Ecotourism disturbance to wildfowl in protected areas: historical, empirical and experimental approaches in the Camargue, Southern France

Ecotourism is becoming very popular, especially in protected areas where wildlife concentrate and is easier to observe, but the consequences of associated disturbance have seldom be quantified other than in the short-term, making the sustainability of this activity untested. We combined a historical, an empirical and an experimental approach to assess the long-, medium- and short-term consequences of disturbance to wintering wildfowl (Anatidae) in a wetland of international importance in the Camargue, Southern France. In the short-term, disturbance made teal (Anas crecca) move away temporarily from observation blinds without leaving the waterbody. Wildfowl fed more after disturbance, disrupting their normal resting activities. In the medium-term, waterbodies with more tourists did not host fewer birds: conversely the most heavily disturbed one hosted the highest wildfowl density. In the long term, wildfowl numbers were not related with the number of visitors. When practiced with appropriate guiding of people, and where appropriate facilities are provided to limit human disturbance as done here, ecotourism may not affect wintering wildfowl other than reversibly in the very short term. The legitimate demand of the public for access, even in fragile protected areas, may therefore be sustainable under some conditions.     

The biology of the common carp, Cyprinus carpio L. in the Camargue, southern France

Carp were sampled monthly during one and a half years in the Etang du Vaccarès, Camargue, southern France using fyke nets. Densities of carp in temporary marshes and canals were determined using rotenone. Carp is the dominant species in biomass in the fresh and brackish waters (14%0) with densities from 11–58 kg/ha in temporary marshes and up to ten times greater in the canals. Scales were used to determine age and growth of 201 carp. The relative condition factor of female and male mature and unsexed fish are not statistically different throughout the year and follow the variations of gonadal development in spring. The best predictive equation of fecundity, Log F = 7.03504 + 0.83829 log ovary weight, was found using a stepwise multiple regression. The sex-ratio in the temporary marshes (♂ : ♀; 1.66–1.76) is greater than the general sex-ratio of the monthly samples (0.67). The diet of carp is characterized by the presence of large numbers of seeds and the absence of large benthic invertebrates. Predation by fish-eating birds and carnivorous fish is minimal, and the annual drying of many marshes is considered the main mortality factor., Accepted 22 July 1980     

Lead poisoning in wild populations of Marsh Harriers Circus aeruginosus in the Camargue and Charente-Maritime, France

Lead poisoning of waterfowl, through the ingestion of spent gunshot, has been recognized as a mortality factor for over a century. However, in Europe relatively little attention has been paid to raptors that may ingest shot embedded in the flesh of prey. The present study examines the incidence of lead poisoning in wild Marsh Harriers Circus aeruginosus trapped at two sites in France, the Camargue and Charente-Maritime.
Eight captive Marsh Harriers (controls) had blood lead (PbB) concentrations of 5.3–10.8 μg per dl. Of 94 wild birds trapped during the winters of 1990/1991 and 1991/1992 either in baited clap traps or in mist nets at night roosts, 29 (31%) had elevated (>30 μg/dl) PbB concentrations and 13 (14%) had concentrations indicative of clinical poisoning (> 60 μg/dl). Similar percentages of birds caught using the two trapping methods had elevated (>30 μg/dl) PbB concentrations. However, clap netting appeared to select for more heavily contaminated birds and for juvenile birds, although the two are not necessarily correlated. In the Camargue, significantly more females than males trapped in clap nets had elevated PbB concentrations.
The incidence of shot in regurgitated Marsh Harrier pellets increased significantly between October and December, indicating increased exposure to lead as the hunting season progressed. There appeared to be a parallel increase in PbB concentrations throughout the hunting season, although this could not be clearly demonstrated as sex ratios were different during different sampling periods.
Elevated PbB concentrations of harriers in the Camargue are likely to result primarily from the ingestion of shot in the flesh of crippled or unretrieved waterfowl and in Charente-Maritime, from eating crippled or dead mammals.
Other raptors at risk from lead poisoning and solutions to this problem are discussed.     

