The altitudes of the sampling stations ranged from 750 to 800 m

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perature) and very low annual rainfall (350 mm mean) – determine the temporary nature of many of the shallow lakes.

The altitudes of the sampling stations ranged from 750 to 800 m. Some of the main environmental characteristics of the sampling sites are given in Table 1.

Vegetation, either emergent, aquatic or both, is abundant and sometimes occupies the entire surface of a shallow lake. The dominant vegetation is formed by emergent macrophytes such as Phragmites australis, Typha domingensis, Typha latifolia, Scirpus lacustris, Scirpus maritimus, Sparganium erectum or Juncus spp. and aquatic species including Drepanocladus

aduncus (a bryophyte), Charaspp. (a charophyte), Polygonum amphibium, Potamogeton spp., Ranunculus spp. or Hippuris vulgaris.

Sampling

Aquatic Coleoptera were collected from 12 shallow lakes throughout a year (spring, summer and autumn 1998 and winter 1999) using a 250 µm mesh D-framed pond net. The captures were made close to the shore by sweeping three 4 m strips, paying attention to particular microhabitats. Two 10 m transects from the shore towards the middle were also sampled at each lake. A fine mesh strainer was used to collect the specimens floating on the surface after sweeping the bottom with the net.

This semiquantitative method (area sampled and time spent sampling were similar at each site) allows structural parameters of the assemblage of aquatic Coleoptera at each site to be com- pared.

Environmental variables

The following environmental variables were recorded in situ at the same time as the Coleoptera were sampled: water temperature, pH, conductivity and dissolved oxygen concentration.

The water temperature was measured at different times during the day and occasionally on different days, and was therefore not used in the analysis. Along with the physical and chemical variables, the following environmental parameters were considered: area, perimeter, maximum depth and vegetation. The following categories were established for water permanence, depending on the number of months per year water was present in the lakes based on observations made over the last six years:

ephemeral (1–2 months), very temporary (> 2–6 months), temporary (> 6–9 months), semipermanent (> 9–12 months) and permanent (always contained water).

The following types of vegetation were considered: emergent macrophytes, aquatic macrophytes other than mosses, bryophytes and filamentous algae. Each type was assigned a value between 0 (absence) and 5 according to the degree of cover.

Data processing

The parameters used to assess the structure of the assemblage were species richness, abundance and diversity, calculated using the Shannon index (H’).

290

Fig. 1. Map of the study area showing the sampling sites.

0 5 0 4 1

30 289

0.33 8.4

1980 8.00

8.50 4

Esclusa n° 4

0 3 4 2 3

50 579

0.74 4.2

948 7.92

10.77 3

Capillas

0 3 0 5 2

30 1044

2.05 3.1

721 7.28

8.45 3

Abarca

0 0 0 5 2

30 654

1.25 3.3

1107 7.30

8.45 Piña de 3

Campos

0 0 1 5 5

250 3454

29.35 2.6

783 7.39

9.70 2

Valdemudo

0 0 0 5 3

100 2022

8.34 3.0

1333 7.43

8.85 2

Valdemoro

0 0 2 2 3

80 317

0.47 3.0

1396 7.59

9.00 2

El Rosillo

0 0 3 4 3

120 367

0.76 6.0

665 7.91

9.38 2

El Juncal

2 0 4 3 5

150 1393

2.22 6.4

466 7.76

11.38 1

Belmonte

0 3 4 3 4

120 1102

2.65 5.0

800 7.63

12.53 1

El Deseo

0 0 4 1 3

200 280

0.52 4.2

841 7.93

13.88 Casas del 1

Rey

0 1 5 2 5

180 990

4.70 6.9

608 8.14

13.13 1

Besana

algae mosses aquatic

macroph.

emergent macroph.

perman.

depth (cm) perimeter

(m) area

(hect.) oxygen

(mg/l) conduct.

(µS/cm) pH

temp.

(^oC) TWN group

TABLE 1. List of sampling sites and values of the environmental variables measured. Temperature, pH, conductivity and oxygen concentration are the average of four samples. Sites are ordered according to the TWINSPAN group (“TWN group” column) to which they belong. The variable “aquatic macrophytes” refers to macrophytes other than mosses.

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All the statistical analyses were carried out on global abun- dances: that is, total number of individuals collected at a site over the sampling period. The TWINSPAN programme was used to classify the localities. The cut-off levels for the pseudo- species were fixed at 0, 3, 10 and 50.

Sites and species were ordinated by DCA (Detrended Correspondence Analysis) using the canoco statistical package for Windows, version 4.02. The Kolmogorov–Smirnov test was used to verify that the DCA scores, the assemblage parameters and the variables, with the exception of “bryophytes” and

“algae”, were normally distributed. A correlation analysis between the site scores and the environmental variables was used to aid the interpretation of the DCA axes. The Pearson product–moment correlation coefficient was used for variables with normal distributions and the Spearman’s rank correlation coefficient for the remaining two. The correlations between assemblage parameters and variables were made in the same way.

RESULTS

Environmental variables

Table 1 summarizes some of the characteristics of the shallow lakes and the average values of the variables measured during the study. The most outstanding features are the differences in area and degree of water permanence. A wide range of macrophyte cover (both emergent and aquatic other than mosses) is also evident, varying from 0 (absence) in some of the sites to 5 (almost com- pletely covered).

