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J Insect Conserv (2008) 12:539548 DOI 10.

1007/s10841-007-9094-5

ORIGINAL PAPER

A preliminary study of the Odonata of SAlbufera Natural Park, Mallorca: status, conservation priorities and bio-indicator potential
Mayumi Sato Nick Riddiford

Received: 22 December 2006 / Accepted: 24 May 2007 / Published online: 12 July 2007 Springer Science+Business Media B.V. 2007

Abstract This study obtained baseline information for adult Odonata and assessed their conservation priorities and suitability as biological indicators in SAlbufera Natural Park in Mallorca, Spain. At this site, human activities in and around the wetland have raised concerns about their impact on the ecosystem. Investigations on adult diversity produced records of 14 species (four Zygoptera and 10 Anisoptera) and included the rst record of Erythromma viridulum for the park. Detrended Correspondence Analysis (DCA) ordination categorised study sites according to their geographical locations in the park and showed clustering of the sites around particular species based on these locations. This pattern might reect the differences in brackishness in water supplied by different water sources. Canonical Correspondence Analysis (CCA) indicated that some environmental factors were related to particular species. Water ow, vegetation, and depth and size of a water body could discriminate stenotopic species from eurytopic species. Only a few species appeared to be tolerant to the sites with high salinity and low oxygen concentration. The ordination results can be useful for establishing conservation priorities with information of

species diversity, abundance, distribution and ight period. Although, with the current limited basic information, the use of Odonata species as biological indicators seems to be difcult, some clear relationships between environmental factors and particular species indicate the great potential of using adult Odonata as biological indicators in the park. Keywords Odonata Species assemblages Environmental factors Indicator species

Introduction Many ecosystems have been subjected to great stress from increasing human activity, resulting in many species disappearing before any conservation actions are taken. Since organisms require certain habitats, the knowledge of their relation to their habitat is critical for establishing conservation and management plans. An understanding of this relationship can also be used to monitor ecosystem changes where the species or species assemblages respond to environmental changes in an easily monitored way. Many studies have reported that using certain species as biological indicators can be an effective way to monitor environmental quality (e.g. Landres et al. 1988; n and EkNoss 1990; Brown 1991; Kremen 1992; Sahle estubbe 2001; Maleque et al. 2006). Monitoring multispecies response provides more valuable perspective in an environment with complex sources of disturbance (Hodkinson and Jackson 2005). Consequently, identifying habitat types and quality based on species presence has potential applications both in terms of choosing and assessing the species as indicators in planning a monitoring programme and establishing conservation priorities for the species.

M. Sato Department of Biology, The University of York, York YO10 5YW, UK Present Address: M. Sato (&) Center for Ecological Research, Kyoto University, Otsu 520-2113, Japan e-mail: m-sato@ecology.kyoto-u.ac.jp N. Riddiford The Albufera International Biodiversity Group, Schoolton, Fair Isle, Shetland ZE2 9JU, UK

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Recently, the potential of using Odonata as convenient and sensitive biological indicators has been increasing (e.g. Carle 1979; Watson et al. 1982; Osada and Tabata 1992; Wildermuth 1994). Several studies have attempted to examine the relationship between habitat preferences and nkova 1997; species or species assemblages (e.g. Bula Clark and Samways 1996; Osborn and Samways 1996; Stewart and Samways 1998; Steytler and Samways 1995; n and Ekestubbe 2001; Osborn 2005; Suhling et al. Sahle 2006). These previous studies indicated the great indicator potential of this taxonomic group. However, species or species assemblages and habitat characteristics vary spatially and temporarily. Therefore, the knowledge of the relationship between them at one place would require further work for its practical application in a monitoring programme at a different place. SAlbufera Natural Park in Mallorca, Spain, is the largest wetland in the Balearic Islands and one of a declining number of major wetlands remaining in the Mediterranean. In recognition of its international importance as a wetland, it was added to the RAMSAR Convention by the Spanish government in 1989. This 1,500 ha freshwater coastal wetland supports a wide range of species (Riddiford 1991, 2003). However, expanding human activities such as intensive agriculture, tourism and a coalred power station situated on the perimeter of the park have resulted in various concerns for maintaining this biodiversity. Specically, the direct usage of water in and around the park and discharge of seawater from the power station to the wetland have caused concern owing to their impact on the ecosystem (Riddiford 1991; Veraart 1999). Thus, monitoring biotic responses to environmental changes is crucial. Various monitoring programmes are currently being done, using plants, birds and insects. Studies on Odonata have been conducted in the park since 1989. A summary paper reported a total of 15 species, comprising three Zygoptera and 12 Anisoptera, conrming the wetland as a key site for the biodiversity of Odonata in the Balearics (Riddiford and Mayol 1997). Although previous studies indicated the potential use of Odonata as a key indicator group for the park (Chapman 1996; Riddiford and Mayol 1997), habitat preference has been reported only for some species, and the relationship between environmental factors and particular species has never been investigated. Also, little is known of the diversity, distribution, abundance and ight period in June and July since no survey had been conducted during that period. Answering these questions will improve our understanding of both their conservation value and their potential as biological indicators to monitor environmental changes. This study had four objectives; rst, to provide baseline information for the summer months on diversity, distribu-

