Estimating distributional patterns of non-marine Ostracoda (Crustacea) and habitat suitability in the Burdur province (Turkey)

Accepted Manuscript Title: Estimating distributional patterns of non-marine Ostracoda (Crustacea) and habitat suitability in the Burdur province (Turkey) Author: Mehmet YAVUZATMACA Okan ¨ ˘ ¨ ¨ GLU KULK OYL UO Ozan YILMAZ PII: DOI: Reference:

S0075-9511(16)30146-3 http://dx.doi.org/doi:10.1016/j.limno.2016.09.006 LIMNO 25533

To appear in: Received date: Revised date: Accepted date:

1-3-2016 26-9-2016 29-9-2016

¨ ˘ ¨ ¨ GLU, Please cite this article as: YAVUZATMACA, Mehmet, KULK OYL UO Okan, YILMAZ, Ozan, Estimating distributional patterns of non-marine Ostracoda (Crustacea) and habitat suitability in the Burdur province (Turkey).Limnologica http://dx.doi.org/10.1016/j.limno.2016.09.006 This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

Estimating distributional patterns of non-marine Ostracoda (Crustacea) and habitat suitability in the Burdur province (Turkey)

Mehmet YAVUZATMACA*, Okan KÜLKÖYLÜOĞLU and Ozan YILMAZ

Department of Biology, Faculty of Arts and Science, Abant İzzet Baysal University, 14280 Bolu, Turkey

*Corresponding author: [email protected]

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ABSTRACT We explored distributional patterns and habitat preferences of ostracods in the Burdur province (Turkey). At 121 sites we recorded 35 taxa (22 recent, 13 sub-recent), of which 23 represent new records for the province. According to the Index of Dispersion and d-statistics, the individual species exhibited clumped distributions. Cosmopolitan species dominated (63.64%). A direct effect of regional factors (e.g., elevation) was not observed, while local factors (e.g., water temperature) best explained species distribution among habitats. Based on alpha diversity values, natural habitats (springs, ponds, creeks) were more suitable than artificial habitats (e.g., troughs, dams), suggesting that natural habitats define regional species diversity. Twenty-two of the recorded species had wider ecological ranges than previously reported. Cosmopolitan species appeared to suppress non-cosmopolitan species due to their wider ecological range.

Keywords: Bioindicator, Clump distribution, Dominant species, Ostracods, Source habitats

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1. Introduction

Ostracods are bivalved aquatic crustaceans that are generally small (0.3 - 5.0 mm, although some marine species may reach up to 30 mm in length) (Meisch, 2000). Their outer chitinous carapace has an epidermis of low magnesium calcium carbonate (calcite) that covers a calcitic shell which can be fossilized in sediments (Chivas et al., 1986). Fossil ostracods can be used to reconstruct paleo-environmental conditions. The first (oldest) undoubted fossil ostracod dates back to the Silurian period about 425 mya. These fossils represent the oldest known microfauna (Delorme, 1991; Siveter, 2008; Williams et al., 2008). Desiccation and freezing resistant eggs, and active and passive dispersal mechanisms contribute to their wide distribution throughout the world (McKenzie and Moroni, 1986; Horne and Martens, 1998; Rossi et al., 2003; Rodriguez-Lazaro and Ruiz-Muñoz, 2012; Külköylüoğlu, 2013) and in a variety of marine and non-marine aquatic habitats (Delorme, 1991; Meisch, 2000; Horne, 2003; Külköylüoğlu, 2013; Escrivá et al., 2014). The distributions of ostracods are effected by multiple factors such as temperature, sediment type, depth, vegetation, elevation, pH, dissolved oxygen, transparency of water and salinity (Malmqvist et al., 1997; Mezquita et al., 2001; Külköylüoğlu, 2005a; Martín-Rubio et al., 2005; Pérez et al., 2010; SzlauerŁukaszewska, 2012). Although species-specific responses to these factors (Benson, 1990; Delorme, 1991), some species are tolerant to a wide range of environmental conditions (e.g., water temperature, dissolved oxygen, etc.) (Uçak et al., 2014). Therefore, ostracods are bioindicators of aquatic conditions and are commonly used in different scientific fields such as geology (biostratigraphy), archeology, palaeobiology, palaeoclimatology, palaeolimnology, palaeoecology, wetland conservation, elemental and isotopes studies and evaluations of anthropogenic pollution (Forester, 1991; Holmes et al., 1998; Külköylüoğlu, 1998; Alvarez Zarikian et al., 2000; Mourguiart and Montenegro, 2002; Padmanabha and Belagali, 2008; Jiang et al., 2008; Sarı and Külköylüoğlu, 2010; Rodriguez-Lazaro and Ruiz-Muñoz, 2012; 3

Ruiz et al., 2013). Ostracods are particularly valuable as indicator species for estimating the past and present environmental changes. However, this requires an understanding and sophisticated knowledge of species-specific ecological requirements and tolerance ranges (limits) across habitats. Additionally, one may also question the type(s) of suitable habitats for ostracods and how ostracods respond to changes in such conditions (Külköylüoğlu, 2003a, 2004). The present study attempts to provide this understanding through a regional evaluation of ecology, distribution and habitat preferences of non-marine ostracods.

Species may exhibit random, clumped (aggregation) and uniform distributional patterns in response to biotic and abiotic factors. Of which, random distribution describes all individuals have equal probability of occurring in habitats. This distribution is also named as “Poisson distribution model” when population variance (s2) equals the mean (µ) (Ludwig and Reynolds, 1988; Zar, 1999). On the other hand, uniform and clumped distributional patterns show equal spacing and accumulation of species in an area and/or habitat, respectively (Ludwig and Reynolds, 1988). Determining the modes of these patterns can provide evidence to the effect of regional and local factors on species occurrence. While local factors (e.g., elevation) can influence the distribution of species residing in a particular habitat (e.g., lake, spring, etc.), regional factors can exert substantial influence on colonization and immigration of the species among the regions (Paradise et al., 2008). Determining type of such distributional patterns with those effective environmental variables on species distribution can help to protect species from extinction in particular areas. Despite this, there are no other extensive regional-scale geographical and ecological studies evaluating the distributional patterns of ostracod species (but see Yavuzatmaca et al., 2015).

Like most regions around the world, our knowledge about ostracod ecology and distribution in Turkey contains large gaps. For example, Burdur province (Fig. 1) has

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received no systematic survey and habitat characterization for its ostracod fauna. Here, we present the results of the first extensive study on Burdur province ostracods. Accordingly, the main objectives of the present study are i) to determine distributional patterns (clumped, uniform, random) of ostracod species in Burdur, ii) to discuss the relationship between habitat suitability and ostracod species diversity, iii) to elucidate the most important environmental factors (local and/or regional) affecting species distribution among habitats along, and iv) to estimate species’ ecological optimum and tolerance levels.

2. Material and Methods

2.1.

Site Description

The province of Burdur with 6887 km2 of surface area (also known as the ‘Lake District Area’) is located in the South Anatolia between 36°53' - 37°50' north latitude and 29°24' - 30°53' east longitude. The province is surrounded by some extentions of the West Toros Mountains in the south, Lake Burdur and the Karakuş Mountain in the north and Lake Acıgöl and Eşeler Mountain in the west. Also, the province has 2.7% upland, 19% lowland, 60.6% mountains and 17.6% hilly lands (Burdur Valiliği, 2014). High mountains seperate the district from the Mediterranean region, and summer is hot but winter is very cold (Burdur, 2014).

2.2.

Sampling and Measurements

Total of 121 sampling sites with six different aquatic habitats (spring, lake, dam, pond, creek and trough) were randomly visited and sampled between August 30 and September 02, 2012 (Fig. 1). Sampling sites were 5 to 10 km apart to prevent bias on similarities in species diversity and distribution. Eight environmental variables (dissolved oxygen (DO, mg L-1), percent oxygen saturation (% sat.), water temperature (Tw, ºC), electrical conductivity (EC, 5

µS cm-1), total dissolved solids (TDS, mg L-1), salinity (Sal, ppt), pH, atmospheric pressure (mmHg)) were recorded with a YSI-Professional Plus before sampling to prevent possible results of “Pseudoreplication” (Hurlbert, 1984). In situ water phsyico-chemical measurements should be taken without any disturbance of the sampling site that can result from ostracod collection and subsequent increased turbidity and water column mixing. Air temperature (Ta, ºC), wind speed (km h-1) and air moisture (%) were obtained by a Testo 410-2 model anemometer, and basic geographical data (elevation, coordinates) were recorded with a geographical positioning system (GARMIN etrex Vista H GPS) (Appendix A).

Ostracod samples were collected from each site with a standard sized hand net (200 µm mesh size). We preserved samples in 250 ml plastic bottles and fixed with 70% of ethanol. In the laboratory, each sample was filtered through four standardized sieves (0.5, 1.0, 1.5 and 2.0 mm mesh size) under tap water and than kept in 70% ethanol for further studies following standard protocol (Danielopol et al., 2002). Subsequently, ostracods were sorted from sediments under a stereomicroscope (Olympus ACH 1X) and their soft body parts were dissected in lactophenol solution for taxonomic identification. Species identification was done using a light microscope (Olympus BX-51). The taxonomic key provided in Meisch (2000) was primarily used for taxonomic classification and species identification, although additional taxonomic keys (e.g., Bronhstein, 1947; Karanovic, 2012) were also used when necessary. All of the ostracod samples were curated at the Limnology Laboratory of Abant İzzet Baysal University, Bolu/TURKEY and are available upon request.

