p<0.05) (Table 3, Figure 10). ponds (63%). Species contributing most to similarity in the CCA axes 1 and 2 jointly explained 37.6% of the total variation stream community were E. balli (23%), S. ciliata (17%), and C. in species structural distribution and diversity among sites. Mean species abundance Mean species abundance International Journal of Zoology 9 Stream River Pond Water type Zygoptera Anisoptera Figure 5: Comparison of mean species richness of the Zygopterans and Anisopterans among the water types. Hc = 3.414, p = 0.169, Kruskal- 15 Wallis test Chao-1 estimate: Streams: 24.33 Rivers: 22 Ponds: 23 100 200 300 400 500 600 700 800 Odonata specimens Figure 6: Standardized comparison of Odonata richness for individual-based rarefaction curves. The data represent summary counts of Odonates that were recorded from the three water types in Ankasa Conservation Area. eTh red, blue, and green lines are the rarefaction curves, calculated from (2) [24], with a 95% confidence interval. eTh dotted vertical lines illustrate a species richness comparison standardized to 24 species and 221 individuals, which was the observed Odonate abundance in the smallest (rivers) of the three water types data set. There was no evident of signicfi ant relationship along axes associated with lentic (Coenagrionidae and Libellulidae) two and three. Following the CCA components, two main or lotic systems (e.g., Calopterygidae, Coenagrionidae, and groups of species were distinguished. The rfi st one (e.g., Libellulidae) [19, 49]. In this study, we observed similar pat- Urothemis edwardsii, Palpopluera lucia, Palpopluera portia, tern of association, where Calopterygidae, Chlorocyphidae, Rhyothemis notate, and Acisoma inflatum) was representative Platycnemididae, and Aeshnidae were found in lotic systems, of the pond community. This group was mainly composed of while Libellulidae and Coenagrionidae were found in both the generalist heliophilic species, which mostly avoid flowing lentic and lotic environments but showed strong affinity water (Figure 10). The second group was represented by to lentic systems (ponds). The presence of Calopterygidae the combined eeff ct of streams and rivers (e.g ., Chlorocypha and Chlorocyphidae exclusively in the lotic systems may be selysi, C. luminosa, Sapho ciliata, and Phaon camerunensis). explained by their strong ani ffi ty to canopied cover and fast The group was mainly composed of Zygopterans which were flowing water bodies, which were characteristics of streams favoured by fast o fl wing water. The only Anisopteran species and rivers in the Ankasa Conservation Area. These features found in group two was the Micromacromia zygoptera,which are well known to represent the preferred habitat type of was also inuen fl ced by fast flowing water body. most species within the Calopterygidae and Chlorocyphidae families [19, 49]. Species from the Aeshnidae family are crepuscular in 4. Discussion nature and are well noted to shun the sun during the day but to come to light at night [26]. This is confirmed in our Several studies have shown that majority of Odonata families and species from anisopterans and zygopterans are either study where most species from the family Aeshnidae showed Mean species richness Taxa (95% confidence) 10 International Journal of Zoology Stream River Pond Freshwater type Figure 7: Mean Odonata species richness among the various water types. 7.0 6.5 6.0 Hc = 1.661, p =0.44, Kruskal-Wallis test 5.5 5.0 4.5 4.0 3.5 3.0 0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 alpha diversity scale parameter Streams Rivers Ponds Figure 8: Renyi diversity ordering that compares Odonata evenness and richness of 1403 individuals, across the three water types. Note that the shape of a habitat profile is an indication of its evenness. a strong association with dense vegetation cover along the in several studies (e.g., [51, 52]) and have been linked to stream banks and utilized the vegetation for perching and higher colonization rate characterized by lentic systems [51, roosting during the day. Also, a large section of Ankasa and 52]. Such is the case observed among lakes in the Brazilian Bonwere rivers that were characterized by rocky substrates Atlantic Forest, where higher Anisoptera species richness was recorded [53]. But our findings rather revealed ponds appeared to support the perching, roosting, and copulating of some zygopterans like Mesocnemis singularis and this proba- to support the least abundance and species richness of bly explains their high abundance. Dijkstra and Clausnitzer damselflies relative to dragonflies which are composed of [25] and Dijkstra [26] reported that Mesocnemis singularis only species from the Libellulidae family. This was probably typically prefersunny rocky substrate, as ecological niches for due to the scale of environmental disturbance and the perching, roosting, and copulating. geographical location of the ponds. For instance, in the Several pond-associated species, such as the Ceriagrions, tropics where this study was conducted, extreme tempera- Agriocnemis species,A.inflatum, C. flavifrons, O. lugubris, T. tures and erratic rainfall in recent times could have wider arteriosa, and P. lucia, have been classified as Heliophilics or ramicfi ations on surface water temperatures of the ponds, stagnant water tolerance species [17, 25, 26], which concur which were beyond the thermal threshold tolerance of the with this current study. Though small in catchment area, species. Lentic environments tend to be geologically less pre- ponds supported several distinct species that were never recorded in other water types and contributed to the greatest dictable through time [54], and this phenomenon tended to Odonata assemblages (abundance and richness) compared exert pressure on species to adapt faster in order to be able to the lotic environments. Globally, these pond-associated to disperse and then persist [5]. Ponds are important refuge species from the families Libellulidae and Coenagrionidae for Odonata conservation because they are relatively isolated are composed of several ubiquitous species that dominate in and show greater heterogeneity in species assemblages [55, unshaded habitats with stagnant water bodies [50]. 56], owing to stochastic effects acting on the colonization Higher Odonata species richness and diversity in lentic process [57]. Variability in pond isolation has the tendency systems relative to lotic environments have been reported to attract good disperser Odonata such as species within Mean species richness Renyi diversity index International Journal of Zoology 11 Transform: Log(X+1) Resemblance: S17 Bray Curtis similarity S7 R3 C 1 R4 S5 S3 S2 C 2 SI S6 R1 C 3 R2 S4 R6 C 4 R5 P10 P4 P6 P9 C 5 P7 P3 P2 P1 P5 P8 0 20 40 60 80 100 Similarity Figure 9: Group average dendrogram of similarity of species composition for Odonata among the water types in the Ankasa conservation area. the Libellulidae family which are ye fl rs and heliothermic in This, however, indicates that Odonata fauna in the Ankasa nature [58]. Conservation Area are restricted to specific water types, Streams and rivers (i.e., typical lotic systems) in con- with each water body supporting some specific species or trast, which were characterized with dense vegetation cover, assemblages not found in other water types. This finding supported the greatest abundance and species richness of supports the importance of maintaining a diversified body of zygopterans relative to anisopterans, as a result of their water, both lentic and lotic, natural or artificial, in ecosystem association with dense vegetation cover along the fringes management to achieve the ultimate goal of conserving of the systems, which provide conducive environment for diverse Odonata fauna and other sympatric freshwater bio- resting, mating, and breeding. The suit of different micro- diversity. habitat complexity along these lotic systems continuum may have contributed in species heterogeneity, largely dominated Data Availability by the Zygopteran functional group. Streams and rivers worldwide have been reported to provide heterogeneous and The data used to support the findings of this study are favourable environmental conditions for diverse Zygoptera available from the corresponding author upon request. species, [55, 58, 59] for their numerous life activities including nocturnal roosting, oviposition, emergence, reproduction, and perching substrate to thermoregulate [55, 58]. Streams Conflicts of Interest and rivers also share similar characteristics linked to their The authors declare that they have no conflicts of interest. geomorphology and o fl w regimes [16]. These systems have extensive catchments as compared to other lentic systems and this dovetailed with similar geomorphological features, flow Acknowledgments rate, and uniform vegetation cover of the waters will ensure less variability in their physicochemical variables [16]. This The authors are grateful to Rufford Foundation (20322- may result in similar colonization and dispersion rate, which 1 and 2) and International Dragonfly Fund for providing may lead to higher overlap in their Odonata fauna as evident financial support for this study. Our heartfelt appreciation in this study. is due to Viola Clausnitzer and Klaas-Douwe B. Dijkstra It was not uncommon that none of the species occurred for providing us with the identification hand books and for in all the three water types which reinforced our hypothesis. their immense contribution, mentoring, and advice towards Sampling sites 12 International Journal of Zoology Ordination Plot −1 −2 −3 −4 −2 −1 0 1 2 Axis 1 Odonata species Sampling sites Figure 10: Canonical correspondence analysis (CCA) ordination diagram, showing the relationship between environmental variables and Odonata species across the three water types in the Ankasa Conservation Area in Ghana. Species names are abbreviated with the first four letters of the genus and the first four letters of the species following Seidu et al., [17]. The blue arrows represent each of the environmental variables plotted pointing in the direction of maximum change of explanatory variables across the water types. Red and green ovals represent specialists’ species in lentic (ponds) and lotic (rivers and streams), respectively. the successful completion of the study. Finally, the authors [6] G. P. Mishra and A. K. Yadav, “A comparative study of physico- chemical characteristics of river and lake water in Central are grateful to David Amaning Kwarteng, Daniel Acquah- India,” Hydrobiologia, vol.59,no. 3,pp.275–278, 1978. Lamptey, Sulemana Bawa, and Emmanuel Amoah, for their special role in eld fi data gathering. [7] W.J. Mitschand J.W.DayJr.,“iTh nkingbig with whole- ecosystem studies and ecosystem restoration - A legacy of H.T. Odum,” Ecological Modelling, vol.178,no.1-2,pp. 133–155, 2004. References [8] T. E. Essington and S. R. Carpenter, “Nutrient cycling in lakes and streams: Insights from a comparative analysis,” Ecosystems, [1] D. L. Strayer and D. Dudgeon, “Freshwater biodiversity con- vol.3, no.2,pp.131–143, 2000. servation: recent progress and future challenges,” Journal of the North American Benthological Society,vol.29,no.1,pp.344–358, [9] R. W.Merritt and K.W. Cummins,Eds., An Introduction to the Aquatic Insects of North America,Kendall Hunt, 1996. [2] V.Clausnitzer, K.-D.B.Dijkstra,R.Kochet al.,“Focus on [10] M. Vincy, R. Brilliant, and A. P. Kumar, “Checklist of odonata African freshwaters: Hotspots of dragonfly diversity and con- species as indicators of riparian ecosystem of a tropical river, servation concern,” Frontiers in Ecology and the Environment, the southern western ghats,” Journal of Entomology and Zoology vol.10,no.3, pp. 129–134,2012. Studies,vol.4,no.2,pp.104–108,2016. [3] C. J. Vor ¨ os ¨ marty, P. B. McIntyre, M. O. Gessner et al., “Global [11] P. S. Corbet, “Dragonflies: behaviour and ecology of odonata,” threats to human water security and river biodiversity,” Nature, Aquatic Insects, vol.23, no.1,pp.83-83, 1999. vol. 467, article no. 7315, pp. 555–561, 2010. [12] A. Dolny, ´ F. Harabiˇs, D. Bar ´ taa, S. Lhota, and P. Drozd, “Aquatic [4] C. Revenga and Y. Kura, Status and Trends of Biodiversity of insects indicate terrestrial habitat degradation: Changes in Inland Water Ecosystems,vol.11of Technical Series,Secretariat taxonomical structure and functional diversity of dragonflies in of the Convention on Biological Diversity, Montreal, Canada, tropical rainforest of East Kalimantan,” Tropical Zoology,vol.25, 2003. no. 3, pp. 141–157, 2013. [5] C.Hof, M.Bra¨ndle, and R. Brandl, “Latitudinal variation of [13] F. Harabiˇsand A. Dolny,´ “Human altered ecosystems: Suitable diversity inEuropeanfreshwater animals is not concordant habitats as well as ecological traps for dragonflies (Odonata): across habitat types,” Global Ecology and Biogeography,vol. 17, The matter of scale,” Journal of Insect Conservation,vol. 16, no. no.4,pp.539–546, 2008. 1, pp. 121–130, 2012. Axis 2 International Journal of Zoology 13 [14] V. Clausnitzer, “Dragonyfl communities in coastal habitats of [32] J. S. Gray and F. B. Mirza, “A possible method for the detection Kenya: Indication of biotope quality and the need of conserva- of pollution-induced disturbance on marine benthic communi- tion measures,” Biodiversity and Conservation, vol. 12,no.2,pp. ties,” Marine Pollution Bulletin, vol. 10, no. 5, pp. 142–146, 1979. 333–356, 2003. [33] A. E. Magurran, Measuring Biological Diversity,vol. 131, Black- well Science Publishers, Oxford, UK, 2004. [15] J. T. Bried, B. D. Herman, and G. N. Ervin, “Umbrella potential of plants and dragonflies for wetland conservation: A quanti- [34] S. Fattorini, “Relations between species rarity, vulnerability, and tative case study using the umbrella index,” Journal of Applied range contraction for a Beetle Group in a Densely populated Ecology,vol.44,no.4,pp.833–842, 2007. region in the mediterranean biodiversity hotspot,” Conservation Biology,vol.28,no.1,pp.169–176,2014. [16] P. Williams, M. Whitfield, J. Biggs et al., “Comparative biodi- [35] S. Fattorini, “A simple method to tfi geometric series and broken versity of rivers, streams, ditches and ponds in an agricultural landscapeinSouthernEngland,” Biological Conservation,vol. stick models in community ecology and island biogeography,” 115, no. 2, pp. 329–341, 2004. A cta Oecologica,vol. 28, no.3, pp. 199–205, 2005. [17] I. Seidu, E. Danquah, C. Ayine Nsor, D. Amaning Kwarteng, and [36] R. MacArthur, “On the relative abundance of species,” e American Naturalist,vol.94,no.874,pp.25–36, 1960. L. T. Lancaster, “Odonata community structure and patterns of land use in the Atewa Range Forest Reserve, Eastern Region [37] S. Fattorini, F. Rigal, P. Cardoso, and P. A. V. Borges, “Using (Ghana),” International Journal of Odonatology,vol. 20, no.3- species abundance distribution models and diversity indices for 4, pp. 173–189, 2017. biogeographical analyses,” Acta Oecologica,vol.70,pp. 21–28, [18] D. Acquah-Lamptey, R. Kyerematen, and E. O. Owusu, “Using Odonates as markers of the environmental health of water and [38] N. J. Gotelli and R. K. Colwell, “Estimating species richness,” in its land related ecotone,” International Journal of Biodiversity Biological Diversity: Frontiers in Measurement And Assessment, and Conservation, vol.5, no.11, pp.761–769, 2013. vol. 12, pp. 39–45, 2011. [39] A. F. Siegel, “Rarefaction curves,” in Encyclopedia of Statistical [19] K.-D. B. Dijkstra and J. Lempert, “Odonate assemblages of Sciences, K. Samuel, C. B. Read, N. Balakrishnan, and B. running waters in the Upper Guinean forest,” Fundamental and Applied Limnology ,vol.157, no. 3,pp.397–412,2003. Vidakovic, Eds., 2006. [40] A. R´enyi, “On measures of entropy and information,” in [20] I. Seidu, C. A. Nsor, E. Danquah, and L. T. Lancaster, “Odonata Proceedings of the Fourth Berkeley Symposium on Mathematics, assemblages along an anthropogenic disturbance gradient in Statistics and Probability,vol.1,pp.1286-1261, University of Ghana’s eastern region,” Odonatologica ,vol.47,no.1-2,pp. 73– California Press, Berkeley, Calif, USA, 1961. 100, 2018. [41] C. E. Shannon, “A mathematical theory of communication,” Bell [21] J. A. Fulan and R. Henry, “A comparative study of Odonata System Technical Journal,vol.27,no. 4,pp. 623–656, 1948. (Insecta) in aquatic ecosystems with distinct characteristics [42] B. Tothm ´ er ´ es ´ z, “Comparison of different methods for diversity Um estudo comparativo de Odonata (Insecta) em ecossistemas aquatic ´ os com distintas caracter´ısticas,” Ambiencia, vol.9,no.3, ordering,” Journal of Vegetation Science,vol. 6,no.2, pp. 283– 290, 1995. pp. 589–604, 2013. [43] Ø. Hammer, D. A. T. Harper, and P. Ryan, “PAST: paleontolog- [22] F. Dowsett-Lemaire and R. J. Dowsett, “An update on the birds ical statistics software package for education and data analysis,” of Kakum National Park and Assin Atandaso Resource Reserve, Palaeontologia Electronica, vol.4,no.1,article 4, 2001. Ghana,” A report prepared for the Wildlife Division, Forestry Commission 75, Dowsett-Lemaire Misc, Accra, Ghana, 2011. [44] J. R.Krebs,J. D. Wilson,R.B. Bradbury, and G.M. Siriwardena, “eTh second silent spring?” Nature,vol.400,article 6745,pp. [23] J. B. Hall and M. Swaine, “Classification and ecology of closed- 611-612, 1999. canopy forest in Ghana,” Journal of Ecology, pp. 913–951, 1976. [45] C. J. F. ter Braak, “Canonical correspondence analysis: a new [24] N. J. Gotelli and R. K. Colwell, “Quantifying biodiversity: eigenvector technique for multivariate direct gradient analysis,” procedures and pitfalls in the measurement and comparison of Ecology, vol.67,no.5,pp.1167–1179, 1986. species richness,” Ecology Letters, vol.4,no.4,pp.379–391, 2001. [46] P. A. Henderson and R. M. H. Seaby, Community Analysis [25] K. D. Dijkstra and V. Clausnitzer, e Dragonflies And Dam- Package ., Pisces Conservation Ltd, Lymington, UK, 2000. selflies of Eastern Africa: Handbook for All Odonata,2014. [47] C. J. F. ter Braak and P.F.M. Verdonschot, “Canonicalcorre- [26] K.-D. B. Dijkstra, “African dragonflies and damselflies online,” spondence analysis and related multivariate methods in aquatic 2017, http://addo.adu.org.za. ecology,” Aquatic Sciences, vol.57,no. 3,pp.255–289,1995. [27] J. H. Zar, Biostatistical Analysis, Prentice Hall, Upper Saddle [48] M. Kent and P. Coker, Vegetation Description And Analysis. A River, NJ, USA, 4th edition, 1999. Practical Approach, John Wiley and Sons Ltd, West Sussex, UK, [28] A. S. Melo and L. U. Hepp, “Ferramentas estat´ısticas para ana´lises de dados provenientes debiomonitoramento,” Oecolo- [49] K.-D. B. Dijkstra, “Dragonflies and damselflies (Odonata) of the gia Brasiliensis, vol.12,pp.463–486, 2008. Atewa Range. A rapid biological assessment of the Atewa Range [29] B. McCune and J. B. Grace, Analysis of Ecological Communities, Forest Reserve, eastern Ghana,” in RAP Bulletin of biological MjM Sow ft are, Gleneden Beach, OR, USA, 2002. Assessment,vol. 47, pp. 50–54, 2007. [30] K. R. Clarke and R. N. Gorley, “Primer V5 (Plymouth rou- [50] V. J. Kalkman, V. Clausnitzer, K. D. B. Dijkstra, A. G. Orr, tines in multivariate ecological research): user manual/tutorial. D. R. Paulson, and J. van Tol, “Global diversity of dragonflies Primer-E,” 2001. (Odonata) in freshwater,” in Freshwater Animal Diversity Assess- ment, pp. 351–363, Springer, Dordrecht, the Netherlands, 2007. [31] B. J. McGill, R. S. Etienne, J. S. Gray et al., “Species abundance distributions: moving beyond single prediction theories to [51] A. S. Niba and M. J. Samways, “Remarkable elevational toler- integration within an ecological framework,” Ecology Letters, ance in an African Odonata larval assemblage,” Odonatologica , vol. 10, no. 10, pp. 995–1015, 2007. vol. 35, no. 3, pp. 265–280, 2006. 