tion, abundance and ight period of each adult Odonata species in sAlbufera; second, to investigate the relationship between species and their habitat types by specically examining major environmental factors that determine Odonata species assemblages; third, to develop wider conservation priorities for Odonata species in the park; and nally, to assess the suitability of adult Odonata as biological indicators of environmental quality in the park.

Study area and methods Study area The eld site was in SAlbufera Natural Park in Mallorca, (3947 N, 36E) Spain (Fig. 1). Four terrestrial transects (Fig. 1) were chosen to investigate diversity of the Odonata species in the park and the ight period of each species, where ight period describes the period when Odonata occur as adults including tenerals. Eleven water bodies were chosen as study sites (Fig. 1) to investigate the abundance and species richness of adult Odonata and their association with particular habitat types. Only sexually mature adults at water bodies were used. Site selection and the site numbers were based on previous sampling sites for adult and larval Odonata species reported by Chapman (1996) and the parks regular water quality sampling sites, with a consideration for accessibility. However, sites 30 and 29 were newly added to ensure the study sites covered as many different habitats as possible. Odonata sampling Data were collected between 12th June and 5th August 2002. Data collection was conducted on clear, windless days between about 10:00 h and 16:00 h when adult

Fig. 1 Map of the study area. , SAlbufera Natural Park boundary; with numbers indicate 11 study sites. Study sites are categorised based on geographical locations in the park; , north-east area; , middle area; , south-west area. with letters (AD) indicate four transects

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Odonata are most active (Watson et al. 1982). Adult individuals were counted when observed within a 5-m band on either side of the transects. The counts were conducted twice a week for each of four transects. The number of individuals for each species was also noted at 11 study sites, each of which consisted of a 100 m2 quadrat. In order to avoid the recording of accidental iers (Schmidt 1985), the number of individuals was determined during the rst 15 min. However, ovipositing females were continuously counted during the next 15 min in order to increase the chance of observing their behaviour. Therefore, each sampling occasion consisted of a 30-min total observation period. Twenty replicates of data were gathered for each study site. Askew (1988) and Riddiford and Mayol (1997) were used to identify species. Nomenclature follows Askew (1988). Habitat characterization

Small, secondary canals: less than 5 m in width or lagoons less than 80 m2

The mean depth of the quadrat was based on ve measurements taken between 50 cm and 200 cm from the edge of the water. The state of water movement was allocated to one of two categories, either still or owing water, and was conrmed by throwing a reed stem into the water 10 times at each study site. In each quadrat, plant species were dened as submerged or emergent, depending on their habit, and the percentage cover of each was estimated. Dissolved oxygen and salinity were measured and averaged as the indicator of water quality. Two measurements were taken, in June and August, and additional data were added from the water quality sampling in June and August of the previous year. Data analysis

