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Diversity, Distinctiveness and Conservation Status of The DDI_122.fm Page 257 Thursday, October 25, 2001 8:58 PM Diversity and Distributions (2001) 7, 257–270 BIODIVERSITY RESEARCH Diversity,Blackwell Science Ltd distinctiveness and conservation status of the Mediterranean coastal dung beetle assemblage in the Regional Natural Park of the Camargue (France) JORGE MIGUEL LOBO1*, JEAN-PIERRE LUMARET2 and PIERRE JAY-ROBERT2 1Department of Biodiversidad y Biología Evolutiva, Museo Nacional de Ciencias Naturales, CSIC, c/José Gutiérrez Abascal 2, 28006 Madrid, Spain. E-mail: [email protected], 2Laboratoire de Zoogéographie, Université Paul Valéry Montpellier III, Route de Mende, F-34199 Montpellier Cedex 5, France. *Corresponding author. Abstract. The Mediterranean region as a whole distinctiveness, dune and marsh edges are charac- has the highest dung beetle species richness terized by the occurrence of two rare, vulnerable, within Europe. Natural coastal habitats in this specialized and large roller dung beetle species of region are among those which have suffered severe the genus Scarabaeus. As with other Mediterranean human disturbance. We studied dung beetle divers- localities, current findings suggest a recent decline ity and distinctiveness within one of the most of Scarabaeus populations and the general loss of important coastal protected areas in the west Euro- coastal dung beetle communities in Camargue. Mediterranean region (the regional Park of Camargue, (3) The comparison of dung beetle assemblages southern France) and made comparisons of dung between the Camargue and Doñana shows that, beetle assemblages with other nearby Mediterra- in spite of the low local dung beetle species rich- nean localities, as well as with other coastal pro- ness in the Camargue, the regional dung beetle tected area (Doñana National Park, Spain). Our diversity is similar between both protected areas. finding showed that: (1) The species richness of Unique historical and geographical factors can coastal habitats in the Camargue is low and only explain the convergence in regional diversity as grasslands showed a similar level of species rich- well as the striking divergence in the composition of ness and abundance to inland habitats of other dung beetle assemblages between both territories. Mediterranean localities. The unique habitats of the coastal area (beaches, dunes and marshes) are Key words. Scarabaeoidea, dung beetles, insect largely colonized by species widely distributed in coastal conservation, Mediterranean region, the the hinterland. (2) In spite of their low general Camargue, Doñana. long history of agropastoralism, the seminatural INTRODUCTION and pastoral ecotopes of open forests, shrublands, The Mediterranean region represent one of the woodlands and grasslands, together with the agricul- most ancient disturbed areas in Europe. For approxim- tural ecotopes of patch- and hand-cultivated rock ately 6000 years (dependent upon geographical polycultures, have created a mosaic of landscapes location), Mediterranean ecosystems have been (Naveh & Vernet, 1991). This mosaic of habitats under the influence of human activities (Vernet & extends into coastal areas, characterised by vari- Thiébault, 1987; Reille et al., 1996) which have ous environmental conditions such as aridity in contributed to their present multidimensional sand dunes and humidity at the edges of marshes, heterogeneity (di Castri, 1981). Throughout the combined with the omnipresence of salt and the © 2001 Blackwell Science Ltd. http://www.blackwell-science.com/ddi 257 DDI_122.fm Page 258 Thursday, October 25, 2001 8:58 PM 258 J. M. Lobo et al. occurrence of several soil textures. Traditionally, To carry out these objectives, the dung beetle sheep and cattle grazing occurred on the hinter- assemblages of coastal habitats of the Camargue land and at the edges of coastal wet marshes, will firstly be reviewed to estimate the species fertilizing these areas and giving dung beetle spe- diversity and to identify the most uncommon and cies (Coleoptera, Scarabaeoidea) the opportunity specialized species, as well as to identify their pre- to extend and increase in abundance. Today, due ferred habitats (local scale analysis). Subsequently, to the development of mass tourism and its correl- the specificity of the coastal dung beetle fauna in ative urbanization (mainly in coastal areas), the the Camargue will be compared with the fauna seminatural habitats have become more and more of the hinterland (regional scale analysis). Then, patchily distributed, and many sandy habitats the species richness and composition of the dung have changed or have been destroyed and grazing beetle fauna of the Camargue will be compared has declined. to that of the Doñana National Park in Spain The dung beetles include three families: (Lobo et al., 1997b), which is the other import- Scarabaeidae, Aphodiidae and Geotrupidae. Geo- ant protected coastal territory located in the trupidae and most of the Scarabaeidae (except western Mediterranean region (interregional Scarabaeini) species are tunnelers, digging vertical scale analysis). The results of these