14 International Journal of Zoology [52] L. E. Stevens and R. A. Bailowitz, “Odonata biogeography in the Grand Canyon ecoregion, southwestern USA,” Annals of the Entomological Society of America,vol.102,no.2,pp. 261–274, [53] S. Renner, E. Perico, and G. Sahlen, “Man-made lakes form species-rich dragonyfl communities in the Brazilian Atlantic Forest (Odonata),” Odonatologica ,vol. 45, no.3-4,pp. 135–154, [54] I. Ribera,G. N.Foster, and A.P.Vogler, “Does habitat use explain large scale species richness patterns of aquatic beetles in europe?” Ecography, vol.26,no. 2,pp. 145–152, 2003. [55] B.Oertli, D.A. Joye,E.Castella, R.Juge, D.Cambin,and J.-B. Lachavanne, “Does size matter? eTh relationship between pond area and biodiversity,” Biological Conservation,vol. 104, no. 1, pp. 59–70, 2002. [56] M. Scheffer, G. J. Van Geest, K. Zimmer et al., “Small habitat size and isolation can promote species richness: Second-order effects on biodiversity in shallow lakes and ponds,” Oikos,vol. 112, no. 1, pp. 227–231, 2006. [57] D. S. Jeffries, J. R. M. Kelso, and I. K. Morrison, “Physical, chemical, and biological characteristics of the Turkey Lakes Watershed, central Ontario, Canada,” Canadian Journal of Fisheries and Aquatic Sciences, vol.45,no.1,pp.s3–s13, 1988. [58] P. S. Corbet and M. L. May, “Fliers and perchers among Odonata: Dichotomy or multidimensional continuum? A pro- visional reappraisal,” International Journal of Odonatology,vol. 11, no. 2, pp. 155–171, 2008. [59] L. Maltchik,C. Stenert,C. B. Kotzian, and M. M. Pires, “Responses of odonate communities to environmental factors in southern Brazil wetlands,” Journal of the Kansas Entomologi- cal Society,vol.83, no. 3,pp.208–220, 2010. 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Patterns of Odonata Assemblages in Lotic and Lentic Systems in the Ankasa Conservation Area, Ghana

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Hindawi Publishing Corporation
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Copyright © 2019 Issah Seidu et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
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1687-8477
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1687-8485
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10.1155/2019/3094787
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Abstract

Hindawi International Journal of Zoology Volume 2019, Article ID 3094787, 14 pages https://doi.org/10.1155/2019/3094787 Research Article Patterns of Odonata Assemblages in Lotic and Lentic Systems in the Ankasa Conservation Area, Ghana 1 2 1 Issah Seidu , Collins Ayine Nsor , Emmanuel Danquah , 1 1 Paul Tehoda, and Samuel K. Oppong Department of Wildlife and Range Management, Faculty of Renewable Natural Resources, Kwame University of Science and Technology, Kumasi, Ghana Department of Ecotourism and Forest Recreation, Kwame Nkrumah University of Science and Technology, Kumasi, Ghana Correspondence should be addressed to Issah Seidu; antwiseidu88@gmail.com Received 5 November 2018; Accepted 15 January 2019; Published 10 February 2019 Academic Editor: Marco Cucco Copyright © 2019 Issah Seidu et al. is Th is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Our study examined Odonata assemblages distribution pattern and the predictive factors that accounted for this in the lotic and lentic water systems within the Ankasa Conservation Area (Ghana). A total of 23 sites with sampling protocol of 2 researchers per hour per sampling site were used to survey Odonata species over two seasons in the three water bodies (streams, rivers, and ponds). Broken stick model, individual-based rarefaction, and Renyi diversity ordering were employed to quantify community assemblages. Ordination technique was also used to determine the Odonata-environmental relationship. A total of 1403 individuals, belonging to 47 species (22 Zygoptera and 25 Anisoptera) in six families, were recorded. Species richness (Hc =3.414, p =0.169) and diversity (Hc =1.661, p = 0.44) generally did not dieff r among the three water systems. However, from individual sites, ponds appeared mostly diverse (𝛼 -scale = 0.04, Renyi index (r) = 5.86 to 𝛼 = 3.5, r = 3.12), in spite of their lowest species abundance and richness. At the suborder level, ponds equally exhibited the highest Anisoptera species richness (9.90 ± SE 0.640) compared with Zygopterans (0.80± SE 0.291). Overall, Anisopterans (K=16.51, p= 0.00026) and Zygopterans richness (K=16.39, p= 0.00023) differed significantly among the three subsystems, while Odonata composition also differed significantly among the various water bodies (ANOSIM: global R=0.94, p<0.001). Flow rate, water temperature, channel width, and turbidity were the key predictive factors that influence the structure of Odonata species assemblages. The results highlight the need to improve the functional status of the lentic and lotic systems, with the ultimate goal of conserving diverse Odonata fauna and other sympatric freshwater biodiversity. 1. Introduction Freshwater habitats present two major differing water systems, lotic (running) and lentic (standing) waters, which Freshwater habitats cover only 1% of the total earth sur- differ in their environmental and spatiotemporal settings [5]. face and contain 10% of the earth biodiversity [1]. Their They are distinguished by physicochemical parameters of the importance in sustaining biodiversity and human welfare is water such as turbidity, organic matter, pH [6], dissolved oxy- undeniable. Freshwater resources are the major sources of gen [7], nutrients content [8], and flow regimes. These water livelihood to Afrotropical rural and periurban folks [2]. They systems together support heterogeneous environment which provide water supplies for human consumption, industrial provide favorable conditions for both vertebrate and inverte- utilization, and ecosystems support for sher fi ies and other brates communities including the amphibious Odonata taxa. aquatic biodiversity. However, wetlands are considered one Odonata are denizen of freshwater environments such of the most jeopardized ecosystems in the world [3]. Many wetlands worldwide are experiencing dramatic anthropic as rivers, lakes, ponds, wetlands, and, to some extent, phy- totelmata and brackish water resources [9, 10]. They play change, mostly for agricultural purpose [4]. Generally, these changes are associated with abiotic conditions which are significant role in freshwater ecosystem functioning, acting as both prey (fed by vertebrates and other large insects) normally not found in nature with cascading impacts on residence aquatic biota. and top predators (feeding on smaller insects in vertebrate 2 International Journal of Zoology free aquatic environment) [11]. Due to the reliability of both July and September to November, with average annual rainfall larval and adult Odonata to specicfi water conditions for of 1700 to 2000 mm [22]. survival [12], and their sensitivity to habitat disturbances, they are eeff ctively used asindicatorsof water quality [13, .. Description of Sampled Water Types in the Study Area. 14]. Odonata, therefore, serve as an umbrella species in Stream (𝑛 = 7): We located all the sampling sites along biodiversity conservation [15] and represent specific biotic the Asufia stream and a stream which is tributary to the wetland assemblages. Ankasa River (Figure 1). Three sites were laid along the Asuafi Sustainability of Afrotropical freshwater resources and stream, while four were located on the other stream. The sites their associated Odonata fauna requires knowledge of the were characterized by sandy substrate. The channel width contribution of different water bodies in particular ecosys- ranged from 1 m to 1.9 m while the depth was from 0.1 m to tems. Theseinclude knowledgeabout thespecies richness, 0.21 m. The water was o fl wing rapidly through dense canopy diversity, and community structure in different water types, cover, with the trees and shrubs being the dominant bank the variability of water systems across the landscape, and vegetation. the net contribution of these water systems to the catchment River (𝑛 = 6): All the sites were located along two biodiversity [16]. In general, such information is practically major rivers (Ankasa and Bonwere River) in the Ankasa scanty worldwide but particularly in West Africa. This is Conservation Area (Figure 1). Three sites were laid along the result of traditional Odonata research being geared each river, representing the total sampling sites. The Ankasa towards specific water body. For example, especially in and Bonwere Rivers are characterized by rocky and sandy Ghana, most current research on Odonata assemblages has substrates. All sites were associated with rapids and highly virtually focused on rivers (see [17–19]), and streams [19, 20] oxygenated, cold water. The channel width was between 2 m with little or no studies describing other natural and artificial and 15 m while the depth ranged from 0.3 m to 0.75 m. The lentic freshwater systems such as ponds, pools, and