Table 1 summarises seven habitat variables included in the Canonical Correspondence Analysis (CCA). The variables were measured for each of 11 study sites. Two types of water body were present: canals, comprising man-made channels connected directly or indirectly to the sea; and lagoons, comprising any open body of water not a canal. Bodies of water were categorised based on size. Three sizes were chosen based on the following criteria: Large, main canals: 15 m or more in width or lagoons 150 m2 and above in area Medium, secondary canals: from 5 m but less than 15 m in width or lagoons from 80 m2 but less than 150 m2 in area

Table 1 Environmental variables included in the CCA analysis Unit of measurement 1 Water body Nominal variables 1 2 3 2 Water ow 3 Water depth 4 Emergent vegetation 5 Submerged vegetaion 6 Oxygen concentration 7 Salinity 1 2 Description

Small sized canals or lagoons Medium sized canals or lagoons Large sized canals or lagoons Still water Flowing water Centimeters % cover (arcsin transformation) % cover (arcsin transformation) mg/l g/l

Detrended Correspondence Analysis (DCA) and CCA were milconducted using CANOCO program (ter Braak and S auer 2002). DCA ordinates sites along axes, arranging sites most similar in taxonomic composition closest together (Gauch 1982). The advantage of using DCA is that it is an indirect gradient analysis technique, requiring only the distribution value of species abundance (Palmer 1993). Therefore, it is a good preliminary test to examine the ability of species distribution data to indicate a particular environmental pattern (Kremen 1992). By using DCA technique, we tested whether sites, categorised based on geographical locations in the park (see Fig. 1), can show any patterns in species composition without studying environmental gradients directly. The water in the northern part of the park is known for its relatively high brackishness because, for a number of years, there was leakage of salt water from the power station supply pipe (Riddiford 1991; Chapman 1996), and sites 30, 19 and 20 were located in this area. Due to its geographic proximity to the shoreline, the eastern part is also known for its relatively high brackish water (Riddiford 1991), and site 3 was located in this sector. Therefore, sites 30, 19, 20 and 3 were allocated to the same, north-east area category. In contrast, water in southern and western parts of the park is fed by an unknown number of underground springs near site 10 and is less brackish (Riddiford 1991), and sites 10 and 12 were allocated to the south-west area category. Sites located in the middle of the park, namely sites 25, 26, 27, 29 and 7, were grouped together into the middle area. In order to further examine the relation of each site to any patterns in species composition, 11 individual sites were also ordinated using DCA.

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Canonical Correspondence Analysis was used to analyse the relationship between species distributions and environmental variables. CCA, a direct gradient analysis technique, ordinates the patterns of variation in the species composition which could be best explained by the observed environmental gradients that are not expressed in DCA ordination (ter Braak and Prentice 1988; Palmer 1993). The statistical signicance of the relation between the species and the whole set of environmental variables was tested by a Monte Carlo permutation test, using an F-ratio of the sum of all eigenvalues as the test statistic (ter Braak and Prentice 1988). Species data for DCA and CCA were log (y + 1) transformed for performing CANOCO. In order to facilitate comparison with species data, environmental variables expressed in percentage were arcsin transformed (Sokal and Rohlf 1994). The acceptable level of signicance (5%) was established for all statistical procedures.

and ten species of Anisoptera, were recorded in this study, including the rst record of E. viridulum in the park. All 14 species were observed during the line transects. However, Ceriagrion tenellum, Aeshna mixta, Selysiothemis nigra, Sympetrum fonscolombii, Sympetrum striolatum were not recorded at any of the study sites. Ischnura elegans, Crocothemis erythraea and Orthetrum cancellatum were at all study sites and line transects. Among the study sites, Orthetrum coerulescens and Aeshna isosceles were found only at site 10, and E. viridulum was found only at site 12. Oviposition Observations of ovipositing females are summarised in Table 2. Ovipositing females were found for all species present at sites 19, 26 and 30. Only certain species were found ovipositing at sites 3, 10, 12, 25, 27 and 29. No ovipositing females were at site 7. Flight period Two species showed a declining trend and disappeared in the middle of the study season. Cercion lindeni started declining after week 2 and was not recorded after week 6 (Fig. 2a). A. isosceles was not found after week 4 (Fig. 2c).