Response of Juniperus phoenicea on sandy dunes in the Camargue (France) to water and saline constraint in summer

The ‘Bois des Rièges’ woodland occurs on the relic littoral dunes in the National Reserve of the Camargue. Although surrounded by brackish ponds and saline lands with very salt groundwater close to the surface, the dunes are covered with non-halophytic vegetation of herbaceous shrubs and trees, including the dominant shrub Juniperus phoenicea. This is due to the presence of a freshwater lens beneath the dunes supplied by rains, and floating in hydrodynamic equilibrium upon the saline aquifer. The importance and duration of the freshwater stock depends on the precipitation-evapotranspiration balance as well as on the size of the dune. From the end of the spring to the autumn rain period this freshwater stock is considerably reduced, while the capillary potential in the zone of aeration of soil and the osmotic potential of the soil solution, influenced by capillary rise of the brackish groundwater, decrease. In summer the vegetation is thus subjected to severe drought caused by lack of water or salt excess, to which it must adjust its biological activity. The water relationships in the soil-plant system have been studied along a transect between the top and the borders of a dune surrounded by saline lands. Using simultaneous water potential measurements of the sunny and shady sides of Juniperus trees daily and seasonal transpiration regulations were studied. The preliminary results indicate that trees on the border of the dune as well as on tops are absorbing water from the same freshwater lens in the middle of the dune.     

Kinetics of FeS-mediated denitrification in sediments from the Camargue (Rhone delta, southern France)

Abstract Denitrification rates were measured in sediments after the addition of different concentrations of FeS. A decrease of the denitrification rate was observed when high concentrations of ferrous iron (> 10 mM) were present. In the experiments with no significant concentrations of free Fe²⁺, the relationship between NO₃⁻ reduction and FeS concentration followed Michaelis and Menten kinetics. The maximum rate was 0.273 mmol l⁻¹ d⁻¹, 6 times as much as the basal rate 0.046 mmol l⁻¹ d⁻¹, which was attributed to organic matter; the K_s was 1.45 mM FeS. The stoichiometry of the overall reaction involving NO₃⁻ reduction and the concomitant S²⁻ oxidation was also investigated. Measured ΔS/ΔN ratios ranged between 0.55 and 0.64, with ΣH₂S + SO₄²⁻ changing less than 10%. These values agree with the theoretically expected value of 0.56.     

Urea fertilization and the N-cycle of rice-fields in the Camargue (S. France)

The use of urea as N-fertilizer for rice production has been studied for European conditions. In field and pot experiments, urea appeared to be more efficient than ammonia or nitrate. The rice yield could be maintained at maximal local production values by broadcasting 100 kg ha^-1 of urea-N instead of using 200–200 kg ha^-1 of ammonia or nitrate nitrogen. The best results were obtained by adding the urea about 5–8 weeks after sowing. When this was done, splitting the dose did not increase the efficiency. In laboratory experiments it was shown that urea diffuses and hydrolyses rapidly to ammonia in the anoxic layers of the flooded soils. Because this happens in the anoxic zones, the ammonia cannot be oxidised to nitrate, and is therefore less prone to denitrification, but it is adsorbed onto the soil particles. The upward diffusion of ammonia is slow, rendering it available to the rice roots for a relatively long time. The diminished use of N fertilizer is profitable for the rice farmer and helps to protect the surrounding environment.     

Prevalence of West Nile virus neutralizing antibodies in wild birds from the Camargue area, southern France

The Camargue area of southern France experienced the re-emergence of West Nile Virus (WNV) in the late summer of 2000 and 2004. Immediately preceding the 2004 outbreak, samples were collected from 432 birds of 32 different species captured in mist nets and from 201 Cattle Egret (Bubulcus ibis) nestlings sampled in their nests between 1 April and 12 June 2004. West Nile virus neutralizing titers of >/=40 were detected in 4.8% (95% confidence limit, 2.9-7.5%) of the adult birds and in 1.6% (0.3-4.6%) of the egret nestlings. Migratory passerines had a higher prevalence of WNV neutralizing antibodies (7.0%) than did resident and short-distance migratory passerines (0.8%), suggesting exposure to WNV or a related flavivirus during overwintering in Africa.     