lakes, sites were laid adjacent to intact secondary forest vegetation although these water bodies are well known to harbor diverse with the margins mainly composed of trees and shrubs, and Odonata fauna and higher Odonata richness elsewhere [21]. small patches of various grasses (Poaceae). The water bodies In order to contribute to the initial understanding of the also pass through dense canopy with low sun exposure except importance and the influence of dieff rent water types on in sun flecks caused by tree falls. the Ghanaian Odonata biodiversity, differences in Odonata Pond (𝑛 = 10): All the 10 ponds were naturally permanent assemblage structure of lotic and lentic systems were investi- water bodies located in and outside the Ankasa Conservation gated. We hypothesized that adult Odonata composition will Area. Four ponds were located in the forest reserve, while be signicfi antly dieff rent among the water types due to their six were outside the forest adjacent to cultivated rubber, preference for different water bodies [12, 13]. Accordingly, we vegetables, and cocoa plantation which were mostly used for addressed two major questions: (1) are there any signicfi ant irrigation by the local communities (Figure 1). The bottoms differences in Odonata abundance, richness, and community were mainly composed of mud/clay and organic matter. Most composition between the lotic (rivers and streams) and lentic of the ponds were surrounded by partial vegetation structure systems (ponds)? and (2) are there signicfi ant differences in with high amount of sun penetration. The dominant plant the abundance and richness of Anisopterans and Zygopterans families in the marginal zones were Cyperaceae and Poaceae. among the water types? In order to address these questions, Pondssurfaceswereassociated with stands ofemergent or we compared adult Odonata assemblages occurring in 7 sites floating vegetation which were utilized by the adult Odonata along two major rivers, 6 sites along three different streams, for perching. and 10 different ponds found in and outside the Ankasa Conservation Area. .. Odonata Sampling Procedures. We sampled adult indi- viduals of all Odonata species at 23 sites with a sampling protocol of 2 researchers per hour per sampling site, along 2. Materials and Methods the three different water types, Rivers, Streams, and Ponds ∘ 󸀠 ∘ .. Study Site. Ankasa Conservation Area (5 17 Nand 2 in the Ankasa Conservation Area. We sampled simulta- 39 W) is a twin Protected Area comprising Nini-Suhien neously, collecting and noting the species occurring, and their abundances in each sampling site until no new species National Park and the Ankasa Resource Reserve [22]. It is about 500 km situated in the Western Region of Ghana, were encountered for approximately one hour for each visit. and the only area in the Wet Evergreen Forest [23]. Ankasa Sampling was done from January, 2017, to March, 2017, for Conservation Area is designated as a Globally Significant thedry season whilethe wetseason sampling took place Biodiversity Area (GSBA) and Important Bird Area (IBA) from May, 2017, to July, 2017. We sampled all adult Odonata [23]. during the day between the hours of 9 am and 5 pm. We Ankasa Conservation Area presents an ideal ecosystem captured all adult Odonata individuals where possible, using forthisstudy, asitboasts ofa signicfi ant number of complex a hand net. We identified each specimen to species level in situ, using Dijkstra, and Clausnitzer, [25] identifiction keys. and diverse freshwater systems including riverine, streams, and ponds. These wetlands and their associated forest envi- Where identification of some species was not possible on ronmentsupport themostbiological diversity of any kind the eld fi , we photographed them and then used the African Dragonflies and Damselflies Online database (ADDO) [26], in Ghana [22]. The climate of the area is characterized by a distinctive bimodal rainfall pattern occurring from April to for subsequent idenitificantion. International Journal of Zoology 3 ∘   ∘   ∘   ∘   ∘   ∘   ∘   2 44 10 W 2 41 40 W 2 39 10 W 2 36 40 W 2 34 10 W 2 31 40 W 2 29 10 W River Stream Roads Towns Ankasa CA Sampling sites Ponds River Kilometers Stream 0 1.25 2.5 5 7.5 10 Figure 1: Map of the study area in and near the Ankasa Conservation Area, Western Region, with the situation of the 23 sampling sites of the three water types. .. Measurement of Biophysical Variables. We recorded abi- Allmultivariate analyses weredoneusing PRIMER 6.1.5 otic variables concurrently during the Odonate sampling, to package [30]. assess their influence on Odonate community structure. Sur- face water temperature ( C), pH, dissolved oxygen (mg/L), ... Species Abundance Distribution (SAD) for Odonata turbidity, conductivity, altitudes, flow rate, channel width Species. The application of species abundance distribution and depth, aquatic vegetation, substrate type, and bankside models in the study of species patterns has been widely vegetation were all measured in all sampling sites following used in community ecology by most scientists [31], as well Seidu et al. [17] procedure. as measuring the impact of disturbance on community structure [32]. In this study, Odonate abundance as a measure .. Data Analysis. We first tested the normality of the abun- of diversity was quantified using rank abundance model [33]. dance data set using Shapiro-Wilk test [27]. The abundance In each site, we listed the number of Odonata species for all of the wet and dry seasons, say 𝑆 data was log(X+1) transformed prior to analysis. Bray-Curtis , represented by one similarity indices and nonparametric multidimensional scal- individual, and the number of species, say 𝑆 , represented ing (NMDS) were used to determine relationships of species by K individuals, where K denotes the abundance of the composition among the sampling sites of the various water most abundant species and 𝑆 + ... + 𝑆 = S [34]. 1 𝐾 bodies. To test for the significant difference in species com- Accordingly, the sequence of relative frequencies 𝑓 = 𝑆 /S (r 𝑟 𝑟 position among the various water types, we employed one- =1...K) constitutes a frequency distribution for the number way analysis of similarities with 999 permutations (ANOSIM; of individuals per species which is usually referred to as the [28, 29]), with Bray-Curtis similarities as dependent and the species-abundance curve [34]. We then tfi ted the MacArthur three different water types (streams, rivers, and ponds) as broken stick model (BS) [35, 36] in the species abundance independent factor. Similarity percentage analysis (SIMPER) data, using the regression model approach [35] to determine routine in primer [30] was used to determine average dis- the pattern of species communities in each of the freshwater similarity between the water bodies and the various species systems. MacArthur [36] suggested that the niche space could contributing to the most similarity within each water body. be compared to a stick of length 1, where n – 1 points would ∘   ∘   ∘   ∘   ∘   ∘   5 10 50 N 5 13 20 N 5 15 50 N 5 18 20 N 5 20 50 N 5 23 20 N 4 International Journal of Zoology randomly generate n segments of lengths proportional to the suborders (Anisoptera and Zygoptera) for streams, rivers, and number of individuals of each species in the community, ponds. given as Renyi [40] extended the concept of Shannon’s entropy [41], by defining the entropy of order 𝛼 (𝛼≥ 0, 𝛼 ≠ 1) of a probability distribution (p , p ...p ). Diversity prolfi e values 𝑁 1 1 2 s 𝑛 = ∗ ∑ (1) (H-alpha) were calculated from the frequencies of each com- 𝑆 𝑛 𝑖=1 ponent species (proportional abundances pi =abundance of species i/ total abundance) and a scale parameter (𝛼 )ranging (see [36]) Where 𝑛 represents the number of individuals of from zero to infinity as the species i; N represents the total number of individuals; and S represents the total number of species in the commu- 𝑠 (log ∑ 𝑝𝑖 ) 𝑖−1 nity. (3) (𝐻 )= (1− 𝛼 ) Thismodel approach wasused in order to test against the null hypothesis (𝐻 ) that species abundance distribution and (See [42]). Odonate abundance, richness, and diversity richness did not differ in each of the three water systems. ordering were performed using PAST version 3.06 sowa ft re All the species in each of the sampling sites per water type package [43], which provides robust algorithm as shown in were ranked from the most to the least abundant on the rank Krebs et al. [44]. abundant curve [37]. Each species rank is plotted on the x- Due to the nonnormal nature of the data set, a Kruskal- axis, and the abundance is plotted on the y-axis. Wallis test was applied to test for the differences in Odonata With the broken stick model, if a log scale is used for and suborders (Anisoptera and Zygoptera) abundance and abundance, the species exactly fall along a straight line, richness among the 23 sites, using PAST version 3.12 [43]. according to the model equation log𝐴= 𝑏 +𝑏 𝑅 ,where 𝑜 1 Homogeneity of species variance among sample plots was Aisthe speciesabundance, R is the respective rank, and evaluated, using Levene test [45], defined as 𝑊 = ((𝑁−𝑘)/(𝑘− b and b are optimized tfi ting parameters [32]. Analysis of 𝑘 𝑘 2 2 ́ ́ ́ 1))(∑ 𝑁𝑖( Z𝑖− Z𝑖) /∑ ∑ (𝑍𝑗−𝑖 Z𝑖) ) where 𝑗𝑍𝑖 can 𝑖=1 𝑖=1 𝐽=1 covariance (ANCOVA) was applied to test for the significant have one of the following three definitions. difference of the slope of the SADs for the three water types, ́ ́ ́ 2 𝑗=𝑍𝑖 ⌈𝑌 −Y ⌉ where Y is mean of the 𝑖𝑡ℎ subgroup; Y is 𝑖 𝑖 𝑖 while Pearson’s Chi-square test (𝜒 ) was applied to determine the median of the 𝑖𝑡ℎ subgroup, and, finally, 𝑗=𝑍𝑖 ⌈𝑌 − Y 𝑖⌉ , whether an observed distribution along the goodness of tfi statistically differed in the BS model. Among the four notable where Y 𝑖 is the 10% trimmed mean of the 𝑖𝑡ℎ subgroup. Z𝑖 SAD models (i.e., geometric, log series, log normal, and BS), are the group means of the 𝑗𝑍𝑖 and Z is the overall mean of the BS model is the only one that fundamentally describes the the 𝑗𝑍𝑖 . process of niche partitioning in a community where species exhibit continuous nonoverlapping niches [33]. .. Environmental Predictors of Odonata Distribution. We Individual-based rarefaction techniques [38] were used determined the relationships between the abiotic variables to compare Odonate richness across the three water sys- recorded and the species occurrence in the various water tems (rarefaction curves). Rarefaction curves are created by bodies using a canonical correspondence analysis (CCA, randomly resampling the pool of N samples multiple times [45]). We used the Environmental Community Analysis and then plotting the average number of species found in (ECOM.exe) version 1.4 packages [46] to perform the CCA each sample (1,2 .. . 