Results Distribution Distributions of adult Odonata are summarised in Table 2. A total of 14 species of Odonata, four species of Zygoptera

Table 2 Distribution and abundance of adult Odonata from 11 study sites and four transects 3 Zygoptera Coenagrionidae lys), (C. lin) Cercion lindeni (Se Ceriagrion tenellum (Villers), (C. ten) Ischnura elegans (Vander Linden), (I. ele) Erythroma viridulum (Charpentier), (E. vir) Anisoptera Aeshnidae Aeshna mixta Latreille, (A. mix) ller), (A. iso) Aeshna isosceles (Mu Anax imperator Leach, (A. imp) lys), (A. par) Anax parthenope (Se Libellulidae ), (C. ery) Crocothemis erythraea (Brulle Orthetrum cancellatum (Linnaeus), (O. can) Orthetrum coerulescens (Fabricius), (O. coe) Selysiothemis nigra (Vander Linden), (S. nig) lys), (S. fon) Sympetrum fonscolombei (Se Sympetrum striolatum (Charpentier), (S. str) Total number of individuals Total number of species 63 4 71 3 66 5 154 6 114 5 29 1 63 5 145 5 113 5 131 5 141 5 7 7 6 12 x x 1 54* 2 9 6 1 23* 29* 2 9* 20* 11 3 19* 31* 7 2 47* 16 4* 23* x x x x x x x x x x x x 4* 1 9* 20* 16* 12* 4 7 12* x x x x x x x x x x x 7 60 32* 17* 87* 30* 29* 23* 41* 65* 39* 51* x 10 35* 10* 42* 35* 22* 51* x x x x x x 7 10 12 19 20 25 26 27 29 30 A B C D

Numbers at the top indicate study sites. Letters (AD) indicate the transects. * indicates presence of ovipositing females. Abbreviations used in the gures are listed in brackets

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(a)
Th e number of Coenagrionidae spp.
25 20 15 10 5 0 1 2 3 4 5 Tim e (w eek) 6 7 C. l i n C. ten E. vir

Abundance and species richness at study sites The total number of individuals, total species richness and total individual number of each species at each study site are summarised in Table 2. Among 11 study sites, the maximum number of individuals and species richness was recorded at site 12; however, E. viridulum constituted the largest number of individuals, consisting of more than half of the proportion of the total individuals at this site. Sites 3, 7 and 20 showed low numbers for both individuals and species richness. I. elegans accounted for 40% of all individuals and was present at all sites. In general, C. lindeni, C. erythraea and O. cancellatum were present in relatively high proportions in total numbers. Ordination of species and sites The eigenvalue, which measured the importance of each of the DCA axes on a scale between 0 and 1, was 0.558 for the rst axis and 0.402 for the second axis. The two principal axes accounted for 36.9% of the total variance (Fig. 3a, b). The DCA ordination of species and sites usually shows that sites, where a particular species has higher frequency, tend to have their site symbols clustered around the species milauer 2002). In this study, sites clustered (ter Braak and S according to the geographical locations in the park although the clusters overlapped each other (Fig. 3a). Individual sites also clustered around particular species (Fig. 3b). Site 12 especially showed high frequency of E. viridulum, and site 10 of O. coerulescens. However, other sites were either fairly equally distributed around certain species or clustered in the middle of the species ordination. The species located at the centre of the DCA ordination are typically ubiquitous species, and the ones found on the edges of the diagram could be rare species or species that prefer extreme environmental conditions (Kremen 1992; ter Braak 1995). I. elegans was the closest to the centre of the diagram, and A. isosceles, O. coerulescens, E. viridulum and O. cancellatum might qualify as the latter category. Ordination with environmental variables The results of the Monte Carlo permutation test based on the rst and the sum of all canonical eigenvalues showed the signicance of the rst ordination axis and that of all canonical axes together (F = 34.99, P < 0.01; F = 19.478, P < 0.01, respectively), showing the relationship between the species and the environmental variables was statistically signicant. The eigenvalue was 0.403 for the rst axis and 0.368 for the second axis. The two axes accounted for 71.1% of the total variance (Fig. 4).