2.3.

Statistical Analyses

Distributional patterns of species among sampling sites were tested by the application of Poisson probabilities along with a Chi-square test (Ludwig and Reynolds, 1988). The observed number of sites (f) with 0, 1, 2, 3, 4 or more (4+) species was computed (Fig. 2). 6

Then, the mean (µ) was then calculated by multiplying the number of species (i.e., 0, 1,...,4 +) by f, then dividing by the total number of sampling sites. The Poisson probability of finding of x individuals in sampling units (P(x)) was calculated using Equation 1.

Equation 1 where e, represents Euler’s number and approximately equals 2.71828; µ, the mean number of successes that occur in a specified region; x, the actual number of successes that occur in a specified region; P(x; µ) the Poisson Probability that exactly x successes occur in a Poisson experiment, when the mean number of successes is µ.

A Chi-square (X2) test was then applied to compare observed (O) and expected (E) frequencies of random distribution at 0.05 critical (α) level. The value of calculated Chisquare was computed using Equation 2.

Additionally, departure from Poisson distribution was tested by application of the Index of dispersion (variance (s2) / mean (µ)) (Ludwig and Reynolds, 1988) where s2/µ = 1 random distribution, s2/µ < 1 = uniform distribution, and s2/µ > 1 = clumped distribution.

To test whether data conformed to the Poisson distribution when the sample size is N30, we calculated d-statistics (Equation 4). To find d-statistics value, we need Chi-square (X2) (it is different from X2 calculated using Equation 2) calculated using Equation 3 (Ludwig and Reynolds, 1988).

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where X2: Chi-square, Xi: the number of individuals in the ith sampling unit; N: total sample size; X : the mean number of successes that occur in a specified region

Equation 4

where X2 is derived from Equation 3

The d-statistics (Elliott, 1973 sensu Ludwig and Reynolds, 1988, p. 28) are interpreted as follows: i) if ΙdΙ < 1.96, accept a random dispersion ii) if d < -1.96, suspect a regular dispersion iii) if d > 1.96, a clump dispersion

The relationships between species and environmental variables (electrical conductivity (EC), water (Tw) and air (Ta) temperatures, dissolved oxygen (DO), elevation (Elev) and pH) were examined by Canonical Correspondence Analysis (CCA). The data were logtransformed (ter Braak, 1987; Birks et al., 1990) and tested with Monte Carlo Permutation tests (499 permutation) where rare species were removed before analyses. Before performing CCA, suitability of data for CCA was tested with Detrended Correspondence Analysis (DCA) (software package CANOCO for windows 4.5).

C2 software was used to estimate species tolerance (tk) and optimum (µk) levels for different ecological variables after using a transfer function of weighted averaging regression (Juggins, 2003). The accuracy of optimum estimates is proportional to species’ prevalence in 8

samples (ter Braak and Barendregt, 1986). Therefore, the optimum and tolerance levels of a species to ecological variables can show differences according to their occurrence frequencies in different geographical areas as this is the case in Burdur and in many other regions in and out of Turkey (see discussion below). This situation may be considered for further evaluation of the ideas about ecological preferences of individual species and using them as indicators of environmental conditions.

The software program Species Diversity & Richness 4 (Seaby and Henderson, 2006) was used to calculate the Shannon-Wiener index value for different habitat types. The range of 1.5 and 3.5 was used to consider low to high index values as suggested by Magurran (1988). Species were classified as eudominant (32 – 100%), dominant (10 – 31%), subdominant (0.32 – 9%), recedent (1 - 3.1%), subrecedent (0.32 - 0.99%) and sporadic (< 0.31%) based on their dominance coefficient (Rombach, 1999)

3. Results

A total of 35 taxa (22 recent and 13 sub-recent) were encountered from 110 of 121 sampling sites in Burdur province. Of these, 23 (10 living and 13 sub-recent) taxa represents new records for Burdur (Appendix A). The occurrence probabilities of P (x = 0), P (x = 1), P (x = 2), P (x = 3) and P (x = 4(+)) of species in Burdur were calculated using Equation 1 and given in Table 1. Chi-square (X2) values of each occurrence (0, 1, 2, 3 and 4 (+)) are presented in Table 2. Ostracod occurrence did not conform to a random distribution or “Poisson distribution” 9

(X2calculated = 8.08 > X2table (3,121,0.05) = 7.81), Index of Dispersion and d-statistics (1.09 and 2.51, respectively) suggested a clump dispersion of ostracod species among sampling sites in Burdur.

The first two axes of the CCA explained 78.80% of relationships between 13 species and six environmental variables with a moderately low variance (9.50) (Table 3).

Among the variables, water temperature (Tw) (F = 3.746, P = 0.002) was the only one with strong effect on species followed by non-significant pH (F = 1.753, P = 0.074), dissolved oxygen (DO) (F = 1.494, P = 0.144), air temperature (Ta) (F = 1.170, P = 0.306), elevation (Elev) (F = 1.021, P = 0.406) and electrical conductivity (EC) (F = 0.4920, P = 0.610) (Fig. 3). Six species (Candona neglecta, Psychrodromus olivaceus, P. fontinalis, Cypria ophtalmica, Ilyocypris bradyi and Prionocypris zenkeri) were located at the left site of first axis where there is only elevation but three species (Limnocythere inopinata, Ilyocypris monstrifica and Physocypria kraepelini ) were located at the site of pH, EC and Ta. The other species (Heterocypris incongruens, H. salina, Potamocypris variegate and Herpetocypris intermedia) were placed at the site of Tw and DO. Two well known cosmoecious species (see discussion for definition) (H. incongruens and I. bradyi) were relatively closer to the center of diagram (Fig. 3).

There was no apparent relationship between species richness and elevation (Fig. 4). The number of sampling sites did not affect species richness at different elevational ranges (e.g., see 886 - 1036 m and 1339 - 1489 m of ranges) where species richness (8) were the same despite differences in the numbers of sites (23 and 11, respectively). Accordingly, there was no clear correlation between numbers of sites and numbers of species encountered within the elevational ranges. Sexually reproducing species were encountered more frequently than asexually reproducing species when elevation increased (Fig. 4). 10

Cosmopolitan species generally displayed relatively higher tolerance and optimum values for environmental variables than other species (Table 4). For example, H. incongruents, H. salina and I. bradyi had the highest tolerance levels for water temperature and L. inopinata had higher tolerance level for pH and electrical conductivity but I. monstrifica had highest tolerance level for pH. In contrast, C. neglecta shows higher than mean optimum values for dissolved oxygen concentration when P. variegata had highest optimum value for dissolved oxygen concentration. These results suggest species-specific tolerance (tk) and optimum (μk) levels of individual species for different environmental variables. Shannon-Wiener index values of ponds (H′ = 2.25), springs (H′ = 1.79) and creeks (H′=1.71) were higher than other sampled habitats (Table 5). Although troughs were sampled more frequently (58 sites with 10 spp.) than the other types of habitats, species richness are higher in ponds (17 sites with 15 spp.). In other words, a clear habitat effect was observed despite differences in sampling effort. Four cosmopolitan species (H. incongruens, H. salina, I. bradyi and P. olivaceus) were dominant over the sub-dominant species (C. neglecta, Cypria ophtalmica, Herpetocypris chevreuxi, H. intermedia, I. monstrifica, L. inopinata, P. variegata and P. zenkeri) while another 10 species were defined as sporadic (Table 5).

Although species abundance (number of individuals) was much higher in troughs than other habitat types (Table 6), species richness per site were high in the other five habitats. Among the habitats, spring and creeks exhibited equal species richness (7) with almost similar numbers of individuals.

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4. Discussion Until now, 24 living species (Gülen, 1985; Altınsaçlı, 2004; Rasouli et al., 2014) have been recorded from Burdur, Turkey. 12 of these were encountered during the present study. Additionally, 23 (10 living and 13 sub-recent) taxa herein are new reports for the region (Appendix A). Thus, 47 ostracod species have been recorded from Burder, to date. Consequently, ostracod diversity in Burdur represents an important region for ostracod diversity in Turkey, comparable to other regions around the world in which systematic surveys have been conducted. For example, 37 taxa have been reported from 114 sites in Çankırı (Külköylüoğlu et al., 2016), 41 taxa have been reported from 111 sites in Adıyaman (Yavuzatmaca et al., 2015), 25 taxa have been reported from 50 sites in Diyarbakır (Akdemir and Külköylüoğlu, 2011), 34 taxa have been reported from 133 sites in Ordu (Külköylüoğlu et al., 2012c), 14 species have been reported from 38 sites in northern Finland (Iglikowska and Namiotko, 2010), 74 taxa have been reported from 320 sites in north east Italy (Pieri et al., 2009), and 54 species have been reported from 132 sites in upper Paraná River, Brazil (Higuti et al., 2009). Accordingly, numbers of species or taxa do not correspond to increasing sampling effort (the Sampling Effect Hypothesis) (Williamson, 1988; Hill et al., 1994). Rather, ostracod diversity patterns in Burdur may be better explained by the "Habitat Diversity Hypothesis" (Williams, 1943) which predicts that species richness increases with the availability and diversity of suitable habitats (see Külköylüoğlu et al., 2012a). Consequently, the numbers of ostracod species or taxa in an area may be related to variety of habitat types and habitat quality as is the case in our study.

