Abstract :
[en] Introduction
The purpose of this study is to verify one of the principal rules of ecology: the principle of exclusive competition, by using the sibling species of long-eared bats present in Belgium as a model: the brown long-eared bat (Plecotus auritus) and the grey long-eared bat (Plecotus austriacus). Two similar species present in the same region and, what is more, share the same roost, must differ according to certain aspects of their ecological niche in order to be able to co-exist (MacNab 1971).
In order to determine the resource sharing mechanisms between the two species, we have compared the three important dimensions of their ecological requirements: the use of trophic resources by means of faecal analysis and the use of space and time by radio-tracking.
Trophic resources
One hundred and thirty samples (4688 droppings, 6388 occurrences) taken from 5 single-species colonies of P. austriacus, 5 single-species colonies of P. auritus and from a mixed colony have been taken into account in the analysis of the summer diet.
With the help of a first model constructed with the aid of generalized linear regressions, we have shown that, independently of the cohabitation conditions (single-species and mixed colonies), the two species have similar dietary requirements. Qualitatively, the same types of prey are consumed : Lepidoptera, Diptera (Craneflies, Cyclorrhapha and others Diptera), Coleoptera, Arachnids and Dermaptera. Small quantitative differences of between 3% to 8%, were seen to exist : P. austriacus consumes slightly more Coleoptera and Lepidoptera and fewer Craneflies than P. auritus.
With the help of a second model enabling interspecific comparison of the diet according to whether the animal belonged to the single-species colonies or the mixed colony, we were able to show that the differences were more marked.
When P. austriacus belong to single-species colonies, with reference to single-species colonies of P. auritus, predicted value indicate that they consume more Coleoptera (+ 5,0 %, p = 0,001) Lepidoptera (+ 14,4 %, p < 0,0005) and fewer Arachnids (- 8,3 %, p < 0,0005), Dermaptera (- 8,8%, p < 0,0005) and Craneflies (- 9,5 %, p = 0,009). This leads us to believe that the proportion of non-flying or diurnal prey gleaned by P. auritus is 2,5 times greater than that of P. austriacus.
On the other hand, when P. austriacus belongs to the mixed colony, in reference to the P. austriacus single-species colonies, it changed its feeding behaviour. The predicted consumption of Arachnids and Dermaptera is higher, 14,5 % (p < 0,0005) and 11,3 % (p < 0,0005) respectively, while the consumption of Lepidoptera is much less (- 25,6 %, p < 0,0005). The syntopic P. austriacus also consume more craneflies (+ 7,9 %, p = 0,034) but fewer Coleoptera (-11,1 %, p = 0,009). This shows that P. austriacus posesses surprising adaptation abilities. The diet of the P. auritus of the mixed colony in relation to the single-species P. auritus, only showed slight variations on the other hand : - 7% (p = 0,006) for the Cyclorrhapha and - 4% (p = 0,002) for the Dermaptera.
In syntopy, the study of seasonal variations has shown that the quantitative differences were significant at the end of gestation and lactation when the energy requirement is at its highest. In June, P. austriacus consumes more Arachnids (p = 0,046), in July, more Arachnids (p = 0,020) and fewer Lepidoptera (p = 0,020). In August, they consume more Dermaptera (p = 0.019), fewer Coleoptera (p = 0.032) and Lepidoptera (p = 0.034).
Winter trophic ecology
Our study has also led to the discovery of a little-known aspect of the ecology of chiroptera: winter trophic ecology. The results have shown that long-eared bats consume at least 70% of their prey which they capture by gleaning (Spiders, Dermaptera, Cyclorrhapha). These results have made it possible to confirm that the two species possess the ability to glean their prey.
Spatial and temporal resources
Twenty-two Plecotus were used for the analysis : seven P. auritus in Gozin, six P. austriacus in Gembes, five P. auritus and four P. austriacus in Pondrome (mixed colony). Eighty-four nights of monitoring corresponding to the discovery of 111 hunting grounds were used for the analyses.
The results show that the use of space by P. auritus and P. austriacus contain similarities. The distance of the hunting grounds, the number of grounds visited per night, the duration of their use, their surface and the individual area of daily activity are similar.
In the same way, contrary to what the literature suggests, it seems that P. auritus possesses the ability to exploit the open or semi-open areas like the meadows surrounded by linear woodland elements, while P. austriacus possesses the ability to exploit equally well the closed areas such as woods.
In syntopy, the two species showed a different selection of habitats (grasslands, leafy woods,, gardens, edges for P. auritus and leafy or coniferous woods, gardens and edges for P. austriacus), P. auritus spending 64% of hunting time above the grassland against 83% in the woods for P. austriacus. On the other hand, in comparison with the two single-species colonies, the division of hunting time around the roost is more spread out in terms of space and the number of hunting grounds visited by night is more important.
No difference in the exploitation of temporal resources was detected; the later departures from the Gozin colony were attributed to the presence of artificial lighting placed in front of the roost.
The mechanisms allowing resources partitioning
In the case of single-species colonies we have shown the existence of trophic resources partitioning. In fact, P. austriacus showed a diet less rich in Arachnids, Dermaptera, Craneflies and richer in Coeloptera and Lepidoptera than P. auritus. However, in syntopy, P. austriacus clearly modified its diet by consuming more Arachnids and Dermaptera to the detriment of Coleoptera and Lepidoptera. The consumption in Arachnids and Dermaptera remains higher than those of single-species P. auritus.
Whether for single-species colonies or mixed colonies, trophic resources partitioning is quite evident. As requirements in terms of syntopy were modified, the results lead us to assume that inter-specific competition exists between this sibling species.
The differences observed in the diet are probably dictated by a different use of the habitats. Indeed, in relation with the ecology of prey, the results of habitats use have made it possible to show that habitat partitioning occurs between the two species of long-eared bat.
Our results therefore constitute a good illustration of the principle of competitive exclusion. Whether in the case of single-species colonies or mixed colonies, the two species share the trophic and spatial resources which allow them to co-exist.
Recommendations for the conservation of the species
To satisfy the requirements of the summer diet, it is necessary to maintain a network of diverse habitats composed of gardens, meadowlands, humid zones, hedges, bushes, tree alignments, leafy or mixed woodlands, preferably hygrophilous, allowing for the development of undergrowth and clearings. The results of the selection of surface habitats have globally confirmed the importance of these habitats used as hunting grounds. The selection of linear habitats such as tree-lines, hedges, bushes and edges also show the importance of ecological networks and the potentially damaging effects of the fragmentation of habitats on the bat population.
In order to integrate the spatial requirements of P. austriacus, we recommend the establishment of a minimum protection perimeter of 3500 metres around the nest. With regard to P. auritus, it would be judicious to protect a minimum perimeter of a radius of 2000 metres.