(b)
The number of Libellu lidae spp.

16 14 12 10 8 6 4 2 0 1 2 3 4 5 6 7 Time (w eek) C. ery O. c an O. coe S. nig S. s tr S. fon

(c)
Th e number of Aeshni dae spp.

12 10 A. mix 8 6 4 2 0 1 2 3 4 5 Tim e (w eek) 6 7 A. iso A. imp A. par

Fig. 2 Flight period and changes in numbers of Odonata species at four transects. Week 1, from June 17th to June 23rd; week 2, from June 24th to June 30th; week 3, July 1st to July 7th; week 4, July 8th to July 14th; week 5, July 15th to July 21st; week 6, July 22nd to 28th; week 7, July 29th to August 4th. (a) Coenagrionidae spp. (b) Libellulidae spp. (c) Aeshnidae spp.

On the other hand, three species, A. mixta (Fig. 2c), S. nigra and S. fonscolombii (Fig. 2b), started appearing in week 7. Some species were seen fairly consistently throughout the study season, namely E. viridulum (Fig. 2a), Anax imperator, Anax parthenope (Fig. 2c), C. erythraea and O. cancellatum (Fig. 2b). The rest of the species were seen sporadically with no clear trend.

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-1.0

(a)
A.iso A.imp C.lin DCA ax is 2

O.coe 10 VEG(EME)

North-east area Middle area South-west area

A.iso
C.ery

27 19
CCA axis 2
O.coe I. ele A.par E.vir

20

26 SALINITY C.lin A.par 30 O.can BODY 25

7 I.ele A.imp 29 DEPTH C.ery

FLOW

O.can

OXYGEN 3

12 E.vir

-1 -1 5

DCA axis 1

VEG(SUB)

(b)
A.iso

A.imp C.lin DCA axis 2

Site 3 Site 7 Site 10 Site 12 Site 19 Site 20

Site 25 Site 26

-1.0 -0.6

x Site 27
Site 29 Site 30

CCA axis 1

1.0

C.ery

Fig. 4 Canonical Correspondence Analysis ordination triplot of the species and 11 study sites with environmental variables represented by arrows. Closed circles and the corresponding numbers indicate each study site. Italic letters indicate the species. The eigenvalues are 0.658 and 0.462 for the rst and second axis

O.coe I. ele

A.par

E.vir

O.can

-1

DCA axis 1

Fig. 3 Detrended Correspondence Analysis ordination biplot of species and 11 study sites. The eigenvalues are 0.804 and 0.440 for the rst and second axis. Letters indicate the species. (a) Study sites are classied based on geographical locations in the park. (b) Each study site is ordinated individually

In the CCA triplot of species, sites and environmental variables (Fig. 4), the lengths of the arrows of the environmental variables indicate the strength of the relation to the pattern of variation in species composition, and value of each environmental variable increases in the direction of each arrow (ter Braak 1995). The ordination projects species points along the arrows at 90 and the order of the projection points corresponds approximately to the ranking of the species with respect to each environmental variable

that each arrow indicates (ter Braak 1995). Thus, E. viridulum occurred in conditions with relatively high oxygen concentration, owing water and submerged vegetation. It also preferred deeper bodies of water to other species. The occurrence of O. coerulescens was inuenced by the water ow and smaller water bodies and was also strongly inuenced by emergent vegetation. The occurrence of A. isosceles was related to emergent vegetation. I. elegans, C. lindeni, A. parthenope, A. imperator, C. erythraea and O. cancellatum were ordinated near the centre of the diagram, showing characteristics of typically ubiquitous species, although O. cancellatum showed a relation to relatively high salinity and large water bodies. Since O. cancellatum did not show this trend on DCA as a typical ubiquitous species, it would be safer to conclude that I. elegans, C. lindeni, A. parthenope, A. imperator and C. erythraea were the most eurytopic species. The location of sites relative to arrows indicates the environmental characteristics of the sites (ter Braak 1990). Both site 12 and site 10 in the south-west area showed relations indicative of a smaller, owing water body. Site 12 was indicative of the presence of submerged vegetation and deeper water body. Site 10 showed a relation to presence of emergent vegetation. Other sites were close to the

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coordinate origin of the ordination diagram, indicating no strong value of any particular environmental variables at these sites. Since the angle between arrows indicates a correlation between variables (ter Braak 1990), presence of submerged vegetation showed a strong correlation with oxygen concentration in the water.