The clumped distribution patterns exhibited by Burdur ostracods mirrors the distributional patterns of several other taxonomic groups. For example, benthic populations (Heip, 1976), marine benthic (Heip, 1975) and some sessile invertebrates (Schmidt, 1982)

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also exhibited clumped distributions. For ostracods, Heip (1976) observed a clumped distributional pattern of Cyrideis torosa (Jones, 1850) in a brackish pool in northern Belgium. Conversely, Yavuzatmaca et al. (2015) identified random distributional patterns of ostracods among sampling sites in Adıyaman (Turkey). These opposing findings may be explained by differences in the types of habitats sampled and the prevalence of non-cosmopolitan species. Specifically, Yavuzatmaca et al. (2015) mostly sampled natural springs (n = 44), where noncosmopolitan species were mostly prevalent (ca 51.85% of all species). In Burdur, habitat destruction was clearly observed where natural spring have been transformed into troughs. This habitat degradation may result in the aggregation of ostracod species in the troughs that we sampled (n = 58). Species inhabiting troughs have decreased opportunities for free distribution. These habitats are generally constructed to store water for animals, for irrigation, and for cleaning and drinking purposes (Külköylüoğlu et al., 2013). In such habitats, a direct anthropogenic effect that alters physico-chemical characteristics of water sources is apparent. Dominance of cosmopolitan species in degraded habitats is well documented in the literature (Külköylüoğlu et al., 2013). This is probably the case in Burdur where the majority of sampling sites were troughs with a higher prevalence of cosmopolitan species. If we consider troughs as a source of point diversity (or diversity at a single point or microenvironment (Meffe and Carroll, 1997)), this may explain the way of aggregated distribution of the ostracods in Burdur.

Canonical correspondence analysis showed that only water temperature significanlty affected ostracod dispersion (Fig. 3). Several previous studies similarly identified water temperature as an important factor affecting ostracod assemblages (e.g., Malmqvist et al., 1997; Viehberg, 2006; Kiss, 2007; Külköylüoğlu et al., 2014; Uçak et al., 2014). Since many (if not all) aquatic physico-chemical variables (e.g., electrical conductivity, dissolved oxygen concentration) are affected by changes in temperature (Wetzel, 2001), such changes may 13

in/directly alter the species occurrences. Unlike our current study, Külköylüoğlu and Sarı (2012) identified pH as the most important predictor of ostracod assemblage structure in a variety of aquatic bodies in Bolu, Turkey. Like water temperature (Roca and Wansard, 1997; Xia et al., 1997; Palacios-Fest and Dettman, 2001; Elmore et al., 2012), pH exerts an important control on the calcification of ostracod valves because solubulity of carbonate and calcium in water is dependent on pH (Wetzel, 2001). In this case, fluctuations in water physico-chemical variables (e.g., temperature) can have severe impacts on species with low tolerance ranges (stenophiles). However, species with a high tolerance to different variables within a large geographical distribution or “cosmoecious” (Külköylüoğlu, 2013)” (I. bradyi, H. incongruens and P. zenkeri) are relatively closer to the center of diagram (Fig. 3), implying that such environmental variables may not have critical influence on the distribution of these species.

There is still debate about the effect of elevation on ostracod species richness and distribution. Stevens (1992) stated that species richness and elavations are negatively correlated. Mezquita et al. (1999a) argued that elevation was a limiting factor for ostracod distribution in different water bodies of Spain. Likewise, Pieri et al. (2009) noted the importance of altitudinal range as a critical factor for the distribution of freshwater ostracods in regional scale in Italy. Also, Poquet and Mesquita-Joanes (2011) stated that elevational range may increase regional diversity in warm or temperate climates because water temperature, alkalinity and conductivity values of aquatic bodies are high at low elevation. Other studies, however, did not support (Malmqvist et al., 1997; Külköylüoğlu et al., 2012a, 2012b; Guo et al., 2013) these previous statements. In the present study elevation did not appear to influence species richness although the effect was not statistically examined. For example, we found the same numbers of species (8 spp.) at 886 - 1036 m and 1339 - 1489 m a.s.l. ranges (Fig. 4). Similarly, elevation was not identified as a significant variable in CCA. 14

This does not mean that elevation does not have potential to affect ostracod distribution, particularly through its’ influence on water temperature and other aquatic physico-chemical variables (Brown and Gibson, 1983; Van der Meeren et al., 2010; Reeves et al., 2007; Rogora et al., 2008). Such kind of changes in waters may affect the occurrence and distribution of species. Species with wide tolerance levels to different environmental variables may have better adaptive values to these changes. This is probably the case in the present study (see Fig. 4). It seems that local factors (e.g., water temperature) are more effective than the regional factors (see CCA diagram, Fig. 3) on species occurrence and distribution where elevation may have indirect effect on species distribution.

The optimum and tolerance levels reported here for Burdur ostracod species differ from those reported for ostracods from different geographical areas (Karakaş Sarı and Külköylüoğlu, 2008; Külköylüoğlu et al., 2013; Rasouli et al., 2014; Uçak et al., 2014), probably because of differences in occurrence frequencies and abundances among ostracods assessed in analyses. Sampling adequacy should be evaluated before drawing conclusions about the ecological preferences of individual species. Knowledge about species-specific tolerance and optimum levels is important for understanding the life history of ostracods, using them in palaeoreconstruction studies, understanding the environmental changes and also important for using ostracod species as bioindicators. Because of the increased sampling effort conducted in this study, the ecological preferences to eight different environmental variables for 22 ostracod species were all wider than previously reported (Appendix B). Hence, use of living species as bioindicators should be done carefully.

As mentioned above, numbers of species (15 spp.) was higher in ponds than the other habitat types. However, dominancy of individuals per species (abundance) was clearly on the side of troughs where the conditions favor cosmopolitan species with higher tolerance and

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optimum levels over non-cosmopolitans (Table 6). This is indeed the case in our samples where we found H. incongruens, a well-known cosmopolitan (also see cosmoecious species concept), in 31 of 58 troughs (Table 5). It appears that H. incongruens increases advantages over other species by means of increasing its abundance in such artificial habitats (i.e., troughs). Aquatic conditions are changeable in troughs where cosmopolitan species can tolerate. Accordingly, they show dominancy in numbers (Table 6).

According to alpha diversity index values, natural habitats (springs, ponds and creeks) were apparently more suitable than those of artificial (e.g., troughs, dams, etc.) habitats (Table 5). Another study done in Kahramanmaraş (Turkey) by Külköylüoğlu et al. (2012a) showed partially similar results with our findings as limnocrene springs (H′ = 2.89), ponds (H′ = 2.2) and creeks (H′ = 1.95). In addition, the high values of Shannon-Wiener Index were also found for ponds (H′ = 1.25), springs (H′ = 1.00) and creeks (H′ = 0.71) in Zonguldak and Bartın (Turkey) (Külköylüoğlu, O. pers. comment). However, species diversity of lakes (3.0 species per sample (sps)), springs (2.8 sps) and ponds (2.1 sps) noted by Van der Meeren et al. (2010) in western Mongolia are higher than the present study (ponds (0.88 sps), springs (0.70 sps) and lakes (0.44 sps) (Table 6)). Such differences may be exlained by several factors such as sampling time, number of sampling sites and geographical differences. In our case, we collected samples in a short time in one season which may cause for such differences. Among the habitats, springs are known as natural biological laboratories with stable ecological conditions (Forester, 1991) if there is no disturbance. Having relatively stable ecological conditions, springs provide different opportunities for organism. Hence, springs have good conditions for ostracod diversity especially for noncosmopolitans. Unlike springs, creeks have spatial and temporal heterogeneity in physico-chemical and biological features. This is possibly because of different water sources merging into the creeks by means of precipitation and dissolved and/or particulate matters produced from drainage basin of flowing waters 16

(Wetzel, 2001). As stated by Lansac-Tôha et al. (2004), lotic habitats are the connection between the lentic habitats (e.g., especially open lakes) and so their fauna comes from all types of lentic habitats. Therefore, this eventually increases heterogeneity of creeks where microhabitats will provide alternative places for organism. Thus, this will make important changes in species richness and biodiversity. Conversely, there can be some negative effects of flowing waters on species diversity. For example, large flood events negatively effect the flowing water environments because they disturb all substrate types and the species as well. All this information may allow us to express the high diversity index value of creeks in the present study. Last habitat that we can consider as suitable for ostracods is “ponds”. Céréghino et al. (2012) pinpointed that biogeographic turnover is higher in ponds than in other freshwater bodies for freshwater species. Similarly, Martín-Rubio et al. (2005) stated most continental ostracods inhabit the stable water of lakes and ponds. In addition, the expression of Marmonier et al. (1994) may be used to elucidate high diversity of ostracods in ponds in the present study as they stated that temporary ponds have species with short life spans, desiccation resistance and tolerance, high migratory ability and have spherical or cylindrical body shape. The life spans of ostracods (e.g., Cypridopsis vidua reach to maturity in 45 days) that change species to species (Delorme, 1991) and the body shape of them (e.g., kidney, bean, elliptical, etc., (for more see Karanovic, 2012) imply why species diversity of ostracods are high in ponds. The well known dispersion abilty of ostracods by birds is another important event for explanation of ponds that are suitable or may be called as source habitats for ostracods. This is because ponds are the stepping stones for migration, dispersion and genetic exchange especialy for wild species (e.g., birds) (Céréghino et al., 2014) and so the eggs and ostracods attached to their feathers disperse large distances in different geographical areas and habitats on the path of migratory birds. If suitable conditions and source habitats for ostracods 17

are known, we may have a chance to prevent their extinction. Also, the number of similar studies should be increase in the world and in Turkey for ostracods and for other taxonomic groups since they are important for the protection of biodiversity. Over all habitats, such as ponds, springs and creeks may be called as suitable (or source) habitats for ostracods but this view should be tested in different geographical areas in future.