The invasive red swamp crayfish as a predictor of Eurasian bittern density in the Camargue, France

Few data exist on the relationships between food levels and breeding density of the Eurasian bittern Botaurus stellaris , a vulnerable species of high-priority concern in Europe. Concurrent data were obtained on male bittern density and relative food abundance over a 3-year period in two wetlands totalling 2500 ha of Mediterranean reed marsh enclosing 25% of the French bittern population. Food abundance was estimated by sampling up to 25 hydrological units using a beach seine in early June of 2002, 2003 and 2004. The density of booming males in each hydrological unit was obtained by point counts and acoustic triangulation in May of the same years. The impact of food abundance on male bittern density was assessed by general regression models using a forward stepwise procedure with mosquitofish Gambusia affinis , carp Cyprinus carpio , other fish species, amphibians, red swamp crayfish Procambarus clarkii and other invertebrates as prey groups. Of these, only to crayfish abundance was bittern density related, contributing to 45% of the variance observed. When the impact of water level was taken into account, the relative abundance of crayfish explained 56% of the inter-annual differences in bittern density. Because crayfish are rich in calcium and well adapted to fluctuating hydroperiods alternating with drought intervals, they provide a good food source for the bitterns throughout the breeding season at the study sites. The loss of diversity and degradation reported from macrophyte-dominated marshes following crayfish invasion does not seem to apply to reed-dominated wetlands. It is further suggested that the recent increase in bittern numbers in the Camargue, while other French populations were decreasing, could in part be related to red swamp crayfish abundance.     

The labyrinth of nutrient cycles and buffers in wetlands: results based on research in the Camargue (southern France)

Wetlands, especially in the Mediterranean area, are subject to severe eutrophication. This may upset the equilibrium between phytoplankton production in undesirable quantities and a quantitatively desirable macrophyte production. In order to manage this equilibrium, a quantitative knowledge of nutrient input and fluxes is essential and the role of sediments in these processes must be understood. This knowledge can be useful even for agriculture, e.g. rice cultivation, where optimal utilization of fertilizers can lead to an economic benefit.In this article different aspects of nutrient cycles are discussed in view of approaching a sufficiently precise quantification. The nutrient input balance of the Camargue was therefore measured which showed that the input of nutrients with the irrigation water, taken from the river Rhone, roughly equals the quantity of fertilizers added.Phytoplankton growth can be approached reasonably with the Monod model, although there are still many practical problems, such as the influence of the pH on P uptake and the problem of measuring P uptake in the field. The situation is worse for macrophyte growth; quantitative data are scarce and studies have often been carried out with unrealistic nutrient concentrations or without addressing the influence of the sediment. This influence can also include negative factors, such as high concentrations of Fe²⁺, H₂S or FeS, but cannot yet be quantified.The nitrogen cycle in wetlands is dominated by denitrification. Most wetlands have sediments with high concentrations of organic matter, therefore with a large reducing capacity. Besides this process, we have shown that denitrification can also be controlled by FeS. In the Camargue sediments this denitrification is mediated by bacteria from the sulfur cycle; this appeared to be the major pathway. It was shown that a stoicheiometric relation exists between nitrate reduced and sulphate produced. The influence of the temperature was quantified and appeared to be stronger at high organic matter concentrations than at lower ones. Denitrification with FeS means that the bacteria use nitrate also for their N demands, while this is not necessarily the case during denitrification with organic matter.Mineralization of macrophytes is a much slower process than that of phytoplankton, probably because of their high C/N ratio. We could, however, not confirm the general assumption that the addition of nitrogen stimulates this mineralization. On the contrary, we found that two amino acids both with a C/N ratio of 6 had different mineralization rates. The amino acid composition of dead macrophytes and the C/N ratio may be of equal importance.Unlike nitrogen, phosphate is always strongly adsorbed onto sediments. The two mechanisms of the adsorption of inorganic phosphate onto sediments, i.e. the adsorption onto Fe(OOH) and the precipitation of apatite, have been quantified. The adsorption of phosphate onto Fe(OOH) can be satisfactory described with the Freundlich adsorption isotherm: P_ads = A^* (o-P)^B. The adsorption coefficient A depends on the pH of the system and the Ca²⁺ concentration of the overlying water and can be quantified preliminarily by A = a.10(^–0.416*pH).(2.86 – (1.86.e^–Ca2+)). B can be approached by 0.333, which means the cube root of the phosphate concentration. The second mechanism is the solubility of apatite. We found a solubility product of 10^–50 for hard waters. The two mechanisms are combined in solubility diagrams which describe equilibrium situations for specific lakes.The conversion of Fe(OOH) to FeS has a strong influence on phosphate adsorption, although the partial reduction of Fe(OOH) P by H₂S does not release significant quantities of phosphate. Even after complete conversion to FeS only a small part of the bound phosphate was released.Besides the two inorganic phosphate compounds, we established the existence of two organic pools, one soluble after extraction with strong acid (ASOP), the other one with strong alkali. The first pool is probably humic bound phosphate, while the larger part of the second pool was phytate. The ASOP was remineralized during the desiccation of a Camargue marsh; this drying up oxidized FeS, thus improving the phosphate adsorption and decreasing the denitrification capacity. It can, therefore, be an important tool for management. The phytate was strongly adsorbed onto Fe(OOH), which explains the non-bioavailability towards bacteria.The fact that the sediment phosphate concentration can be approached by multiplying the relevant sediment adsorption constant with % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0Jf9crFfpeea0xh9v8qiW7rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaWaaOqaaeaaca% WGVbGaeyOeI0IaamiuaaWcbaGaaG4maaaaaaa!3B8D!\[\sqrt[3]{{o - P}}\] concentration has the consequence that much larger quantities of phosphate accumulate in the sediments than in the overlying water. This means that even if the phosphate input is stopped, the eutrophication will only be reversed very slowly, and not at all, if the shallow waters in wetlands have no through flow — as is often the case in many marshes in Mediterranean wetlands.Abbreviations used o-P = dissolved ortho phosphate (or its concentration) - N_part, P_part = particulate N or P - Tot-N_inorg = Total inorganic nitrogen (= NH₃ + NO ₂ ^– + NO ₃ ^– ) This paper, giving an overview of the research in the sediments of the Camargue, was read during the symposium Nutrient Cycles — A Joy Forever, on the occasion of my retirement, 19th of May 1993 at the I.H.E. in Delft (Netherlands).     

