Josep M. Bas

Structure of ground-foraging ant assemblages in relation to land-use change in the northwestern Mediterranean region

The abandonment by humans of marginal and less productive zones signifies an important change in land use in North Mediterranean agroecosystems. Human perturbations have led to a highly diversified landscape, with a mosaic made up of patches of land at different stages of succession, from cultivated fields to closed forest. Our aim here is to characterize ant assemblages and their functional groups in response to these land-use changes. This progressive abandonment results in an initial increase in ant richness and abundance, which can reach high levels if the succession proceeds as far as woodland. In terms of the ant functional groups, this land-use change implies: (1) the appearance of Subordinate Camponotini; (2) an increase in Generalized Myrmicinae, Cryptics and Cold-climate specialists in terms of ant species richness, overall abundance and, for Generalized Myrmicinae and Cryptics, an increase in abundance percentage; (3) a decrease in percentage abundance of Opportunists; (4) a progressive decrease in species richness as well as overall and percentage abundance of Hot-climate Specialists throughout the transformation from crops to woodlands; and (5) an initial increase of Dominant Dolichoderinae followed by a decrease in ant species richness, overall abundance and percentage abundance. Using the ant functional group approach for the clearly separate stages of the regeneration process is a promising method for comparing responses of ant communities to human land use.

Ant behaviour and seed morphology: a missing link of myrmecochory

Seed dispersal by ants (myrmecochory) is mediated by the presence of a lipid-rich appendage (elaiosome) on the seed that induces a variety of ants to collect the diaspores. When seeds mature or fall onto the ground, these ant species transport them to their nest. After eating the elaiosome, the seed is discarded in nest galleries or outside, in the midden or farther away, where seeds can potentially germinate. The final location of seeds with their elaiosomes removed was evaluated to assess the importance of possible handles (structures that ants can grasp to carry) in transporting ants during re-dispersal experiments of seeds from nests of six species of ants. The results indicate that seeds remained within the nest because the ants were not able to transport them out of the nest. As a consequence of the elaiosome being removed, small ant species could not take Euphorbia characias seeds out of their nests. Only large ant species could remove E. characias seeds from their nests. Attaching an artificial handle to E. characias seeds allowed small ant species to redistribute the seeds from their nests. On the other hand, Rhamnus alaternus seeds that have a natural handle after the elaiosome removal were removed from the nests by both groups of ant species. If a seed has an element that acts as a handle, it will eventually get taken out of the nest. The ants’ size and their mandible gap can determine the outcome of the interaction (i.e. the pattern of the final seed shadow) and as a consequence, could influence the events that take place after the dispersal process.

Exclusive frugivory and seed dispersal of Rhamnus alaternus in the bird breeding season

We studied avian frugivory and seed dispersal in a dioecious shrub, Rhamnus alaternus, focusing on the quantitative and qualitative components of effectiveness. The study took place at three locations in the northeast of the Iberian Peninsula, and examined bird behaviour, intensity of feeding, and the consequences for seedling emergence. The coincidence between the bird breeding season and fruit ripening of R. alaternus in the absence of other ripe fruit, generates a monospecific interaction. The extant frugivorous species were mainly legitimate seed dispersers and their abundance was low. Sylvia melanocephala and S. undata were the most important at one site whereas S. atricapilla, Erithacus rubecula and Turdus merula predominated at the other two sites. Fruit handling took place directly on the branches. Bird species used microhabitats differently as first post-feeding perch, which usually was a short distance away. The low density of frugivorous birds in all localities, among others factors, resulted in satiation of the disperser community and many mature fruits unconsumed. Both adults and juveniles feed upon the plants and their foraging patterns are similar. Adults of S. melanocephala were observed to feed fruit to nestlings and consequently a second phase of dispersal potentially arises from the transport of fecal sacs. Pulp removal and passage through the digestive tract increased the probability of seedling emergence. This plant-dispersal interaction has important consequences, both positive and negative for the plant. Positively, the fruiting of R. alaternus at a time when other ripe fruits are not available avoids interspecific competition for seed dispersers. In addition, a low density of seed rain may reduce intraspecific competition. Negatively, the low density and small size of the breeding frugivorous bird community limit fruit handling and removal away from the parent plants, while the territorial behaviour of birds at that time of the year reduces the potential distances of seed dispersal.