5. CONCLUSION

The number of taxa in Burdur were increased up to 47. Dominant species were generally cosmopolitans when they have higher abundance values than non-cosmopolitans in the present study. In other words, they seem to suppress the occurrence of non-cosmopolitans. Accordingly, they show aggregational pattern among the habitats where they occurred. As a result, frequent occurrences of cosmopolitans in aquatic bodies may be related the quality of these habitats, usually showing tendency for decrease. These conditions should be carefully monitored and should be taken under consideration for the protection of biodiversity. The sampling of natural habitats (for non-cosmopolitans), frequent presence of cosmopolitans and dispersal ability of ostracods may contribute to and be used as a way of explaining spatial patterns of ostracods among sampling sites. In spite that some of ponds, springs and creeks are under human impact, they may be showed as source habitats for ostracods. This should be tested in the future. As seen in the present study, the local factors (e.g., water temperature) are more effective than regional factors (e.g., elevation) on distribution of ostracods. Finally, ecological information was gathered from literature and from the present study of each species in Appendices A and B and are very important for using ostracods as bioindicators and for palaeoenvironmental reconstruction studies.

18

Acknowledgement

We would like to thank Daniel Hering (Germany) for his constructive review and comments on the earlier version of this manuscript and two anonymous reviewers. We also thank to Dr. Randy Gibson (USFWS, Texas) and Dr. Benjamin T. Hutchins (TPWD, Texas) for their comments and suggestions on English of the first draft of this study. Special thanks go to Mrs. Sinem YILMAZ for her help during field studies. This study is funded by the Scientific Project Research Agency of Abant İzzet Baysal University (Project no: 2012.03.01.534).

This

is

a

part

of

19

Ph.D.

dissertation

of

M.Y.

Appendix A. Ecological variables and taxa were reported from different aquatic bodies in Burdur. St. no

St. Ty

1

6

2

1

3

5

4

6

5

4

6

6

7

6

8

1

9

4

10

6

11

6

12

4

13

6

14

6

15

4

16

6

17

6

pH 7.2 4 7.2 5 7.1 3 7.8 6 7.8 3 7.6 2 7.9 6 7.1 5 7.9 3 8.5 1 8.2 3 8.6 2 8.1 9 8.1 1 7.8 9 7.9 9 7.7

DO

%D O

8.7

79.5 496.7

EC

Sp. EC

Sal

659.7

0.32

9.3

87.3 512

670

0.33

5.77

55.1 480.8

620

0.3

8.53

88.6 451.7

530.2

0.26

8.66

84.6 355.3

450.5

0.22

11.5 5

122. 665 3

760

0.37

8.94

91.9 279

328

0.16

4.55

43.3 581

752

0.37

7.39

71.5 603

766

0.38

8.41

81.5 181.3

228.8

0.11

9.85

89.5 179.4

245

0.12

8.78

96.7 347.2

386.5

0.19

11.3 3

135. 483 6

486.7

0.23

7.11

83.5 677

703

0.34

2.65

34.4 1390

1296

0.64

9.04

98.9 536

598

0.29

10.9

133. 525

513

0.25

Tw Ta 12. 1 12. 6 13. 2 17. 3 13. 9 18. 9 17. 1 13. 1 13. 9 14. 1 11 19. 8 24. 6 23. 1 28. 6 19. 8 26.

TDS

Atm.

Moist W. . s.

25.6 0.429 685.7 26.7

2.2

0.435 23.6 680.5 26.3 5

2.7

22.8 0.403 683

5.5

27.2

0.345 23.6 671.5 20.9 2 0.294 27.2 661.7 21 4 28.4 0.494 669.2 19.2 30.8

0.213 2 0.487 5 0.500 5 0.148 9 0.159 2 0.250 9 0.317 2

4.3 6.5 2.5

664.2 19.5

5.1

665.5 20.2

3.5

665.1 23.5

1.6

635.4 25.2

4.5

635

22.9

4.9

671.2 19.4

3.2

684.3 13.7

5.5

31.3 0.455 672.5 16.9

2.5

0.838 31.7 681.2 12.6 5

3.2

33.2 0.39

675.1 14.4

8.2

31.8 0.331 685.1 28.7

3.9

26.7 28 26.7 28.3 30.5 33.3

20

Elev Coordinate . N37°31'837'', 855 E036°02'383'' N37°38'791'', 921 E030°41'594'' N37°38'107'', 907 E030°42'167'' 105 N37°38'569'', 0 E030°35'921'' 118 N37°39'551'', 1 E030°35'551'' 108 N37°38'919'', 3 E030°33'190'' 113 N37°38'926'', 9 E030°30'906'' 112 N37°38'581'', 4 E030°30'393'' 112 N37°38'388'', 3 E030°30'493'' 153 N37°40'638'', 8 E030°31'080'' 152 N37°40'694'', 1 E030°31'196'' 108 N37°37'626'', 3 E030°31'253'' N37°32'555'', 882 E030°31'127'' 106 N37°34'509'', 1 E030°28'308'' N37°33'680'', 946 E030°28'235'' 102 N37°34'564'', 7 E030°26'708'' 898 N37°31'227'',

Date 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012

Taxa Po; (Isp) (Hsp; Isp; Psp; Pysp) Cn; Pzi; (Isp) Hi (Csp; Isp; Pzi) Hi; Ib (Esp; Hts; Hsp; Isp) Pfo; (Isp) Cn; Ib; Pf; (Pzi) Po Po; (Hsp) (Isp) Hi; Pv Hin; Hi

30.08.2012 30.08.2012

Hi; Ib; Pzi; Po 30.08.2012 Hi; Ib; Pv

18

5

19

6

20

6

21

6

22

3

23

1

24

6

25

1

26

6

27

3

28

6

29

3

30

6

31

6

32

6

33

4

34

4

35

5

36

6

6 8.0 1 8.0 6 8.0 6 7.9 8 8.7 6 7.3 6 7.7 4 7.8 2 7.8 4 8.1 3 7.3 5 8.3 8 7.3 9 7.6 7.3 3 8.0 8 7.8 2 7.6 8 7.4

3 3.2

40.4 800

762

0.37

11.5 7

138. 487 5

493.7

0.24

9.35

92.1 303.8

370

0.18

8.86

85.7 288

364.8

0.18

5.6

67.9 327.7

326.6

0.15

4.77

48.2 978

1176

0.59

7.63

76.5 472.2

579.4

0.28

7.95

73.3 290.6

367

0.18

8.94

89.4 301.6

365.2

0.18

6.6

72.4 351.4

388

0.19

8.26

103. 623 1

603

0.29

5.04

62

6.09

344.7

331.4

0.16

71.2 615

649

0.32

5.94

66.7 597

650

0.32

5.71

59.8 590

685

0.34

7.08

66.8 672

874

0.43

4.07

41.6 324.1

387.8

0.19

9.1

83.7 289.5

390.9

0.19

7.95

92.3 1068

1130

0.56

3 27. 8 24. 2 15. 8 13. 9 25. 2 16. 2 15. 3 14. 2 15. 9 20. 1 26. 6 27. 1 22. 3 20. 6 17. 7 12. 9 16. 4 11. 4 22

5 32.9 0.494 684.3 13

2.7

906

683.9 12.8

2.4

878

682.4 13.6

3.6

904

681.9 14

0

905

686.5 22.2

0

843

32.8 0.767 684.8 14.3

5.5

907

33.4 0.377 688.4 17.5

3.3

813

0

637

0

460

0

287

1.5

348

2.4

282

2.2

443

1.5

792

0

792

691.2 42.3

0

808

693

43.3

0

785

690.2 50.5

2

815

690.6 33.4

4.6

815

35.7 33.6 32.9 34.4

0.321 1 0.241 2 0.237 9 0.211 9

0.239 32.1 707.5 23.7 2 0.237 31.4 721.4 26.5 3 0.252 30.3 737.8 24.8 2 30.5 0.39

728.8 21.8

0.215 30.7 732.2 21.7 2 0.422 30.1 769.9 21.2 5 0.422 12.4 692.8 54.2 5 13.8 0.442 692.5 51.7 17.1 16.4 17.4 18.9