𝑁) [24]. us, Th rarefaction generates analysis. The significance of the rfi st two axes generated the expected number of species in a small collection of n in the analysis was validated through the Monte Carlo test individuals (or n samples) drawn at random from the large (using 5000 iterations) [47]. Environmental variables utilized pool of N samples. The rarefaction curve 𝑓 is defined as in the CCA were water temperature, dissolved oxygen, pH, turbidity, conductivity, o fl w rate, and channel width and −1 𝑁 𝑁− 𝑁𝑖 depth. CCA is a direct method of ordination with the (2) 𝑓 =𝐸[𝑋 ]=𝐾 − ( ) ∑ ( ) 𝑛 𝑛 resulting outcome being the variability of the environmental 𝑛 𝑛 𝑖=1 data, aswell asthe variability ofspecies data [48]. (See [38]). Where 𝑋 = the number of groups still present in the subsample of “n”lessthan 𝐾 whenever 3. Results at least one group is missing from this subsample, 𝑁= total number of items, 𝐾= total number of groups, 𝑁𝑖 = .. General Pattern of Odonata Composition and Abundance total number of items in group 𝑖 (𝑖 = 1,... 𝑘) [24, 39]. u Th s, Distribution across the Streams, Rivers, and Ponds. Atotal the linear model for the BS was tfi ted for each rarefied rank of 1403 adult Odonata specimens belonging to 47 species, in order to build the 95% cond fi ence limits for the slopes of and six families, were registered in streams, rivers, and all sampling sites. ponds in the study area (Tables 1(a) and 1(b)). Of the 47 Rarefaction methods, both sample based and individual species recorded, 22 Zygoptera species belonging to four based, allow for meaningful standardization and compari- families (Calopterygidae, Chlorocyphidae, Coenagrionidae, son of datasets [24]. We compared the estimated Odonata and Platycnemididae) and 25 Anisopterans from two families abundance and species richness, as well as the estimated (Aeshnidae and Libellulidae) were recorded (Tables 1(a) and abundance and number of species belonging to the respective 1(b)). Libellulidae was the dominant family with 13 species, 𝐼𝐽 𝐼𝐽 𝑁𝐼 International Journal of Zoology 5 Table 1 (a) Checklist and abundance of Zygoptera (damselflies) species recorded in streams, rivers, and ponds in the Ankasa Conservation Area. Species that o ccurred exclusively in streams are represented by (∗), exclusively in rivers (#), and exclusively in ponds (!). Species shared between streams and rivers are represented by (∗#), between streams and ponds (∗!), and between rivers and ponds (#!) Family Zygopterans Stream River Ponds Total Calopterygidae Phaon camerunensis Sjosted ¨ t, 1900∗ 19 0 0 19 Phaon iridipennis (Burmeister, 1839)∗#6 16 0 22 Sapho bicolor Selys, 1853∗ 20 0 2 Sapho ciliata (Fabricius, 1781)∗#39 10 0 49 Umma cincta (Hagen in Selys, 1853)∗ 20 0 0 20 Chlorocyphidae Chlorocypha luminosa (Karsch, 1893)∗#33 19 0 52 Chlorocypha radix Longfild, 1959 ∗#16 8 0 24 Chlorocypha selysi Karsch, 1899∗#10 29 0 39 Coenagrionidae Agriocnemis exilis Selys, 1872! 0 0 4 4 Agriocnemis zerafica Le Roi, 1915! 0 0 13 13 Ceriagrion corallinum Campion, 1914∗!8 0 13 21 Ceriagrion glabrum (Burmeister, 1839)! 0 0 26 26 Ceriagrion rubellocerinum Fraser, 1947∗#7 6 0 13 Pseudagrion hamoni Fraser, 1955∗#3 9 0 12 Pseudagrion isidromorai Sart, 1967∗ 15 0 6 Pseudagrion kersteni (Gerstack ¨ er, 1869)∗ 50 0 5 Pseudagrion melanicterum Selys, 1876∗#23 24 0 47 Pseudagrion hamoni Fraser, 1955∗ 26 0 8 Pseudagrion sjoestedti F¨orster, 1906 02 0 2 Platycnemididae Mesocnemis singularis Karsch, 1891# 0 26 0 26 Elattoneura balli Kimmins, 1938∗#38 16 0 54 Elattoneura villiersi (Fraser,1948)∗ 34 3 0 37 Total number of individuals Total number of species (b) Checklist and abundance of Anisoptera (dragonflies) species recorded in streams, rivers, and ponds in the Ankasa Conservation Area. Species that occurred exclusively in streams are represented by (∗), exclusively in river (#), and exclusively in pond (!). Species shared between streams and rivers are represented by (∗#), between streams and ponds (∗!), and between rivers and ponds (#!) Family Anisopterans Stream River Ponds Total Aeshnidae Gynacantha bullata Karsch, 1891∗ 50 0 5 Gynacantha cylindrata Karsch, 1891∗ 10 0 1 Libellulidae Acisoma inflatum Selys, 1882! 00 147 147 Aethriamanta rezia Kirby, 1889! 0 0 33 33 Chalcostephia flavifrons Kirby, 1889! 0 0 90 90 Cyanothemis simpsoni Ris, 1915# 0 9 0 9 Eleuthemis buettikoferi Ris, 1910# 0 9 0 9 Neodythemis klingi (Karsch, 1890)∗ 14 0 0 14 Micromacromia zygoptera (Ris, 1909)∗ 1000 10 Olpogastra lugubris (Karsch, 1895)#! 0 3 22 25 Orthetrum austeni (Kirby, 1900)! 0 0 19 19 Orthetrum julia Kirby, 1900∗!7 0 13 20 Orthetrum microstigma Ris, 1911! 0 0 6 6 Orthetrum stemmale (Burmeister, 1839)∗!6 0 7 13 Orthetrum trinacria (Selys, 1841)! 00 6 6 Palpopleura lucia (Drury, 1773)! 0 0 81 81 Palpopleura portia (Drury,1773)! 0 0 71 71 Pantala flavescens (Fabricius, 1798)! 0 0 5 5 6 International Journal of Zoology (b) Continued. Family Anisopterans Stream River Ponds Total Rhyothemis notata (Fabricius, 1781)#! 0 3 100 103 Rhyothemis semihyalina (Desjardins, 1832)! 0 0 38 38 Trithemis aconita Lieinc ft k, 1969∗!3 0 17 20 Trithemis arteriosa (Burmeister, 1839)#! 0 9 86 95 Trithemis bifida Pinhey, 1970#! 0 3 2 5 Trithemis dichroa Karsch, 1893#! 0 3 43 46 Urothemis edwardsii (Selys, 1849)#! 0 3 28 31 Total number of individuals Total number of species Table 2: Results of the broken stick model for the abundance rank distribution of Odonata species, calculated for each of the three water types. Sample Intercept ±S.E. Slope ±S.E. R Prob. Streams 4.05 ± 1.75 0.55 ± 0.19 0.37 0.009 Rivers 2.96 ± 1.19 0.26 ± 0.09 0.38 0.008 Ponds 25.27 ± 5.41 -1.02 ± 0.43 -0.33 0.02 Slope of SAD: 𝐹 = 6.22, p (regr): 0.002 2,138 (ANCOVA interactions x species rank) Monte-Carlo Permutation (n = 99999): p<0.0014 Levene test for homogeneity of variance:p<0.0015 However, from three water types, we observed Odonate followed by Coenagrionidae (n =12) andCalopterygidae (n = 4), in rivers and ponds. Community assemblages across abundance in ponds to be the highest (n =870), but their the three sites were ranked from the most abundant to the spatial distribution did not differ significantly along the least abundant (Figure 2). Their abundance distribution fitted slopes of the curve (𝜒 P = 25.07, P = 0.24). Similar abundance well in the broken stick distribution (BS) model and generally and distribution trends were observed in streams (n =312, showed significant difference in the slopes of the three water 𝜒 P =7.12, P = 0.99) and rivers (n = 221, 𝜒 P = 4.11, P = systems (𝐹 = 6.22, p(regr) = 0.002, ANCOVA interactions 0.99) (Table 2, Figure 2). Individuals per sample site, in ponds 2,138 xspecies rank)(Table 2,Figure 2). Further Monte Carlotest (87.00 ± SE 8.83), streams (44.6± SE 4.4), and rivers (36.8 ± SE (n = 99999) revealed signicfi ant dieff rence in SAD slopes ( p 4.23), equally followed similar trend (Hc =16.72, P = 0.0002, = 0.001). Kruskal-Wallis test)(Figure 3). Pairwise comparisontest showed a significant difference between ponds and streams At the suborder level, streams had the greatest mean (P = 0.003) and ponds and rivers (P = 0.004). However, Zygopterans abundance (38.0± SE 4.29) (e.g., E. balli =54, C. there was no significant difference in Odonata abundance luminosa =52, and S. ciliata =49),compared with Anisopter- between rivers and streams (P > 0.05). Comparison of the ans (6.57± SE 2.05). Conversely, the ponds exhibited the SADs for the three water systems helps to distinguish a greatest Anisoptera abundance (81.40± SE 8.264) (e.g., A. specific habitat quality, in relation to its influence on Odonate inflatum =147, R. notata =103, and T. arteriosa =95) while abundance, while the shape of the rank abundance curve zygopterans were the least abundant (5.60± SE 1.96) (Table 3 generally revealed differences in Odonate dominance and and Figure 4). Sapho bicolor and P. sjoestedti represented by evenness from individual habitats, and which reflects in their double individuals (doubleton) and Gynacantha cylindrata, relative tolerance to disturbances. single individual (singleton), T. bifida (𝑛 = 5) and G. bullata (n = 5), were the least dominant Zygopterans and Anisopterans, respectively, in the study area (Tables 1(a) .. Comparison between Zygopterans and Anisopterans Spe- and 1(b), Figure 2). There was a significant difference in cies Richness among the Water Types. Ponds exhibited the the abundance of Zygopterans (K =16.5, p = 0.00025) and highest Anisoptera species richness (9.90± SE 0.640) but Anisopterans (K = 16.28, p= 