Discussion Assessing Odonata diversity in the park The Albufera International Biodiversity Group (formerly Earthwatch Europe Project SAlbufera) has been monitoring the occurrence of Odonata species every spring since 1989 and every October and November since 1991. Surveys in summer were conducted only once in August 1993. The lack of information in summer especially in June, July and September has resulted in some ambiguity about the diversity of Odonata in the park. Riddiford and Mayol (1997) report in their summary paper that further recordings of new Odonata species might be expected. Therefore, this study evaluated the reported Odonata diversity in the park and also established a baseline of information for the summer season. The result added one more Zygoptera species, E. viridulum, to the park list. Certain evidence points to the conclusion that this species has probably existed historically in the park: (1) Zygoptera species are generally weak iers compared to Anisoptera and their ability to disperse seems to be limited to between water bodies less than 1 km apart (Conrad et al. 2002); (2) throughout the study period, the species was found in the middle of the park, and its presence was restricted to a small area (site 12 or the beginning of transect D); (3) consistent numbers of individuals were found during each sampling occasion throughout the study period; (4) oviposition was observed during sampling (Table 2); (5) the existence of the species has been reported in Mallorca (Askew 1988); (6) E. viridulum could easily be overlooked amongst the numerous I. elegans occurring in the park, as this latter species is similar in body colour and size and lastly, (7) monitoring of adult Odonata species in the park has never been conducted in June and July (Riddiford and Mayol 1997). Therefore, it is highly unlikely that dispersed individuals from a population outside the park were found in this study. Transects covering a wider range of the park would be helpful to determine if the distribution of E. viridulum is restricted only to site 12 (i.e. the beginning of transect D). S. nigra, a rare species in Europe where it is largely restricted to the Mediterranean region (see Askew 1988), was found in the park in August 1995 (Riddiford and

Mayol 1997) but not subsequently. In this study, two individuals were found during week 7 (Fig. 2b). Because no individuals were found in June and July, it is likely that their ight period starts in August. However, Compte Sart (1963) gives June as the start of the ight period in the Balearic Islands. We suggest further research should be conducted to establish its ight period in order to examine whether the species rarity is due to it being on the wing for a restricted period and thus overlooked, or due to other biological or abiological factors. Aeshna cyanea and Sympetrum sanguineum are two species which have been reported in the park but were not found during this study. The only record of A. cyanea in the park is a report of one individual in May 1993 (Riddiford and Mayol 1997), and there is only one previous record of the species in Mallorca (Compte Sart 1963). Therefore, this species seems to be very rare not only in the park but also in the whole island. Unlike A. cyanea, S. sanguineum was reported regularly in spring and autumn by Earthwatch Europe Project SAlbufera. However, it has not been found in recent years, and the early records may be / are now thought to be largely or all misidentications of brighter S. striolatum and / or S. fonscolombii. In the absence of specimens or photographs, the occurrence of S. sanguineum in the park must be considered unconrmed and doubtful. Establishing conservation priorities The CCA ordination in this study (Fig. 4) indicates some clear relationships between each species and certain environmental characteristics. With baseline information, such as species occurrence, abundance, distribution and ight period, the ordination results can answer more specically what kind of habitats should be protected for a particular species and what areas in the park are more important for their conservation. The current population status of O. coerulescens and A. isosceles, which were found only at site 10, do not appear to be threatened based on the results of the total number of individuals (Table 2) and the change in numbers throughout their ight period (Fig. 2b, c). Although A. isosceles was not seen after week 4, and the total number seen at the study sites was low, the relatively high number at week 1 suggests an established population in the park. This is borne out by the previous report (Riddiford and Mayol 1997) which gave the peak month for this species in sAlbufera as May. Therefore, the disappearance of A. isosceles at week 4 may indicate the end period of its normal range of temporal distribution. Nevertheless, the results of this study indicate that careful monitoring might be required because they were found only at one site. In the CCA ordination (Fig. 4), both species show a strong