0.565 5 0.252 2 0.254 2 0.734

21

E030°28'277'' N37°30'531'', E030°27'003'' N37°29'825'', E030°26'687'' N37°30'527'', E030°25'110'' N37°30'453'', E030°24'693'' N37°30'715'', E030°32'463'' N37°30'156'', E030°33'286'' N37°28'623'', E030°32'090'' N37°18'918'', E030°46'278'' N37°19'051'', E030°48'237'' N37°20'007'', E030°48'767'' N37°22'519'', E030°48'876'' N37°22'239'', E030°49'626'' N37°23'364'', E030°47'747'' N37°26'745'', E030°29'863'' N37°26'310'', E030°29'447'' N37°25'078'', E030°26'957'' N37°25'205'', E030°25'556'' N37°25'859'', E030°23'902'' N37°23'448'',

30.08.2012 30.08.2012

(Hsp; Pos) Hi; Pv; Po; (Isp)

30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012 30.08.2012

(Csp; Im) Cn; (Hsp; Isp; Psp) Ib; Po; (Pzi) Cn Po (Hsp; Isp)

30.08.2012

Hi; Po Im; Li; Pk; (Hsp; Pos; Psp; 30.08.2012 Pysp) 30.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012

Hin; Hi; Ib (Hsp; Pzi) Hi; Ib Cn; Ib; Im; Tc; (Hsp; Psp)

Pfo 31.08.2012 Hs

37

4

38

6

39 40

6 6

41

4

42

5

43

4

44

5

45

3

46

6

47 48 49

4 5 1

50

6

51

4

52

5

53

4

54

4

55

6

1 8.1 6 7.4 2 7.9 4 8.1 8.0 3 8.3 1 8.1 6 8.1 3 8.3 8 7.0 5 8.2 1 8.5 1 7.8 2 8.1 5 8.6 5 8.5 1 8.2 8.4 5 7.6

9.53

92.6 612

767

0.38

7.6

74.4 832

1036

0.52

9.3

100

372.5

429.7

0.21

7.42

67.6 338.6

435.7

0.02

6.35

64.7 1824

2181

1.12

7.58

84.5 466.4

509.1

0.25

11.2 9

130. 420.9 4

442.9

0.21

6.96

76.3 525

586

0.29

7.02

81.5 352.8

371.2

0.18

4.54

48.7 1010

1158

0.58

5.95

66.2 346.9

381.3

0.18

9.74

110. 658 7

705

0.34

4.7

59

556

532

0.26

9.5

110. 407.1 4

421.7

0.2

9.5

96.3 198.1

235.4

0.11

5.88

72.9 395.6

386.6

0.18

4.16

49.7 329.6

338.2

0.16

5.77

69.1 491.4

492.1

0.24

7.18

73.2 497.7

608.3

0.3

14. 5 14. 8 18 12. 5 16. 5 20. 6 22. 5 19. 6 22. 4 18. 3 20. 3 21. 5 27. 2 23. 1 16. 7 26. 2 23. 7 24. 9 15.

5 0.487 21.4 691.3 28.5 5 24

0.676 684.8 28.7

0.282 25.6 672.3 24.7 1 0.270 25.6 651 21.9 4 26

1477

0.330 8 0.286 27.9 7 0.383 31.2 5 0.241 27.1 2 27

0

810

0

884

1.6 7

674.2 23.5

5

674.6 35.4

1.6

663.5 20.5

6.1

663.9 29

1.6

660.7 26

12

103 3 131 0 103 2 103 3 117 1 117 2 119 4 112 9 103 8

31.7 0.754 665.8 22.3

1.6

0.247 32.5 672.3 20.1 7

2.5

29.7 0.455 676.1 18.9

17.5 990

32.3 33.1 30.5 31.5 30.5 33 33.8

0.344 5 0.274 3 0.153 4 0.251 6 0.219 7 0.320 4 0.395

676.1 17.8

1.8

671.1 15.8

2.7

661.8 16.3

0

655.5 22.2

1.6

657.5 10.4

2.5

666.4 19.9

1.6

662.7 14.2

0

22

988 104 2 115 9 123 9 122 4 110 4 114

E030°24'430'' N37°22'132'', E030°23'441'' N37°22'527'', E030°20'878'' N37°21'242'', E030°14'817'' N37°23'290'', E030°14'920'' N37°26'205'', E030°05'593'' N37°27'908'', E030°05'668'' N37°29'728'', E030°08'682'' N37°28'535'', E030°10'296'' N37°28'261'', E030°10'481'' N37°29'210'', E030°09'476'' N37°29'505'', E030°06'228'' N37°28'640'', E030°04'570'' N37°27'177'', E030°03'511'' N37°23'500'', E030°03'182'' N37°21'135'', E030°03'711'' N37°20'154'', E030°04'103'' N37°20'443'', E030°04'510'' N37°21'289'', E030°02'458'' N37°18'747'',

31.08.2012 31.08.2012 31.08.2012 31.08.2012

Hs; Po Hi; (Isp; Pysp) Hi; (Isp; Tsp)

31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012

(Cps; Hsp; Isp) Pk; (Isp; Psp) Cn; Hi; Ib; Po Li; (Cps; Im) Hi; Ib; (Csp; Psp) (Csp) Ib; Po; (Csp; Hsp) Ds; (Ig) Hi Hi; Pa Ib; Pzi; Po; (Csp; Cps; Hsp; Lis) Li; (Isp)

Ig; Isb; Li; Pv; (Cps) 31.08.2012 Hi; Ib; (Po)

56

1

57

6

58

4

59

2

60

2

61

6

62

6

63

3

64

6

65

3

66

6

67

6

68

6

69

5

70

5

71 72

3 4

73

5

74

5

3 7.6 3 7.8 5 8.7 9 8.9 6 9.0 2 8.2 6 7.8 9 8.6 1 8.0 2 8.7 3 7.6 1 8.2 9 7.9 4 8.4 1 8.2 5 8.5 8.0 1 8.4 2 8.3

5 11. 8 22. 3 28. 9

7.26

67.2 335.1

447.8

0.22

8.25

94.4 992

518.7

0.25

5.34

69

989

0.48

12.0 5 10.7 9 10.5 9

154. 628 1 135. 395.2 2 131. 500 2

594

0.29

28

29.8

396.2

0.17

26

34.5

7.26

84

1064

471.2

0.22

329.5

343.2

0.17

7.04

83.6 358.3

373.4

0.18

5.6

67.6 469.7

470.1

0.23

7.52

84.6 580

621

0.3

5.28

51

981

0.49

77.5

6.21

65.3 459.1

532

0.26

8.01

93.9 488.2

568.9

0.28

9.93

88.4 391

543.9

0.26

9.5

88.5 269.1

356.9

0.17

7.17

79.7 351.1

383.6

0.18

6.57

63.1 438.7

563

0.27

7.71

77.7 540

653

0.32

8.16

81.4 421.4

518

0.25

27. 6 23. 1 22. 9 24. 9 21. 5

34.5 33.3 32.3

31.4 29.5 30.5 30.5

2 0.291 2 0.336 7 0.643 5 0.383 5 0.247 8 0.307 5 0.256 7 0.241 8 0.304 8

640.3 12.6 644.9 15.5 669.6 25.8 670.1 26 670

14.8

668.8 13.2 665.2 13 659.1 14.4 665.4 14.4

28.3 0.403 660.2 23

14

14.7 0.637 660

17. 6 17. 6 10. 3 12. 1 20. 5 13. 5 15. 9 15.

0.346 16.8 657.2 43.7 4 0.369 17.4 656.3 38.2 8 0.353 20.9 646.1 32.6 6 23

43.7

0.232 651.3 34.1

0.218 25.6 662.3 28.9 9 24.6 0.366 661.4 36.5 0.422 25.9 659.2 32.2 5 25.7 0.338 652.7 30.3

23

5 142 2 8 137 0 5 105 1.9 9 104 6 2 104 6 2 105 11 8 110 2.6 1 119 5.7 1 111 12.7 0 118 3.8 3 118 2 7 123 1.6 6 123 0 5 139 0 1 134 0 0 120 1.8 1 120 0 4 122 0 9 4 131

E029°59'996'' N37°16'436'', E029°59'061'' N37°17'829'', E030°00'243'' N37°20'620'', E029°57'475'' N37°21'607'', E029°59'053'' N37°23'158'', E029°59'084'' N37°24'593'', E029°59'093'' N37°24'720'', E029°56'417'' N37°26'076'', E029°54'596'' N37°23'091'', E029°54'010'' N37°23'920'', E029°50'076'' N37°23'921'', E029°50'077'' N37°21'217'', E029°45'737'' N37°21'347'', E029°45'673'' N37°11'685'', E029°57'708'' N37°13'363'', E029°56'692'' N37°13'363'', E029°56'691'' N37°14'513'', E029°52'186'' N37°14'095'', E029°50'124'' N37°12'345'',

31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 31.08.2012 01.09.2012 01.09.2012

Co; Ib; (Hsp; Pzi; Psp; Pysp) Cn; Hi; Ib (Pzi) (Cps; Hsp; Isp; Lis) Co; Im; Li; (Hsp; Pos) Hs; Ib; (Hts; Im) (Hsp) (Cps; Isp; Isbp; Lis; Psp) Hi Li; (Cps; Pzi; Psp) Hc; Hi; Pzi Hi