0.0003) among the three sites. the lowest number of Zygopterans (0.80± SE 0.291) (Fig- Zygopteran abundance in ponds differed signicfi antly in ure 5). The streams had the highest Zygopteran richness pairwise comparison with streams (p= 0.0007) and rivers (p= (7.57± SE 0.481) but exhibited almost similar Anisoptera 0.0018) but showed no difference between rivers and streams species richness (2.0± SE 0.577) with rivers (1.8± SE 1.014) (p= 0.174). Similarly, the Anisoptera abundance in ponds (Figure 5). Kruskal-Wallis test showed a significant differ- varied significantly in the pairwise comparison with streams ence in Zygoptera species richness (K=16.39, p= 0.0002) (p= 0.0007) and rivers (p= 0.001) but no significant difference and Anisoptera richness (K=16.51, p= 0.0003) among the occurred between the rivers and streams (p = 0.825). water types. Pairwise comparison test showed a significant International Journal of Zoology 7 2.0 2.0 3.0 1.5 1.5 2.5 1.0 1.0 2.0 0.5 0.5 1.5 Rivers 0.0 0.0 1.0 Streams Ponds −0.5 −0.5 0.5 −1.0 −1.0 0.0 Species rank order Species rank order Species rank order Figure 2: Broken stick model for Odonata rank abundance distribution across the three water types in Ankasa Conservation Area. Abundance is based on cumulative values per species test sites. Notice that SADs are ordered in decreasing magnitude and plotted against the corresponding rank order. Table 3: Canonical coefficients and the correlations with the first three axes of the environmental variables of the canonical correspondence analysis (CCA) for the three water types. Interset correlations were significant ( p <0.05∗) for the three axes. Axis I Axis II Axis III Correlation 0.573∗ 0.098 -0.156 Turbidity 0.549∗ 0.081 0.114 Flow rate Width -0.706∗ -0.076 0.164 -0.260 -0.009 0.114 Depth -0.421 0.054 -0.092 Conductivity 0.203 0.079 0.145 Do Temperature -0.748∗ -0.060 0.070 0.154 -0.137 0.091 pH Canonical Eigen value 0.651 0.445 0.185 22.3 15.29 6.338 % variance explained 22.3 37.6 43.96 Cumulative % variance Pearson correlation 0.979 0.708 0.781 species/environment scores Kendal rank correlation of 0.684 0.463 0.597 species/environment scores difference in Zygopteran richness between ponds and streams However, mean species richness per sample site was rather (p= 0.0006), and between rivers and ponds (p= 0.001), the highest in ponds (10.7 ± SE 0.56), while rivers had the least but no difference existed between streams and rivers ( p= number (8.7 ± SE 0.92) (Figure 7). Homogeneity of species 0.56). Similarly, Anisoptera species richness in ponds differed variance among the three water systems differed significantly significantly with streams ( p= 0.00071) and rivers (p= 0.001), (p<0.0002, Levene test)(Table 2). but there was no significant difference between streams and Observed trends in Odonate structural assemblages (i.e., rivers (p= 0.82). abundance, evenness, and richness) reflected in the Renyi diversity ordering (from higher to lower indices; along an .. Trends in Odonata Richness and Diversity in the ree increasing alpha scale values) (Figure 8). Overall, Odonate Water Systems. Interpolating the SADs across the streams, diversity did not differ significantly ( Hc =1.661, p = 0.44) rivers, and ponds, with sample-based rarefaction, revealed across the three water types. However, from individual sites, that Odonate richness among the three systems was not we observed that Odonates from ponds appeared mostly diverse (𝛼 -scale = 0.04, Renyi index (r)= 5.86 to 𝛼 =3.5, r = signicfi antly dieff rent ( Hc =3.414, p =0.169, Kruskal-Wallis test) (Figure 6) and did not follow similar pattern observed 3.12), in spite of their lowest species abundance and richness in individual abundance. Chao-1 estimated species richness (Figure 5). This was linked to the shallower SAD curve for the three sites showed streams to be the highest (n = observed in Figure 2. u Th s, species abundance distributions, 24.33), followed by ponds (n =23) andrivers (n = 22). with shallower curve, tended to be highest in diversity, while log Abundance 0.0 2.5 5.0 7.5 10.0 12.5 15.0 17.5 20.0 22.5 25.0 log Abundance 0.0 2.5 5.0 7.5 10.0 12.5 15.0 17.5 20.0 22.5 log Abundance 0.0 2.5 5.0 7.5 10.0 12.5 15.0 17.5 20.0 22.5 8 International Journal of Zoology Hc = 16.72, P = 0.0002, Kruskal Wallis test Stream River Pond Water type Figure 3: Mean Odonata species abundance among the various water types. Stream River Pond Water type Zygoptera Anisoptera Figure 4: Comparison of mean species abundance of Zygopterans and Anisopterans among the water types. those with steeper curves were less diverse (Figure 6). Species luminosa (16%). Cholorocypha selysi (26%), P. melanicterum from the riverine systems were the least diverse and ranged (19%), and M. singularis (13%) contributed most to similarity from 𝛼 = 0.04, r =5.83to 𝛼 =3.5, r = 3.08 and were found at the in river community, whereas T. arteriosa (16%), A. inflatum bottom of the Renyi index curve (Figure 6). Odonate diversity (15%), and P. lucia (14%) were greatest contributing species in in streams (𝛼 = 0.04, r =5.84 to 𝛼 =3.5, r = 3.07) could barely pond communities. be distinguished from those in the riverine systems, as their The species composition of Odonata dieff red signicfi antly curves were spatially similar. between the various water bodies (ANOSIM: global R= 0.94, p<0.001). Pairwise comparison test showed a significant difference in species composition between rivers and ponds .. Similarity in Odonata Composition among Streams, (R= 0.98, p= 0.002). Also, streams revealed weak significant Rivers, and Ponds. The Nonparametric Hierarchical Cluster dieff rence with rivers ( R= 0.52, p= 0.02) but higher significant analysis of species occurrence showed vfi e different clusters difference with ponds ( R=0.99, p= 0.001). (P8, P5, P1, P2, P3, P7, P9, P6, P4, and P10), (R5 and R6), (S4, R2, R1, and S6), (S1 and S2), and (S3, S5, R4, R3, and R7) at 40% similarity index (Figure 9). The species .. Environmental Predictors of Odonata Structural Dis- tribution and Diversity. Canonical correspondence analysis occurrence in ponds showed a strong significant separation from streams and rivers communities. However, the sampling (CCA) showed the overall relationships between species sites of stream and river were ecologically less distinct and distribution and the biophysical variables recorded (Table 3, showed a higher species overlap with each other (Figure 9). Figure 10). Among the eight biophysical variables initially The Similarity Percentage (SIMPER) analysis revealed a included in the analysis, only four biophysical variables, similar trend, suggesting that streams and ponds (98.72%) namely, o fl w rate, water temperature, channel width, and and rivers and ponds (93.87%) exhibited greatest average turbidity, were shown to strongly inu fl ence the structure dissimilarity in species composition to one another. Streams of species assemblages. Species assemblages along the rfi st and rivers (67.31%) were relatively similar to each other axis correlated significantly with water temperature (r =- in Odonata species composition. SIMPER also revealed an 0.74, p<0.05), channel width (r = -0.70), o fl w rate ( r = 0.54, average similarity within the streams (49%), rivers (43%), and p<0.05), and turbidity (r = 0.57, p<0.05) (Table 3, Figure 10). ponds (63%). Species contributing most to similarity in the CCA axes 1 and 2 jointly explained 37.6% of the total variation stream community were E. balli (23%), S. ciliata (17%), and C. in species structural distribution and diversity among sites. Mean species abundance Mean species abundance International Journal of Zoology 9 Stream River Pond Water type Zygoptera Anisoptera Figure 5: Comparison of mean species richness of the Zygopterans and Anisopterans among the water types. Hc = 3.414, p = 0.169, Kruskal- 15 Wallis test Chao-1 estimate: Streams: 24.33 Rivers: 22 Ponds: 23 100 200 300 400 500 600 700 800 Odonata specimens Figure 6: Standardized comparison of Odonata richness for individual-based rarefaction curves. The data represent summary counts of Odonates that were recorded from the three water types in Ankasa Conservation Area. eTh red, blue, and green lines are the rarefaction curves, calculated from (2) [24], with a 95% confidence interval. eTh dotted vertical lines illustrate a species richness comparison standardized to 24 species and 221 individuals, which was the observed Odonate abundance in the smallest (rivers) of the three water types data set. There was no evident of signicfi ant relationship along axes associated with lentic (Coenagrionidae and Libellulidae) two and three. Following the CCA components, two main or lotic systems (e.g., Calopterygidae, Coenagrionidae, and groups of species were distinguished. The rfi st one (e.g., Libellulidae) [19, 49]. In this study, we observed similar pat- Urothemis edwardsii, Palpopluera lucia, Palpopluera portia, tern of association, where Calopterygidae, Chlorocyphidae, Rhyothemis notate, and Acisoma inflatum) was representative Platycnemididae, and Aeshnidae were found in lotic systems, of the pond community. This group was mainly composed of while Libellulidae and Coenagrionidae were found in both the generalist heliophilic species, which mostly avoid flowing lentic and lotic environments but showed strong affinity water (Figure 10). The second group was represented by to lentic systems (ponds). The presence of Calopterygidae the combined eeff ct of streams and rivers (e.g ., Chlorocypha and Chlorocyphidae exclusively in the lotic systems may be selysi, C. luminosa, Sapho ciliata, and Phaon camerunensis). explained by their strong ani ffi ty to canopied cover and fast The group was mainly composed of Zygopterans which were flowing water bodies, which were characteristics of streams favoured by fast o fl wing water. The only Anisopteran species and rivers in the Ankasa Conservation Area. These features found in group two was the Micromacromia zygoptera,which are well known to represent the preferred habitat type of was also inuen fl ced by fast flowing water body. most species within the Calopterygidae and Chlorocyphidae families [19, 49]. Species from the Aeshnidae family are crepuscular in 4. Discussion nature and are well noted to shun the sun during the day but to come to light at night [26]. This is confirmed in our Several studies have shown that majority of Odonata families and species from anisopterans and zygopterans are either study where most species from the family Aeshnidae showed Mean species richness Taxa (95% confidence) 10 International Journal of Zoology Stream River Pond Freshwater type Figure 7: Mean Odonata species richness among the various water types. 