Creation of a nature reserve, its effects on hunting management and waterfowl distribution in the Camargue (southern France)

The eastern portion of the Camargue in southern France is divided into two parts: a natural wetland area of 2800 ha and an agricultural area of 5045 ha. In 1984 and 1989, a new protected area (1000 ha) was created on two contiguous former hunting estates. Analysis of aerial photographs (1968–1998) in conjunction with a field survey revealed, from 1984 to 1998, an increase of management for waterfowl hunting in natural wetlands located on the periphery of the new reserve, and a similar increase in the agricultural area amongst residual wetlands and former rice fields. Based on monthly aerial censuses, the size of the waterfowl population in the winter increased in this part of the delta by a factor of 3.8 following the creation of the reserve. Our results suggest that the creation of a reserve on former hunting estates resulted in greater attractiveness for ducks but also in the development of commercial hunting activity around the edge. This development increased fragmentation and uniformity of natural areas, including the loss of the unique Cladium mariscus habitat in the Camargue and resulted in the spatial expansion of waterfowl hunting areas in agricultural areas. It is associated with an increase in financial profit from waterfowl hunting. Our study highlights unexpected costs for nature conservation associated with the creation of a reserve. It illustrates the need for new models of wetland conservation where protection goals are not restricted to reserves, but also integrate conservation and economic development outside the protected areas.     

In vitro and in situ studies on nitrate disappearance in water-sediment systems of the Camargue (southern France)

Results of in vitro and in situ experiments on nitrate disappearance from water-sediment systems in the Camargue are described.In the in vitro experiments two factors were studied: temperature and organic matter. After a first addition of KNO₃ to these sediments, the concentration of organic matter exerted a strong influence on the disappearance rate of nitrate at 25 °C and 15 °C but not at 2 °C. After a second addition of nitrate at 25 °C and 15 °C the denitrification rate increased by approximately 10%, probably because the activity of the bacterial population had increased.Experiments in situ in freshwater temporary marshes showed that nitrate disappeared at approximately twice the rate at similar temperature in vitro.After the first addition of nitrate in the in vitro experiments the concentration of nitrite in the water above the sediment reached about 10% of the concentration of total dissolved inorganic nitrogen at 2 °C and 15 °C. These high concentrations were not found after the first addition at 25 °C or after the second addition of nitrate at 25 °C and 15 °C. In the in situ experiments, however, high concentrations of nitrite were found.     