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correlation with the presence of emergent vegetation, which distinguishes site 10 from other sites. Site 10 appears to provide important habitat for these species. Site 10 has been excluded from the annual aquatic invertebrate surveys since 1995 (Chapman 1998). Considering its value for the occurrence of particular Odonata species, re-inclusion of this site for the regular survey should probably be considered. The CCA ordination (Fig. 4) shows that E. viridulum also requires a particular type of habitat that is found only in site 12. Flowing water and presence of submerged vegetation are the most important environmental characteristics for this species, but the key habitat characters, distinguishing this species occurrence at site 12 from site 10, are the presence of submerged vegetation and higher concentrations of dissolved oxygen. Any change in these habitat characteristics at site 12 might lead to a direct threat of the existence of this species in the park. In addition, site 12 yielded the highest total number of individuals and species richness (Table 2). Therefore, this site is important, not only for the conservation of E. viridulum, but also due to its high Odonata diversity. According to Watson et al. (1982), dragonies that usually breed in owing water such as streams or rivers tend to stay close to their breeding grounds and do not disperse widely. Accordingly, when their limited dispersal ability is considered, protecting site 10 and site 12 is crucial for maintaining their populations. More importantly, environmental quality in the south-west area should be carefully monitored as the particular environmental factors shown at these two sites in CCA are probably related to the underground springs in the area. Some sites show relatively high number of individuals, namely sites 12, 19, 26, 27, 29 and 30 (Table 2). In the CCA ordination (Fig. 4), sites 19, 26, 27, 29 and 30 do not show any particular environmental patterns. Therefore, more eurytopic species such as A. parthenope and C. erythraea appear to favour these sites. However, the specic environmental characteristics at site 12 which attract E. viridulum do not seem to prevent eurytopic species from coming to the site, although relatively small numbers of eurytopic species at this site and their relationship to environmental factors suggest that they have a lower preference for those habitat characteristics. The availability of certain habitat types is therefore important for stenotopic species such as O. coerulescens, A. isosceles and E. viridulum. Although eurytopic species such as A. parthenope and C. erythraea do not require any specic environmental characteristics, high salinity and low oxygen concentration in water could limit the availability of potential habitat. Site 20, which had the highest mean of salinity (37.45 g/l), was found to have only one species (I. elegans) and site 3, which had the least dis-

solved oxygen (0.05 mg/l), showed the second least number of species (I. elegans and O. cancellatum) when A. parthenope and C. erythraea, for both of which only one individual was found, are excluded. Both sites also showed the least and the second least number of total individuals. Site 27 had the second lowest oxygen concentration (0.76 mg/l) and site 19 had the second highest salinity (12.86 g/l) and, contrary to our expectation, both sites had higher number of species and individuals indicating that salinity for site 20 and oxygen concentration for site 3 may have been beyond their tolerance level. The very low number of species at sites 20 and 3 could raise two hypotheses, each of which is not necessarily exclusive of the other. One is that only certain species of larvae which have high tolerance for high salinity and low oxygen concentration may have hatched from these two sites. In fact, larvae of I. elegans are thought to have high tolerance of anoxic and brackish water (Solimini 1997; Askew 1988), which presumably explains the preponderance of this species in the park. The other hypothesis is that adults intent on breeding may have some mechanism for judging water quality and do not stay there. Little is known regarding whether there is any relationship between water chemistry factors and habitat choice of adult Odonata species. Nevertheless, the latter hypothesis seems very plausible. Many Anisoptera species are strong iers and thought to colonise new water bodies readily (Michiels and Dhondt 1991). Yet, despite this mobility, Anisoptera were generally absent from the north-easternmost corner of the park where water bodies were highly saline because of direct contact with the sea (personal observation). Recent remedial measures have been enacted to stop the direct discharge of seawater from the power station into the park (M Rebassa, personal communication). Monitoring for species assemblages in the north-east area of the park should therefore be continued since the high salinity at site 20 may have been related directly to this discharge. Assessing the indicator properties of Odonata Stewart and Samways (1998) stated that, given a set of species, it was fairly easy to predict the type of water body from which they came. In this study, the indicator properties of Odonata species were established using DCA and CCA. The result of DCA ordination (Fig. 3b) showed that certain sites were correlated with particular species, indicating the potential of Odonata as indicators. The properties of Odonata as indicators were more clearly specied by matching species composition to environmental factors. The CCA ordination (Fig. 4) showed signicant correlation between species and environmental data. For example, if more E. viridulum were found than O. cancellatum, one can easily predict that the