7.0 6.5 6.0 Hc = 1.661, p =0.44, Kruskal-Wallis test 5.5 5.0 4.5 4.0 3.5 3.0 0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 alpha diversity scale parameter Streams Rivers Ponds Figure 8: Renyi diversity ordering that compares Odonata evenness and richness of 1403 individuals, across the three water types. Note that the shape of a habitat profile is an indication of its evenness. a strong association with dense vegetation cover along the in several studies (e.g., [51, 52]) and have been linked to stream banks and utilized the vegetation for perching and higher colonization rate characterized by lentic systems [51, roosting during the day. Also, a large section of Ankasa and 52]. Such is the case observed among lakes in the Brazilian Bonwere rivers that were characterized by rocky substrates Atlantic Forest, where higher Anisoptera species richness was recorded [53]. But our findings rather revealed ponds appeared to support the perching, roosting, and copulating of some zygopterans like Mesocnemis singularis and this proba- to support the least abundance and species richness of bly explains their high abundance. Dijkstra and Clausnitzer damselflies relative to dragonflies which are composed of [25] and Dijkstra [26] reported that Mesocnemis singularis only species from the Libellulidae family. This was probably typically prefersunny rocky substrate, as ecological niches for due to the scale of environmental disturbance and the perching, roosting, and copulating. geographical location of the ponds. For instance, in the Several pond-associated species, such as the Ceriagrions, tropics where this study was conducted, extreme tempera- Agriocnemis species,A.inflatum, C. flavifrons, O. lugubris, T. tures and erratic rainfall in recent times could have wider arteriosa, and P. lucia, have been classified as Heliophilics or ramicfi ations on surface water temperatures of the ponds, stagnant water tolerance species [17, 25, 26], which concur which were beyond the thermal threshold tolerance of the with this current study. Though small in catchment area, species. Lentic environments tend to be geologically less pre- ponds supported several distinct species that were never recorded in other water types and contributed to the greatest dictable through time [54], and this phenomenon tended to Odonata assemblages (abundance and richness) compared exert pressure on species to adapt faster in order to be able to the lotic environments. Globally, these pond-associated to disperse and then persist [5]. Ponds are important refuge species from the families Libellulidae and Coenagrionidae for Odonata conservation because they are relatively isolated are composed of several ubiquitous species that dominate in and show greater heterogeneity in species assemblages [55, unshaded habitats with stagnant water bodies [50]. 56], owing to stochastic effects acting on the colonization Higher Odonata species richness and diversity in lentic process [57]. Variability in pond isolation has the tendency systems relative to lotic environments have been reported to attract good disperser Odonata such as species within Mean species richness Renyi diversity index International Journal of Zoology 11 Transform: Log(X+1) Resemblance: S17 Bray Curtis similarity S7 R3 C 1 R4 S5 S3 S2 C 2 SI S6 R1 C 3 R2 S4 R6 C 4 R5 P10 P4 P6 P9 C 5 P7 P3 P2 P1 P5 P8 0 20 40 60 80 100 Similarity Figure 9: Group average dendrogram of similarity of species composition for Odonata among the water types in the Ankasa conservation area. the Libellulidae family which are ye fl rs and heliothermic in This, however, indicates that Odonata fauna in the Ankasa nature [58]. Conservation Area are restricted to specific water types, Streams and rivers (i.e., typical lotic systems) in con- with each water body supporting some specific species or trast, which were characterized with dense vegetation cover, assemblages not found in other water types. This finding supported the greatest abundance and species richness of supports the importance of maintaining a diversified body of zygopterans relative to anisopterans, as a result of their water, both lentic and lotic, natural or artificial, in ecosystem association with dense vegetation cover along the fringes management to achieve the ultimate goal of conserving of the systems, which provide conducive environment for diverse Odonata fauna and other sympatric freshwater bio- resting, mating, and breeding. The suit of different micro- diversity. habitat complexity along these lotic systems continuum may have contributed in species heterogeneity, largely dominated Data Availability by the Zygopteran functional group. Streams and rivers worldwide have been reported to provide heterogeneous and The data used to support the findings of this study are favourable environmental conditions for diverse Zygoptera available from the corresponding author upon request. species, [55, 58, 59] for their numerous life activities including nocturnal roosting, oviposition, emergence, reproduction, and perching substrate to thermoregulate [55, 58]. Streams Conflicts of Interest and rivers also share similar characteristics linked to their The authors declare that they have no conflicts of interest. geomorphology and o fl w regimes [16]. These systems have extensive catchments as compared to other lentic systems and this dovetailed with similar geomorphological features, flow Acknowledgments rate, and uniform vegetation cover of the waters will ensure less variability in their physicochemical variables [16]. This The authors are grateful to Rufford Foundation (20322- may result in similar colonization and dispersion rate, which 1 and 2) and International Dragonfly Fund for providing may lead to higher overlap in their Odonata fauna as evident financial support for this study. Our heartfelt appreciation in this study. is due to Viola Clausnitzer and Klaas-Douwe B. Dijkstra It was not uncommon that none of the species occurred for providing us with the identification hand books and for in all the three water types which reinforced our hypothesis. their immense contribution, mentoring, and advice towards Sampling sites 12 International Journal of Zoology Ordination Plot −1 −2 −3 −4 −2 −1 0 1 2 Axis 1 Odonata species Sampling sites Figure 10: Canonical correspondence analysis (CCA) ordination diagram, showing the relationship between environmental variables and Odonata species across the three water types in the Ankasa Conservation Area in Ghana. Species names are abbreviated with the first four letters of the genus and the first four letters of the species following Seidu et al., [17]. The blue arrows represent each of the environmental variables plotted pointing in the direction of maximum change of explanatory variables across the water types. Red and green ovals represent specialists’ species in lentic (ponds) and lotic (rivers and streams), respectively. the successful completion of the study. Finally, the authors [6] G. P. Mishra and A. K. Yadav, “A comparative study of physico- chemical characteristics of river and lake water in Central are grateful to David Amaning Kwarteng, Daniel Acquah- India,” Hydrobiologia, vol.59,no. 3,pp.275–278, 1978. Lamptey, Sulemana Bawa, and Emmanuel Amoah, for their special role in eld fi data gathering. [7] W.J. Mitschand J.W.DayJr.,“iTh nkingbig with whole- ecosystem studies and ecosystem restoration - A legacy of H.T. Odum,” Ecological Modelling, vol.178,no.1-2,pp. 133–155, 2004. References [8] T. E. Essington and S. R. Carpenter, “Nutrient cycling in lakes and streams: Insights from a comparative analysis,” Ecosystems, [1] D. L. Strayer and D. Dudgeon, “Freshwater biodiversity con- vol.3, no.2,pp.131–143, 2000. servation: recent progress and future challenges,” Journal of the North American Benthological Society,vol.29,no.1,pp.344–358, [9] R. W.Merritt and K.W. Cummins,Eds., An Introduction to the Aquatic Insects of North America,Kendall Hunt, 1996. [2] V.Clausnitzer, K.