Fuelling rates of garganey ( Anas querquedula) staging in the Camargue, southern France, during spring migration

Most species of long-distance migratory birds put on energy stores to fuel their travels. However, recent studies have highlighted the potential costs associated with carrying too much fuel, either through increased predation risk or decreased flight efficiency. Consequently, it is now widely accepted that migratory birds should carry optimal rather than maximum fuel loads. Information from 372 garganey (Anas querquedula) ringed and recaptured at least once during the same spring in the Camargue, southern France, was used to document fuelling rates of individual ducks in relation to environmental variation and individual variation in condition. On average, garganey added very little fuel stores in the Camargue (mean gain per day=0.33 g, less than 0.5% of mean body-mass in total over an average stay of 5 days). Fuelling rates were negatively correlated with body mass at capture, but it cannot be excluded that this pattern was a statistical artefact. Given their body-mass at ringing, garganey could potentially still fly long distances when they stop in the Camargue. It is therefore likely that the aim of their stay in southern France is more for resting than refuelling, a finding that may have implications for the proper management of stop-over sites.Communicated by F. Bairlein     

Rapid response of a long-lived species to improved water and grazing management: The case of the European pond turtle (Emys orbicularis) in the Camargue,France

Among human activities, the effect of habitat management by grazing on population viability is ambiguous. Indeed, beneficial effects of grazing are expected by maintaining open meadows, but overgrazing is supposed to increase mortality by trampling. Grazing has been shown to negatively impact the survival of European pond turtle (Emys orbicularis) in the Camargue. Consequently, a new management plan was defined. We investigated the consequences of this management using capture–recapture methods to estimate variations of population sizes in this managed site and a control site over a 17 years period. Results show an increase of the number of adults and juveniles on the managed site after the management change. Our results suggest that improved water management with flooding in autumn provided better hibernation conditions, and that reduced grazing intensity in autumn/winter likely decreased the risk of trampling. Population size significantly increased in less than 4 years following the management change, probably by the relaxation of density-dependence. It is an original result for a long lived-species supposed to have an important time of resilience to perturbations.     

Regulation of photosynthesis and oxygen consumption in a hypersaline cyanobacterial mat (Camargue, France) by irradiance, temperature and salinity

Short-term effects of irradiance (0-1560 micromol photons m(-2) s(-1)), temperature (10-25 degrees C), and salinity (40-160) on oxygenic photosynthesis and oxygen consumption in a hypersaline mat (Salin-de-Giraud, France) were investigated with microsensors under controlled laboratory conditions. Dark O(2) consumption rates were mainly regulated by the mass transfer limitations imposed by the diffusive boundary layer. Areal rates of net photosynthesis increased with irradiance and saturated at irradiances >400 micromol photons m(-2) s(-1). At low irradiances, oxygen consumption increased more strongly with temperature than photosynthesis, whereas the opposite was observed at saturating irradiances. Net photosynthesis vs. irradiance curves were almost unaffected by decreasing salinity (100 to 40), whereas increasing salinities (100 to 160) led to a decrease of net photosynthesis at each irradiance. Dark O(2) consumption rates, maximal gross and net photosynthesis at light saturation were relatively constant over a broad salinity range (60-100) and decreased at salinities above the in situ salinity of 100. Within the range of natural variation, temperature was more important than salinity in regulating photosynthesis and oxygen consumption. At higher salinities the inhibitory impact of salinity on these processes and therefore the importance of salinity as a regulating environmental parameter increased, indicating that in more hypersaline systems, salinity has a stronger limiting effect on microbial activity.     

Changes in clutch size, brood size and numbers of nesting Squacco Herons Ardeola ralloides over a 32-year period in the Camargue, southern France

Changes in numbers of nesting pairs, clutch size, and brood size of Squacco Herons Ardeola ralloides in the Camargue, southern France were analysed over a 32-year period. The annual numbers of breeding pairs [average 84 pairs pL 30 sd, n = 32 years) exhibited a possible increasing trend, but with considerable variability. This variability is associated with local spring rainfall, which may influence the available surface area of their preferred freshwater habitats. In contrast to population size, clutch size decreased substantially in recent years. Our data do not provide a causal explanation for this change in clutch size, although the reduction temporally corresponds with increasing rice cultivation and with a dramatic increase in the number of Cattle Egrets Bubulcus ibis in the Camargue.