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water body is relatively small with less brackish, owing water and submerged vegetation. Disappearance of E. viridulum could indicate that one of these environmental characteristics changed. One can also imagine that the water has become more brackish when E. viridulum population levels decrease and O. cancellatum population levels increase. In such a case, environmental conditions can be predicted not only by which species are present, but also by the proportion of eurytopic species to stenotopic species (Stewart and Samways 1998). Relatively high brackishness in the north-east area and low brackishness in the south-west area of the park can be expected to be related to the different water sources in the park. This might explain local patterns in species compositions in the DCA ordination reected by sites based on the geographical locations (Fig. 3a). Moreover, the CCA ordination shows that the presence of vegetation and owing water at site 12 and site 10 in the south-west area, and the relatively high salinity at site 3 in the north-east area, appear to be related to these known water sources. Since species composition at each site or each geographical location can be predicted based on the DDC and CCA ordinations, any change in species composition might indicate changes in environmental qualities inuenced by these different water sources in the park. In this study, adult Odonata were investigated. However, conicting opinions have been expressed regarding the use of larval versus adult Odonata as indicators. Watson et al. (1982) point out that larvae might be cryptic or nocturnal, and have microhabitat preferences depending on species, which requires a variety of methods for sampling larvae (Hawking and New 1999). However, the differences in sampling methods make it difcult to compare the data. In addition, it is virtually impossible to identify all early-instar individuals. Therefore, using larvae as indicators does not seem to be practical. Adults are, on the other hand, more conspicuous and easy to observe, and sexually mature adults usually stay near water bodies, suggesting the greater potential of adult Odonata as indicators. Samways (1995) reports the advantage of using adult Odonata in investigating their relationships to environmental characteristics. One possible major disadvantage of using adult Odonata as indicators could be that, despite the numerous studies reporting the relationship of adult Odonata to the features (structures and appearance) of the plant, little is known about the relationship between adult Odonata and water quality in their habitat selection. Although only two water quality variables were measured in this study, results of this study indicated the importance of water quality in inuencing the adult assemblages. One should bear in mind that biological indicators also have a number of limitations (Grillas 1996). Even changes

in a particularly well known species or species assemblages cannot tell us the mechanism by which habitat disturbance has occurred. We cannot use them to determine whether the disturbance is single or interactive, direct or indirect, or whether it is natural or anthropogenic. In other words, it is impossible to use indicator species to pinpoint a specic type of effect or disturbance (Stewart and Samways 1998). More importantly, as this study suggests, indicator species should be chosen or used by criteria that are clearly dened (Landres et al. 1988). Firmly established baseline information, showing spatial and temporal abundance and distribution based on long-term observations, is critical in this regard. The current limited basic information restricts the use of Odonata as biological indicators in sAlbufera at this stage. However, some clear relationships between environmental factors and particular species shown in this study indicate the great potential of using adult Odonata as biological indicators in the park. Spatial and temporal replication of the study would be needed to assess more precisely the potential of adult Odonata as indicators.
Acknowledgements The authors gratefully acknowledge the Balearic Conselleria de Medi Ambient for permission to study at the site, and the park staff for their warm welcome and support. We also thank Pere Vicens for his help with water sampling, Dr. Richard Furnas for his advice on data analysis, Dr. Jeremy Levitt, Mike Mayer, Dr. Peter Mayhew and two anonymous referees for their comments on the manuscript and Mikio Hisamatsu for the provision of literature.

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