01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012

(Hsp; Isp; Pysp) Cn; (Esp; Isp; Pysp) (Isp; Lis) Hi; Po; (Esp; Isp) Hi; (Isp; Pysp)

75

6

76

6

77

2

78

5

79

5

80

1

81

4

82

6

83

6

84

3

85

6

86

6

87

3

88

6

89

6

90

2

91

6

92

5

93

6

9 8.1 6 7.7 7 7.4 4 8.4 9 8.0 2 8.2 3 8.2 9 7.8 7 8.2 3 8.3 8.3 1 7.7 5 8.4 7 7.6 5 7.7 9 8.0 1 7.6 7 7.6 5 7.5

7.08

67.3 261.6

339

0.16

8.65

84.2 293.4

369.9

0.18

2.27

22.4 490.3

617.9

0.3

7.34

78.1 266.1

304.9

0.15

7.33

73.4 625

510

0.31

6.36

67.4 272.1

312.8

0.15

8.4

89.2 365.3

407.3

0.2

8.58

86.3 449

546.4

0.27

7.52

70.7 223.6

290.6

0.14

5.91

67.4 311.3

330.8

0.16

7.75

84.6 327

364

0.17

6.37

68.2 588

678

0.33

5.87

68.7 394.7

407.8

0.2

5.12

54.9 870

982

0.49

6.27

71.9 831

882

0.43

5.65

72.6 750

724

0.35

8.41

85.6 535

643

0.31

9.3

86.1 401.9

535.9

0.26

3.1

30.9 700

870

0.43

3 13. 1 14. 2 14. 2 18. 4 15. 4 18. 2 19. 6 15. 3 12. 8 21. 8 19. 7 18. 1 23. 3 19. 1 22 25. 8 16. 4 11. 9 14.

27 26.7 25.8 26.4 29.1 28.7 30.1 30.4 31.5 31.4 32.6

0.220 3 0.240 5 0.401 7 0.197 6 0.409 5 0.202 8 0.264 6 0.354 9 0.191 1 0.215 2 0.236 6

648.2 27.9

0

655.6 26.5

5

660.6 27.5

3

648

12

20

649.7 30.7

1.5

646.1 22.7

7.4

650.8 22.9

0

654

3

25

641.9 15.5

3.5

638

2.3

14.8

649.8 14.8

4

33.6 0.442 654.6 16.9

0

0.265 34.1 667 2

6

16.8

3 136 7 127 7 121 6 138 0 135 6 139 8 134 1 128 9 146 4 151 5 135 7 130 0 114 3 112 1 111 9

32.6 0.637 667.4 18.9

1.9

33.2 0.572 666.4 15.3

9.5

36.6 0.468 679.7 15.2

0

955

37.2 0.416 678.1 14.4

0

973

0

960

0

965

0.348 36.6 667.5 24 4 35.8 0.565 678.7 19

24

E029°48'594'' N37°10'954'', E029°48'759'' N37°10'496'', E029°46'906'' N37°11'069'', E029°45'365'' N37°08'460'', E029°45'521'' N37°08'438'', E029°46'046'' N37°07'890'', E029°46'191'' N37°06'883'', E029°46'313'' N37°05'643'', E029°46'500'' N37°00'046'', E029°49'821'' N37°59'365'', E029°51'134'' N37°01'750'', E029°46'891'' N37°01'454'', E029°46'683'' N37°04'445'', E029°44'149'' N37°04'485'', E029°42'489'' N37°05'647'', E029°39'705'' N37°06'497'', E029°36'491'' N37°06'422'', E029°36'713'' N37°05'412'', E029°35'351'' N37°06'691'',

01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012

Hi (Csp; Hsp) Co; Pos; Po; (Hsp) (Isp) Cn; Ib; Pv; Po; (Psp) Cn; Hi; Po Hs; Par (Pysp) Hi Li; (Cps; Esp; Psp) Hi; (Hts; Isp) (Isp; Psp; Pysp) (Hsp; Isp; Lis) (Hsp; Isp) Hi; Ib; Po; (Hts) (Cps; Isp; Psp) Hi; Ib; (Cn; Hts; Pzi; Psp)

Cn; Ib; (Psp) 01.09.2012 Hi; (Csp; Hts; Isp)

94

6

95

6

96

3

97

1

98

1

99

6

100

6

101

6

102

6

103

6

104

6

105

6

106

6

107

2

108

5

109

2

110

6

111

6

112

1

6 7.6 3 7.7 7 8.2 2 7.5 7 7.4 1 7.4 9 7.9 3 7.4 9 7.5 3 7.5 9 7.5 5 8.5 9 7.7 8 9.1 8.7 8 9.1 3 8.0 8 8.4 4 8.0

2.54

26.7 866

998

0.5

5.94

66.7 568

611

0.3

5.76

71.1 438.9

432.7

0.21

7.3

68

470.9

628.2

0.31

7.16

66.1 446.1

599.3

0.29

5.56

61.8 672

729

0.36

6.08

73.1 784

793

0.39

7.6

90.1 801

803

0.39

6.95

80.3 628

655

0.32

4.09

45

1185

0.59

2.31

23.5 852

1021

0.51

8.42

92

6.21

1072

606

681

0.33

63.8 463.2

550.9

0.27

7.34

82

2735

1.42

8.98

85.8 568

729

0.36

6.94

79.3 2411

2617

1.35

2478

8 18. 1 21. 3 25. 8 11. 8 11. 7 20. 9 24. 6 24. 5 22. 9 20. 1 16. 4 19. 3 16. 7 20. 1 13. 4

5 36.2 0.65 0.396 35.7 5 0.281 31 5 0.410 34.1 1 0.389 30.8 3

676.3 16.2

4

654.9 29.1

0

670.4 21.1

6

668.4 17.2

4.2

668.1 16.6

2

30.6 0.468 667.6 17.7

1.7

0.513 31.5 663.5 15.9 5

2.3

28.8 0.52

2.4

659.5 17

0.422 28.7 655.8 16.5 5

2.2

30

0

0.767 673.3 17.7

29.6 0.663 676.6 18.5 25.5 0.442 662.5 21.3 15.9

0.358 658.9 29.9 1

18.2 1774

668.7 48.7

2.2 9.4 0 1.7

21.4 0.481 663.1 39.5

4

21

22.9 1729

0

23.8 0.884 666.8 41.8

1.3

668.7 49.4

5.21

67.8 1387

1360

0.68

25. 8

7.34

85.9 321.7

334.2

0.16

23

24.7 0.217 664.4 28.6

2.8

5.67

55.8 774

958

0.48

15

25.9 0.624 669

2.8

32.4

25

990 126 6 107 3 107 9 107 9 110 6 116 1 120 9 125 9 103 8 995 116 4 120 8 111 6 120 4 113 4 115 3 118 0 111

E029°33'537'' N37°05'560'', E029°31'836'' N37°01'434'', E029°32'172'' N37°59'959'', E029°27'499'' N36°58'770'', E029°28'495'' N36°59'158'', E029°29'133'' N36°59'334'', E029°29'031'' N37°00'084'', E029°30'448'' N37°00'659'', E029°31'922'' N37°00'939'', E029°32'130'' N37°04'203'', E029°32'179'' N37°05'805'', E029°31'858'' N37°09'657'', E029°29'727'' N37°09'655'', E029°29'726'' N37°31'175'', E029°43'107'' N37°32'774'', E029°35'720'' N37°33'781'', E029°38'509'' N37°35'255'', E029°40'590'' N37°33'909'', E029°45'243'' N37°37'823'',

01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012 01.09.2012

Hi; Po; (Ig) Ib; (Hsp) (Isp) (Csp; Hsp; Isp; Pysp) (Pysp) Hi; Ib; Po Hs; (Isp) Hi; Ps (Esp) Hi; Po; (Isp) Hi; Po; (Hsp)

01.09.2012 02.09.2012 02.09.2012 02.09.2012 02.09.2012 02.09.2012 02.09.2012

Po; (Isp) Po Po; (Csp; Isp; Lis) Hs; (Pysp)

Hi 02.09.2012 Cn; Ib; Po

113

6

114

4

115

6

116

5

117

2

118

6

119

2

120

2

121

5 Max Min

3 8.6 4 9.2 6 7.8 5 8.1 4 8.9 5 7.9 1 9.4 4 9.0 5 8.2 7 9.4 4 7.0 5

6.59

62.7 720

903

0.45

6.18

67

1361

0.68

5.64

57.7 739

880

0.43

7.3

74.2 896

1080

0.54

5.61

38905 380.7 81.1 6 4

23.3 3

7.82

88.5 762

0.4

6.55

90

1216

816

14.2 24265 23430 2

14. 5 19. 4 16. 6 16. 1 28. 2 21. 6 26. 8 14. 2

8.13

79.2 435.7

557

0.26

7.94

89.1 486.5

526.9

0.25

21

12.0 5

154. 38905 23.3 23430 1 6 3

2.27

22.4 77.5

28. 9 10. 3

228.8

0.02

9 100 9 106 4 115 6 106 0

26.5 0.585 678.6 34.4

0

26.7 0.884 674.4 26.9

2.7

27.8 0.572 667.5 27.5

5.5

27.8 0.702 676.2 31.7

1.5

23.03 29.2 691.3 28.1 7

3.7

848

31.1 0.533 687.4 21.1

3

890

15.28 29.4 684.9 26.5 8

2.7

917

29

0.352 663.2 39

27.1

0.342 669 5

37.2 1774 12.4

21.3

769.9 54.2

0.148 635 9

10.4

118 0 3 112 16.3 0 153 17.5 8 0

E029°45'988'' N37°39'861'', E029°44'858'' N37°40'544'', E029°51'051'' N37°43'174'', E029°58'959'' N37°42'829'', E030°00'849'' N37°41'883'', E030°04'609'' N37°39'077'', E030°02'855'' N37°35'143'', E029°58'931'' N37°39'565'', E030°22'474'' N37°45'645'', E030°23'855''