Assemblage composition

Ninety two species of aquatic Coleoptera were identi- fied among the 3,551 adults captured (Table 2). Of these, 44 belong to the Hydradephaga (Gyrinidae, Haliplidae, Noteridae, Hygrobiidae and Dytiscidae) and 48 to the Polyphaga (Hydrochidae, Helophoridae, Hydrophilidae, Hydraenidae, Dryopidae, Elmidae and Heteroceridae).

Most of them have a Palaearctic distribution. Only two species endemic to Iberia were collected: Graptodytes castilianus, restricted to the northern Iberian Meseta, Ibe- rian Cordillera and the Ebro depression, and Helophorus seidlitzii, which is widely distributed throughout the Pen- insula (Ribera et al., 1998). Whilst H. seidlitzii was common in the shallow lakes, only 4 G. castilianus speci-

mens were captured. Ochthebius minimus, Limnebius fur- catus, Helophorus brevipalpis, Berosus affinis, Enochrus nigritus, Noterus laevis and Graptodytes bilineatus were the most abundant species.

Ecological parameters of the community

Table 3 gives the global values recorded during the study for the structural parameters of the community.

Richness for each site ranged from 22 (Esclusa n° 4) to 43 (Belmonte), though most of them had more than 30 species. Abundance values varied between 108 specimens captured in Valdemoro and 702 in Deseo, while diversity values ranged between 2.62 (Esclusa n° 4) and 4.43 (Abarca). There were more Coleoptera in spring (74 species, 1,406 specimens) and winter (72 species, 1,049 specimens). Abundance and richness decreased substan- tially in summer (488 specimens belonging to 42 species) and autumn (608 specimens from 47 species), partly because some of the lakes dried out in these seasons (4 during summer and 3 in autumn).

Richness was positively correlated with permanence (r

= 0.737, P = 0.006) and aquatic macrophyte cover (r = 0.716, P = 0.009) and negatively correlated with conductivity (r = –0.67, P = 0.017). Diversity showed a negative correlation with conductivity (r = –0.753, P = 0.005). No significant correlations were detected between abundance and any of the variables.

Classification and ordination

TWINSPAN classified the sites into four groups (Fig.

2). Table 1 shows the values of the variables obtained at each locality. Variables that apparently discriminate the TWINSPAN groups are water permanence, conductivity and to a lesser extent, depth. However, sites are not clearly separated by any variable, with the exception of Esclusa n° 4. Group 1 includes all the permanent (permanence value = 5) and semipermanent (permanence value

= 4) lakes except Valdemudo; they are all very deep.

Group 2 is formed by most of the localities with interme- diate persistence and that are comparatively deep. The third group includes two very temporary shallow lakes (permanence = 2), and Capillas (permanence = 3).

Finally, the fourth group is formed exclusively by Esclusa nº 4, the most ephemeral lake. The lakes in the last two groups are shallow.

Table 2 shows the percentage of localities within each TWINSPAN group at which each species was collected.

Most species were widely distributed within the study area and are indifferent to the characteristics of the lakes, though they were often absent from ephemeral ponds (group 4). One group of species including mainly Ochthe- bius viridis 2, Hydrophilus pistaceus, Noterus clavicornis, Hygrobia hermanni, Bidessus goudoti and Cybister later- alimarginalis was associated with the more permanent shallow lakes (TWINSPAN group 1). In contrast, only a few species appear to be associated with very temporary or ephemeral waterbodies: Hydraena rugosa, Helophorus grandis and Haliplus variegatus.

The eigenvalues of the first four DCA axes were 0.687, 0.402, 0.125 and 0.024, and the cumulative percentage of Fig. 2. Dendrogram of TWINSPAN end-groups for sites.

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variance corresponding to the species data 24.7, 39.2, 43.7 and 44.6. Figs 3 and 4 show the ordination plots for sites and for species on axes 1 and 2. The TWINSPAN groups are also shown in Fig. 3. Group 4 is clearly separated from the others at the positive end of the first ordi-

nation axis. Group 1 is separated at the opposite end, though less obviously so, while groups 2 and 3 overlap on the first axis. The first axis has a long gradient (4.07 stan- dard deviation units), but only Esclusa n° 4 is signifi- cantly separated from the other localities. Generally speaking, localities differing by 4 s.d. are not expected to have any species in common (Ter Braak, 1995), so it would be reasonable to expect different assemblages of species at the extremes of the gradient represented by axis 1. This points to the existence of an important faunistic turnover along the first ordination axis. This axis is positively correlated with conductivity (r = 0.621, P = 0.031) and negatively correlated with permanence (r = –0.617, P

= 0.032). There were no significant correlations with the other variables. We must therefore assume that this axis represents a water-permanence gradient: most temporary waterbodies are located at the right end of axis 1, and the permanent and semipermanent ones at the left end of this axis. The length of the gradient of the second axis is 2.97 s.d. No significant correlations were found between the scores corresponding to this axis and any of the environmental variables. According to the ordination analysis, the following species were particularly abundant in the ephemeral ponds (positive end of axis 1): Hydraena rugosa, Berosus signaticollis, Dryops algiricus, Dryops similaris, Haliplus variegatus, Graptodytes bilineatus and

292

Fig. 3. Plot of the site scores for axes 1 and 2 of the DCA.

Also shown are the TWINSPAN groups.

Fig. 4. Plot of the species scores for axes 1 and 2 of the DCA.

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