-D.B.Dijkstra,R.Kochet al.,“Focus on [10] M. Vincy, R. Brilliant, and A. P. Kumar, “Checklist of odonata African freshwaters: Hotspots of dragonfly diversity and con- species as indicators of riparian ecosystem of a tropical river, servation concern,” Frontiers in Ecology and the Environment, the southern western ghats,” Journal of Entomology and Zoology vol.10,no.3, pp. 129–134,2012. Studies,vol.4,no.2,pp.104–108,2016. [3] C. J. Vor ¨ os ¨ marty, P. B. McIntyre, M. O. Gessner et al., “Global [11] P. S. Corbet, “Dragonflies: behaviour and ecology of odonata,” threats to human water security and river biodiversity,” Nature, Aquatic Insects, vol.23, no.1,pp.83-83, 1999. vol. 467, article no. 7315, pp. 555–561, 2010. [12] A. Dolny, ´ F. Harabiˇs, D. Bar ´ taa, S. Lhota, and P. Drozd, “Aquatic [4] C. Revenga and Y. Kura, Status and Trends of Biodiversity of insects indicate terrestrial habitat degradation: Changes in Inland Water Ecosystems,vol.11of Technical Series,Secretariat taxonomical structure and functional diversity of dragonflies in of the Convention on Biological Diversity, Montreal, Canada, tropical rainforest of East Kalimantan,” Tropical Zoology,vol.25, 2003. no. 3, pp. 141–157, 2013. [5] C.Hof, M.Bra¨ndle, and R. Brandl, “Latitudinal variation of [13] F. Harabiˇsand A. Dolny,´ “Human altered ecosystems: Suitable diversity inEuropeanfreshwater animals is not concordant habitats as well as ecological traps for dragonflies (Odonata): across habitat types,” Global Ecology and Biogeography,vol. 17, The matter of scale,” Journal of Insect Conservation,vol. 16, no. no.4,pp.539–546, 2008. 1, pp. 121–130, 2012. Axis 2 International Journal of Zoology 13 [14] V. Clausnitzer, “Dragonyfl communities in coastal habitats of [32] J. S. Gray and F. B. Mirza, “A possible method for the detection Kenya: Indication of biotope quality and the need of conserva- of pollution-induced disturbance on marine benthic communi- tion measures,” Biodiversity and Conservation, vol. 12,no.2,pp. ties,” Marine Pollution Bulletin, vol. 10, no. 5, pp. 142–146, 1979. 333–356, 2003. [33] A. E. Magurran, Measuring Biological Diversity,vol. 131, Black- well Science Publishers, Oxford, UK, 2004. [15] J. T. Bried, B. D. Herman, and G. N. Ervin, “Umbrella potential of plants and dragonflies for wetland conservation: A quanti- [34] S. Fattorini, “Relations between species rarity, vulnerability, and tative case study using the umbrella index,” Journal of Applied range contraction for a Beetle Group in a Densely populated Ecology,vol.44,no.4,pp.833–842, 2007. region in the mediterranean biodiversity hotspot,” Conservation Biology,vol.28,no.1,pp.169–176,2014. [16] P. Williams, M. Whitfield, J. Biggs et al., “Comparative biodi- [35] S. Fattorini, “A simple method to tfi geometric series and broken versity of rivers, streams, ditches and ponds in an agricultural landscapeinSouthernEngland,” Biological Conservation,vol. stick models in community ecology and island biogeography,” 115, no. 2, pp. 329–341, 2004. A cta Oecologica,vol. 28, no.3, pp. 199–205, 2005. [17] I. Seidu, E. Danquah, C. Ayine Nsor, D. Amaning Kwarteng, and [36] R. MacArthur, “On the relative abundance of species,” e American Naturalist,vol.94,no.874,pp.25–36, 1960. L. T. Lancaster, “Odonata community structure and patterns of land use in the Atewa Range Forest Reserve, Eastern Region [37] S. Fattorini, F. Rigal, P. Cardoso, and P. A. V. Borges, “Using (Ghana),” International Journal of Odonatology,vol. 20, no.3- species abundance distribution models and diversity indices for 4, pp. 173–189, 2017. biogeographical analyses,” Acta Oecologica,vol.70,pp. 21–28, [18] D. Acquah-Lamptey, R. Kyerematen, and E. O. Owusu, “Using Odonates as markers of the environmental health of water and [38] N. J. Gotelli and R. K. Colwell, “Estimating species richness,” in its land related ecotone,” International Journal of Biodiversity Biological Diversity: Frontiers in Measurement And Assessment, and Conservation, vol.5, no.11, pp.761–769, 2013. vol. 12, pp. 39–45, 2011. [39] A. F. Siegel, “Rarefaction curves,” in Encyclopedia of Statistical [19] K.-D. B. Dijkstra and J. Lempert, “Odonate assemblages of Sciences, K. Samuel, C. B. Read, N. Balakrishnan, and B. running waters in the Upper Guinean forest,” Fundamental and Applied Limnology ,vol.157, no. 3,pp.397–412,2003. Vidakovic, Eds., 2006. [40] A. R´enyi, “On measures of entropy and information,” in [20] I. Seidu, C. A. Nsor, E. Danquah, and L. T. Lancaster, “Odonata Proceedings of the Fourth Berkeley Symposium on Mathematics, assemblages along an anthropogenic disturbance gradient in Statistics and Probability,vol.1,pp.1286-1261, University of Ghana’s eastern region,” Odonatologica ,vol.47,no.1-2,pp. 73– California Press, Berkeley, Calif, USA, 1961. 100, 2018. [41] C. E. Shannon, “A mathematical theory of communication,” Bell [21] J. A. Fulan and R. Henry, “A comparative study of Odonata System Technical Journal,vol.27,no. 4,pp. 623–656, 1948. (Insecta) in aquatic ecosystems with distinct characteristics [42] B. Tothm ´ er ´ es ´ z, “Comparison of different methods for diversity Um estudo comparativo de Odonata (Insecta) em ecossistemas aquatic ´ os com distintas caracter´ısticas,” Ambiencia, vol.9,no.3, ordering,” Journal of Vegetation Science,vol. 6,no.2, pp. 283– 290, 1995. pp. 589–604, 2013. [43] Ø. Hammer, D. A. T. Harper, and P. Ryan, “PAST: paleontolog- [22] F. Dowsett-Lemaire and R. J. Dowsett, “An update on the birds ical statistics software package for education and data analysis,” of Kakum National Park and Assin Atandaso Resource Reserve, Palaeontologia Electronica, vol.4,no.1,article 4, 2001. Ghana,” A report prepared for the Wildlife Division, Forestry Commission 75, Dowsett-Lemaire Misc, Accra, Ghana, 2011. [44] J. R.Krebs,J. D. Wilson,R.B. Bradbury, and G.M. Siriwardena, “eTh second silent spring?” Nature,vol.400,article 6745,pp. [23] J. B. Hall and M. Swaine, “Classification and ecology of closed- 611-612, 1999. canopy forest in Ghana,” Journal of Ecology, pp. 913–951, 1976. [45] C. J. F. ter Braak, “Canonical correspondence analysis: a new [24] N. J. Gotelli and R. K. Colwell, “Quantifying biodiversity: eigenvector technique for multivariate direct gradient analysis,” procedures and pitfalls in the measurement and comparison of Ecology, vol.67,no.5,pp.1167–1179, 1986. species richness,” Ecology Letters, vol.4,no.4,pp.379–391, 2001. [46] P. A. Henderson and R. M. H. Seaby, Community Analysis [25] K. D. Dijkstra and V. Clausnitzer, e Dragonflies And Dam- Package ., Pisces Conservation Ltd, Lymington, UK, 2000. selflies of Eastern Africa: Handbook for All Odonata,2014. [47] C. J. F. ter Braak and P.F.M. Verdonschot, “Canonicalcorre- [26] K.-D. B. Dijkstra, “African dragonflies and damselflies online,” spondence analysis and related multivariate methods in aquatic 2017, http://addo.adu.org.za. ecology,” Aquatic Sciences, vol.57,no. 3,pp.255–289,1995. [27] J. H. Zar, Biostatistical Analysis, Prentice Hall, Upper Saddle [48] M. Kent and P. Coker, Vegetation Description And Analysis. A River, NJ, USA, 4th edition, 1999. Practical Approach, John Wiley and Sons Ltd, West Sussex, UK, [28] A. S. Melo and L. U. Hepp, “Ferramentas estat´ısticas para ana´lises de dados provenientes debiomonitoramento,” Oecolo- [49] K.-D. B. Dijkstra, “Dragonflies and damselflies (Odonata) of the gia Brasiliensis, vol.12,pp.463–486, 2008. Atewa Range. A rapid biological assessment of the Atewa Range [29] B. McCune and J. B. Grace, Analysis of Ecological Communities, Forest Reserve, eastern Ghana,” in RAP Bulletin of biological MjM Sow ft are, Gleneden Beach, OR, USA, 2002. Assessment,vol. 47, pp. 50–54, 2007. [30] K. R. Clarke and R. N. Gorley, “Primer V5 (Plymouth rou- [50] V. J. Kalkman, V. Clausnitzer, K. D. B. Dijkstra, A. G. Orr, tines in multivariate ecological research): user manual/tutorial. D. R. Paulson, and J. van Tol, “Global diversity of dragonflies Primer-E,” 2001. (Odonata) in freshwater,” in Freshwater Animal Diversity Assess- ment, pp. 351–363, Springer, Dordrecht, the Netherlands, 2007. [31] B. J. McGill, R. S. Etienne, J. S. Gray et al., “Species abundance distributions: moving beyond single prediction theories to [51] A. S. Niba and M. J. Samways, “Remarkable elevational toler- integration within an ecological framework,” Ecology Letters, ance in an African Odonata larval assemblage,” Odonatologica , vol. 10, no. 10, pp. 995–1015, 2007. vol. 35, no. 3, pp. 265–280, 2006. 14 International Journal of Zoology [52] L. E. Stevens and R. A. Bailowitz, “Odonata biogeography in the Grand Canyon ecoregion, southwestern USA,” Annals of the Entomological Society of America,vol.102,no.2,pp. 261–274, [53] S. Renner, E. Perico, and G. Sahlen, “Man-made lakes form species-rich dragonyfl communities in the Brazilian Atlantic Forest (Odonata),” Odonatologica ,vol. 45, no.3-4,pp. 135–154, [54] I. Ribera,G. N.Foster, and A.P.Vogler, “Does habitat use explain large scale species richness patterns of aquatic beetles in europe?” Ecography, vol.26,no. 2,pp. 145–152, 2003. [55] B.Oertli, D.A. Joye,E.Castella, R.Juge, D.Cambin,and J.-B. Lachavanne, “Does size matter? eTh relationship between pond area and biodiversity,” Biological Conservation,vol. 104, no. 1, pp. 59–70, 2002. [56] M. Scheffer, G. J. Van Geest, K. Zimmer et al., “Small habitat size and isolation can promote species richness: Second-order effects on biodiversity in shallow lakes and ponds,” Oikos,vol. 112, no. 1, pp. 227–231, 2006. [57] D. S. Jeffries, J. R. M. Kelso, and I. K. Morrison, “Physical, chemical, and biological characteristics of the Turkey Lakes Watershed, central Ontario, Canada,” Canadian Journal of Fisheries and Aquatic Sciences, vol.45,no.1,pp.s3–s13, 1988. [58] P. S. Corbet and M. L. May, “Fliers and perchers among Odonata: Dichotomy or multidimensional continuum? A pro- visional reappraisal,” International Journal of Odonatology,vol. 11, no. 2, pp. 155–171, 2008. [59] L. Maltchik,C. Stenert,C. B. Kotzian, and M. M. Pires, “Responses of odonate communities to environmental factors in southern Brazil wetlands,” Journal of the Kansas Entomologi- cal Society,vol.83, no. 3,pp.208–220, 2010. 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