02.09.2012 02.09.2012 02.09.2012 02.09.2012 02.09.2012 02.09.2012 02.09.2012

(Psp) Hi; Im (Esp; Isp; Pysp) Ib; Po; (Hsp; Psp) (Hts; Hsp; Lis; Pas) Hi; Ib; (Hts) Li; (Isp)

02.09.2012 02.09.2012

Ib; (Cn; Hsp; Pysp)

282

Abbreviations: St. no, site number; St. Ty., site type; DO, dissolved oxygen, mg L-1; % DO, percent saturation; EC, electrical conductivity, µS cm-1; Sp. EC, specific electrical conductivity; Sal, salinity, ppt; Tw, water temperature, ºC; Ta, air temperature, ºC; TDS, total dissolved solid, mg L-1; Atm., atmospheric pressure, mmHg; Moist., Moisture, %; W. s., wind speed, km h-1; Elev., Elevation, m a.s.l; Cn, C. neglecta; Csp, *Candona sp.; Co, Cypria ophtalmica; Cps, *Cypria sp.; Ds, Darwinula stevensoni; Esp, *Eucypris sp.; Hc, Herpetocypris chevreuxi; Hin, *H. intermedia; Hts, *Herpetocypris sp.; Hi, Heterocypris incongruens; Hs, H. salina; Hsp, *Heterocypris sp.; Ib, Ilyocypris bradyi; Ig, *I. gibba; Im, I. monstrifica; Isp, *Ilyocypris sp.; Isb, *Isocypris beauchampi; Isbp, *Isocypris sp.; Li, Limnocythere inopinata; Lis, *Limnocythere sp.; 26

Pas, *Paralimnocythere sp.; Pk, *Physocypria kraepelini; Par, Potamocypris arcuata; Pf, *P. fallax; Ps, *P. similis; Pv, *P. variegata; Pos, *Potamocypris sp.; Pzi, Prionocypris zenkeri; Pa, *Pseudocandona albicans; Psp, *Pseudocandona sp.; Pfo, *Psychrodromus fontinalis; Po, P. olivaceus; Pysp, *Psychrodromus sp.; Tc, *Trajancypris clavata; Tsp, *Trajancypris sp. The sub-recent form of taxa were shown in parenthesis. Aquatic types; 1, spring; 2, lake; 3, dam; 4, pond (or pool); 5, creek; 6, trough. * represents new reports for Burdur.

27

Appendix B Minimum and maximum values of eight different ecological variables for 22 species recorded in Burdur. Species Darwinula stevensoni Candona neglecta Pseudocandona albicans Cypria ophtalmica Physocypria kraepelini

Tw 41-352 2.1316-28.917 2.925-29.212 1.134-332 0.937-31.438

EC 863-96003 4.512-529018 92.926-506327 40.82-52602 64.2939-79937

pH 5.54-9.675 619-11.812 6.426-925 4.714-132 6.5940-10.4437

DO 0.326-16.4787 0.322-15.42 0.7528-15.829 02-202 1.0730-19.1237

Ta 128-30.29 6.620-42.321 9.330-36.321 635-34.563 4.1030-3438

Elev. 510-14402 017-31942 6131-229032 036-250014 0.52-16632

TDS 4.611-34212 0.052022-169123 0.153463-580.533 0.247863-450.533 0.215263-70241

Sal. 113-1514(0-3915g/L) 014-4024 0.118-5.515 04-2514 039-2.414

Ilyocypris gibba

3.727-422

122.217-1381027

5.84-9.82

2.0342-142

11.230-42.3542

12-31992

0.09943-575.333

04-3.643

Ilyocypris monstrifica

10.544-3544

30018-526018

6.844-9.2663

4.0763-10.7963

1535-34.563

745-139818

0.215263-0.300328

0.118-3.3018

Ilyocypris bradyi Prionocypris zenkeri Trajancypris clavata

1.6834-33.821 7.1430-31.722 3.727-30.647

15.642-529018 15.9422-139322 187.433-352927

5.432-9.8922 6.2933-9.362 7.252-8.822

0.282-20.732 2.012-20.702 1.8421-19.2321

8.420-42.821 2.230-33.322 16.463-38.721

045-31942 1045-298046 1045-242641

0.148228-177641 0.214522-61512 0.252263-12241

04-4.543 032-0.8322 0.133-1.347

Herpetocypris chevreuxi

5.748-33.949

74.728-147549

6.350-9.334

2.1151-175

1035-42.821

045-192123

0.135221-958.7549

0.0428-414

Herpetocypris intermedia

552-28.321

221.222-332052

633-9.1822

1.410-12.110

12.463-4221

12517-142010

0.147522-49433

0.133-0.433

Psychrodromus olivaceus Psychrodromus fontinalis

1.682-36.728 7.454-29.823

033-247863 84.917-386622

553-11.412 619-9.9217

1.742-202 2.5523-17.149

1828-40.521 15.922-31.122

0.52-170014 28518-223532

033-177463 0.254263-565.523

033-2.112 017-0.3763

Heterocypris incongruens

3.727-33.921

11.0422-1005018

0.2828-2012

12.463-42.921

017-31942(457055)

0.131322-2003441

07-5018

27

22

18

5.333-12.82

Heterocypris salina Isocypris beauchampi Potamocypris fallax Potamocypris similis

14

28

41

4

22

28

22

45

41

55

22

41

3.7 -34 1527-25.918 10.641-26.923 12.928-3321

15.94 -10050 254.19-123456 178.421-107412 033-386622

6.05 -9.9 7.3418-8.4563 5.0423-9.7317 6.533-8.8417

0 -1884 5.0523-10.857 1.4721-10.3058 3.2723-13.5428

18.5 -39.1 179-3363 24.223-42.321 19.6049-42.921

0 -2079 (4570 ) 30056-156023 60517-195423 11428-178923

0.1748 -20034 0.320463-186.039 0.133921-177641 033-414.723

0.122-5018 0.118-0.2463 0.112-1.1023 033-0.3963

Potamocypris variegata Potamocypris arcuata

9.959-2938 1327-2822

137.422-408527 16310-127927

6.558-9.1522 6.3261-9.9561

0.960-1429 7.1122-1227

12.559-35.763 27.222-39.522

11018-168412 561-201841

0.089022-41812 0.131322-163141

0.0622-0.3618(128) 0.122-0.522

Limnocythere inopinata

4.7530-3544

28.232-2426563

6.44-10.462

2.9123-13.2639

4.1030-3438

545-237641(457055)

0.215263-2397341

04-254

Abbreviations: Tw, water temperature (°C); EC, electrical conductivity (μS cm-1); DO, dissolved oxygen (mg L-1); Ta, air temperature (°C); Elev., elevation (m a.s.l); TDS, total dissolved solids (mg L-1); Sal., salinity (‰); 1Van Doninck et al., 2003; 2Külköylüoğlu, 2013; 3Gandolfi et al., 2001; 4

Ruiz et al., 2013; 5Rossetti et al., 2004; 6Külköylüoğlu et al., 2007; 7Külköylüoğlu, 2009; 8Horne, 2007; 9Külköylüoğlu, 2005b; 10Mezquita et al., 28

1999c;

11

Mischke et al., 2012;

12

Külköylüoğlu et al., 2013;

13

Keyser, 1976;

14

Meisch, 2000;

15

Van Doninck et al., 2002;

16

Külköylüoğlu, 2005a;

17

Külköylüoğlu et al., 2012c; 18Rasouli et al., 2014; 19Mazzini et al., 2014; 20Külköylüoğlu, 2005c; 21Yavuzatmaca et al., 2015; 22Uçak et al., 2014;

23

Akdemir and Külköylüoğlu, 2014; 24Gao and Hailei, 2014; 25Scharf and Brunke, 2013; 26Iglikowska and Namiotko, 2012; 27Mezquita et al., 2001;

28

Yılmaz, 2014; 29Delorme, 1991; 30Külköylüoğlu et al., 2014; 31Mezquita et al., 1999a; 32Külköylüoğlu et al., 2012b; 33Külköylüoğlu et al., 2012a;

34

Dügel et al., 2008; 35Horne and Mezquita, 2008; 36Pieri et al., 2009; 37Kiss, 2007; 38Özuluğ, 2011; 39Yılmaz and Külköylüoğlu, 2006; 40Yu et al.,

2009; 41Van der Meeren et al., 2010; 42Narasimha Ramulu et al., 2011; 43De Deckker, 1981; 44Karan-Žnidaršič and Petrov, 2007; 45Altınsaçlı, 2004; 46

Aygen et al., 2012;

47

Külköylüoğlu, 2008;

52

57

Valls et al., 2014;

48

Külköylüoğlu and Vinyard, 2000;

Mezquita et al., 1999b;

53

Boomer et al., 2006;

54

49

Sarı, 2007;

50

Fernandes Martins et al., 2010;

Roca and Baltanás, 1993;

55

Guo et al., 2013;

56

51

Karakaş Sarı and

Escrivá et al., 2014;

Külköylüoğlu, 2003b; 58Özuluğ, 2012; 59Külköylüoğlu and Dügel, 2004; 60Creuzé des Châtelliers and Marmonier, 1993; 61Pieri et al., 2006; 62Van

der Meeren et al. 2011; 63the present study.