studies were tunnels below the dung pat, and carrying dung used to estimate the probable current conserva- into the bottom of the burrow. Scarabaeini are tion status of coastal dung beetle species within rollers, making and rolling away a ball of dung the Camargue. and digging a tunnel outside the dung pat. The small bodied Aphodiidae belong a third guild (the dwellers) as they feed and nest inside the MATERIALS AND METHODS dung pats. The decline of agropastoralism in the last few decades has led many species that were Dung beetle information once very abundant at the beginning of 20th century to become very scarce (Johnson, 1962; Local scale Leclerc et al., 1980; Lumaret, 1990; Väisänen & Dung beetles were trapped from 10 May to 19 June Rassi, 1990; Biström et al., 1991; Lumaret & Kirk, 1996 in seven coastal localities in the Camargue 1991; Miessen, 1997; Lobo, 2001). Coastal dung (Fig. 1). These constituted a representative transect beetle assemblages are very sensitive to human that covered the major coastal habitats, from disturbance, especially when these disturbances seaside to inland. Six distinct habitats were have impacted upon Mediterranean biotopes identified and sampled, namely beach, pre-dune, inhabited for rare and localized species (Pyle et al., dune, marsh, forest and grassland habitats 1981; Balleto & Casale, 1991; Gray, 1991). In the (Table 1). There were good weather conditions case of dung beetles, alteration of most of the coastal (i.e. low wind, and sun) throughout the sampling biotopes of the west Mediterranean area since the period. Pitfall traps baited with dung were used 1960s has strongly affected many roller popula- to collect beetles. Each trap consisted of a plastic tions, reducing the size of their range, with a con- basin 210 mm in diameter containing a preserving sequent decrease of species richness (Lobo, 2001). fluid (a water–liquid soap mixture) and buried to The aims of the present paper are to (i) estab- its rim in the soil. Fresh cattle dung (c. 350 g) lish the current dung beetle fauna composition was placed on a wire grill on the top of the and species richness of the Regional Park of basin. All the pitfall traps were exposed only Camargue (southern France), one of the most once during the trapping period. The sampling important protected coastal areas in the west Euro- effort (number of pitfall traps per habitat) was Mediterranean region; (ii) estimate the dung beetle roughly proportional to the habitat area (between diversity and the faunal distinctiveness of this 3 and 58 traps per habitat). In total, 115 baited area with regard both to nearby Mediterranean pitfall traps were set up. A small number of traps localities and to other Mediterranean coastal attracted the beetles within 24 h (n = 30), whilst area with similar characteristics; and (iii) make a most traps were in position for 48 hours before preliminary evaluation of the current conserva- beetles were caught. The mean distance between tion status of the Camargue dung beetle species. the pitfall traps within a locality was c. 10 m. © 2001 Blackwell Science Ltd, Diversity and Distributions, 7, 257–270 DDI_122.fm Page 259 Thursday, October 25, 2001 8:58 PM Dung beetle species diversity in the Camargue 259 over 24 or 48 h being minimal. Moreover, in the Mediterranean, only a few dung-baited pitfall traps are needed to give a good picture of the structure and composition of local dung beetle assemblages. Lobo et al. (1998) demonstrated that 15 traps allowed the collection of 95% of the species present within a c. 1 km2 localitiy, and that only 2 traps resulted in the collection of 53% of species; in terms of total dung beetle abundance and biomass, these species accounted for c. 85% of the complete inventory. With regard to the effect of season on sampling, previous studies have shown that the dung beetle inventory of a Fig. 1 Location of the seven sample sites in local assemblage when recorded in spring ranged southern France. E: Espiguette; Q: Quatre Maries; from 70% to 80% of the complete annual inventory G: Grand Radeau; B: Baisse de Mouillot; M: (Martín-Piera et al., 1992; Lobo & Martín-Piera, Montille de Charles; L: Longue Montille; R: Radeau 1993; Lobo et al., 1997b). To test if the total de la Foux Vieille. The Mediterranean region, which number of trapped species was an underestimate included the whole of the Camargue area was divided into 17 squares of 0.18° × 0.18° (approximately of the true species richness along the transect, we 304 km2). Insert shows the relative positions of the calculated the species accumulation curve, i.e. the Camargue (France) and the Doñana Park (Spain) cumulative number of recorded species as a compared in the interregional scale analysis. function of the randomised number of pitfall traps (Colwell & Coddington, 1995). For this analysis, the EstimateS statistical software pack- The sampling undertaken at a local scale is age (Colwell, 1997) was used. To eliminate the evidently not comprehensive. However, due to effect of pitfall trap order, 100 randomizations of the adaptation of dung beetles to colonize highly the pitfall order were computed.
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