29

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Fig. 1. 121 ramdomly selected sampling sites from 11 counties (Merkez, Ağlasun, Çeltikçi, Bucak, Kemer, Yeşilova, Karamanlı, Tefenni, Çavdır, Gölhisar, and Altınyayla) of Burdur.

45

Fig. 2. Numbers of sites carrying 0, 1, 2, 3, 4 or more (4+) species of ostracods.

46

Fig. 3. Graph of CCA showing the ordination of 13 species and the six environmental variables (arrows) (Tw, pH, DO, EC, Elev and Ta) from Burdur by first and second axes. Triangles show species code. For abreviations see Appendix A.

47

50 Sta.Type Sta. Taxa Sp. Sex Asex

40

Value

30

20

10

0

0 9 8 7 6 85 34 83 32 64 48 33 18 03 5-8 4-7 3-5 2-4 0-1 9-1 8-1 7-1 6-1 9 3 8 3 8 73 58 43 28 8 14 13 11 10

Elevational range

Fig. 4. The number of taxa, species, site type, sampled site and species with sexual and asexual reproduction at the nine different 150 m a.s.l. elevational ranges in Burdur. Abbreviations: Sta. (number of sites), Sta. Type (site type), Sp. (species number), Sex (species sexually reproduce) and Asex (species asexually reproduce).

48

Table 1 Poisson probabilities of 0, 1, 2, 3 and 4(+) occurrence of ostracod species calculated using Equation 1 in Burdur. P(x=0) probability of no occurrence 0.29 P(x=1) probability of one occurrence 0.41 P(x=2) probability of two occurrences 0.29 P(x=3) probability of three occurrences 0.14 P(x=4+) probability of four(+) occurrences 0.05

49

Table 2 Calculated Chi-square values of 0, 1, 2, 3 and 4(+) occurrences of species in Burdur. Expected (E) probability = N x Poisson Probability; five classes (n) with two constants (habitat and species) so degrees of freedom, df = n-2 => 5-2 = 3. Class x f(obs.freq.) Exp. Prob. O-E (O-E)2 (O-E)2/E 1 0 41 35.32 5.68 32.29 0.91 2 1 35 49.74 -14.74 217.24 4.37 3 2 26 35.02 -9.02 81.44 2.33 4 3 14 16.44 -2.44 5.96 0.36 5 4 5 5.79 -0.79 0.62 0.11 (X2)

N=121

50

8.08

Table 3 Summary table of the CCA for 13 species (with two or more occurrences) from 79 sites and six environmental variables in Burdur (* shows the results of DCA). Axes 1 2 3 4 Total inertia *Lengths of gradient 0.00 5.75 3.98 2.25 Eigenvalues 0.35 0.24 0.10 0.04 6.19 Species-environment correlations 0.65 0.60 0.30 0.32 Cumulative percentage variance of species data 5.60 9.50 11.10 11.80 of species-environment relation 46.90 78.80 92.60 98.50 Sum of all eigenvalues 6.19 Sum of all canonical eigenvalues 0.74

51

Table 4 Optimum (uk) and tolerance (tk) levels of 13 species to four different ecological variables in Burdur. N2 represents Hill’s coefficient value as the measure of effective number of occurrences. Abbreviations: DO (dissolved oxygen concentration, mg L-1), EC (electrical conductivity, µS cm-1), Tw (water temperature, ºC), Max (maximum) and Min (minimum).

Species H. incongruens P. olivaceus I. bradyi L. inopinata C. neglecta H. salina C. ophtalmica P. kraepelini I. monstrifica P. variegata P. zenkeri P. fontinalis H. intermedia

Count 38 26 26 8 12 6 3 2 4 5 4 2 2

Max 171 121 143 14 41 121 19 2 18 135 25 9 74

N2 11.61 11.12 7.60 5.30 4.53 4.25 2.07 1.80 1.78 1.75 1.57 1.42 1.05 Mean Max Min

pH uk tk 7.95 0.29 7.89 0.44 7.80 0.27 8.50 0.44 7.76 0.31 7.94 0.43 7.56 0.37 8.31 0.16 8.85 0.51 8.12 0.23 7.92 0.37 7.25 0.37 8.10 0.36 8.00 0.35 8.85 0.51 7.25 0.16

DO uk tk 7.83 2.39 6.68 2.19 7.35 1.69 5.88 2.16 7.54 1.97 8.39 2.01 4.58 3.70 7.12 4.42 9.19 3.91 11.21 1.01 8.25 2.48 5.38 3.22 7.08 0.83 7.42 2.46 11.21 4.42 4.58 0.83

52

EC uk 610.90 771.07 632.91 2619.36 633.33 741.77 541.34 368.57 421.50 491.36 651.93 686.35 701.61 759.38 2619.36 368.57

tk 120.46 202.11 120.67 4492.43 175.75 203.18 56.85 34.51 127.03 6.04 96.09 91.44 5.57 440.93 4492.43 5.57

Tw uk tk 19.25 3.98 17.21 3.53 17.91 4.32 24.45 2.13 14.69 3.18 22.01 6.01 13.56 3.42 25.57 3.25 25.57 3.54 24.75 1.42 18.89 4.41 12.79 1.20 23.03 1.77 19.97 3.24 25.57 6.01 12.79 1.20

Table 5 Dominance percentage (%) of 22 ostracod species among 3516 individual, their occurrence frequencies in six different aquatic habitat, and Shannon-Wiener index (H′) value of each habitat. Abbreviations: ASI (All Sample Index) and JSE (Jackknife Standard Error). Species

%

Spring (n=10)

C. neglecta

3.01

4

C. ophtalmica

0.97

1

D. stevensoni

0.03

1

H. chevreuxi

0.77

H. intermedia

2.16

H. incongruens

30.55

H. salina

10.41

I. bradyi

18.15

I. gibba

0.11

I. monstrifica

0.71

I. beauchampi

0.06

L. inopinata

1.17

P. kraepelini

0.09

P. arcuata

0.20

1

P. fallax

0.14

1

P. similis

0.06

P. variegata

5.23

P. zenkeri

0.91

P. albicans

0.28

P. fontinalis

0.31

1

P. olivaceus

24.49

2

T. clavata

0.20

Shannon-Wiener

Lake (n=9)

Dam Pond Creek Trough (n=10) (n=17) (n=17) (n=58) 2

5

1

2 1 2

1

4

2

1 2

3

31 5

7

14

1 1

1

2 1

2

4

2

1

1

1 1

1

3

2

2

1 1 3

1

6

14

1

H

1.79

1.32

1.42

2.25

1.71

1.68

Variance H

0.05

0.04

0.09

0.03

0.02

0.01

Exp. H

6.00

3.75

4.14

9.44

5.54

5.35

53

ASI

JSE

2.33

0.25

Table 6 Number of species (or species richness, N. spp.), individuals (N. ind.), individual per species (ind./spp.) and species per site (spp./site) with minimum and maximum values of pH, dissolved oxygen (DO, mg L-1), electrical conductivity (EC, S cm-1), salinity (Sal, ppt), water (Tw) and air (Ta, C) temperatures and elevation (Elev., m a.s.l.) in six different habitat types.

Habitats N. spp.

Pond Trough Spring Creek Lake Dam

15 10 7 7 4 3

N. ind. 156 2582 382 322 50 24

ind./spp. 10.40 258.20 54.57 46.00 12.50 8.00

spp./site

pH

DO

EC

Sal

Tw

Ta

Elev.

0.88

7.82-9.26 7.05-8.64 7.15-8.23 7.13-8.78 7.44-9.44 8.13-8.76

2.65-11.29 2.31-11.57 4.55-9.30 3.20-9.93 2.27-12.05 5.04-7.52

198.10-1824 77.50-1387 272.10-978 266.10-896 395.20-389056 311.30-580

0.11-1.12 0.02-0.68 0.15-0.59 0.15-0.54 0.17-23.33 0.15-0.30

12.90-28.90 11.00-27.60 11.70-27.20 10.30-27.80 14.20-28.20 20.10-27.10

16.40-33.00 12.40-37.20 23.60-34.50 17.40-36.60 18.20-36.60 25.60-34.40

785-1341 348-1538 637-1428 815-1391 848-1216 282-1515

0.17 0.70 0.41 0.44 0.30

54