Territory characteristics and coexistence with heterospecifics in the Dartford warbler Sylvia undata across a habitat gradient

The study of successional gradients may help to understand the relative influence of habitat structure and competition on territory characteristics. Here, we evaluate the effects of vegetation cover, conspecific and heterospecific densities, and distance to the nearest neighbor on territory size, shape, and overlap in insectivorous birds. We studied these effects along a gradient of postfire habitat regeneration in which foliage cover and densities of focal species varied several-fold. We delineate 197 territories (minimum convex polygons) of the shrub-dwelling Dartford warbler (Sylvia undata) and 255 of the syntopic Sardinian (Sylvia melanocephala), subalpine (Sylvia cantillans) and melodious (Hippolais polyglotta) warblers at three plots in NE Catalonia (Spain and France) in 1987–2005. After accounting for the effect of the number of locations used to delineate polygons, generalized linear mixed models (GLMM) showed a reduction in territory area of the Dartford warbler as conspecific density increased and distance to nearest neighbor decreased, in accordance with the contender pressure hypothesis for territory size regulation. Heterospecific density was not included in the final model of territory size and the effect of habitat structure was marginal. Territory roundness was positively correlated with its size and with conspecific density, probably in relation to energetic constraints, and negatively with heterospecific density. Territorial exclusion was almost complete among Dartford warblers, whereas interspecific territory overlap was extensive and tended to increase with heterospecific density and with structural diversity along the gradient. Our results support the hypothesis that Mediterranean warbler coexistence derives from ecological segregation and not from interspecific territoriality.

Home range and territory of the Sardinian Warbler Sylvia melanocephala in Mediterranean shrubland

Capsule Singing territories were well separated. Aims To examine the spatial distribution of Sardinian Warbler males during the breeding period in Mediterranean shrubland and, specifically, their territories, home ranges and spatial overlaps. Methods We studied a 12-ha plot of Mediterranean shrubland in the 1997 to 1999 breeding seasons. Sardinian Warblers were captured using mist-nets, colour-ringed and their territories mapped. In 1999, seven breeding males were radiotracked in order to map home ranges. Results The Sardinian Warbler had an average of 7.3 breeding pairs/10 ha in the study plot. The mean territory size was 8779 m2 and the mean home range was 22 321 m2. A positive relationship was found between the area of the home range and singing territory. Home ranges of males born in 1998 were about half the size of those of the oldest males. Paired males who moved with a female had smaller home ranges than those that were either unpaired or whose mate was incubating. The degree of home range overlap was high with some overlap between neighbouring territories. The estimate of home range area increased by 10% when the information generated by a mapping method was added and the estimated territory area increased by 31% when data generated by radiotracking were added. Transmitters remained attached to birds for an average of 9.63 ± 3.46 days (mean ± se). Conclusions Singing territories were segregated to a considerable degree. In contrast, the wide overlaps among home ranges was best explained by the presence of food resources that the males exploit at the same time and also by the search for extra-pair copulation in nearby territories. We consider radiotracking in this species to be feasible and valid, with no evidence of negative effects on activity levels, weight or mortality.

Mechanical defence in seeds to avoid predation by a granivorous ant

Harvester ants have traditionally been considered as seed predators that negatively affect plants. In some cases, however, they can also act as positive seed dispersers. During field observations, we noted that a portion of Psoralea bituminosa seeds that were collected and carried to the nest by the granivorous harvester ant Messor barbarus were discarded intact in refuse piles outside the nest. We analyzed and compared the physical characteristics of size, mass and toughness in P. bituminosa seeds from two different origins: intact seeds found in the ant’s refuse piles and seeds collected directly from the plants. Seeds from refuse piles were similar in width but lighter and tougher than seeds from the plant. Our results point to a mechanical defence based on seed toughness to avoid predation by M. barbarus and suggest that an elevated proportion (~69%) of the seeds produced by P. bituminosa could be too tough to be consumed by this ant. These transported but uneaten seeds could benefit by being moved far from the mother plant and this could act as a selective evolutionary pressure towards tough seeds.

Coping with invasive alien species: the Argentine ant and the insectivorous bird assemblage of Mediterranean oak forests

Cork oak forests invaded by the Argentine ant Linepithema humile have a lower abundance and biomass of arthropod prey for birds than uninvaded forests. We studied whether the biomass of breeding insectivorous birds was also lower in invaded areas. We explored this and other possible effects of the ant invasion on the bird community by censusing birds in transects located in four invaded and four uninvaded forest sites in Catalonia (NE Spain) for 3 years. Redundancy analysis showed only slight differences in the community composition between forests. Two insectivorous species, Luscinia megarhynchos and Fringilla coelebs, tended to be less abundant in invaded areas although two others, Phylloscopus bonelli and Sylvia melanocephala, showed the opposite trend. Overall, the differences in prey biomass between invaded and uninvaded areas did not entail a biomass shift in the guild of insectivorous birds, regardless of whether they were shrub or canopy foragers. The main role of the habitat structure in determining bird densities and food resources being non-limiting in the studied forests are two possible non-exclusive explanations for this inconsistency. At today’s levels, the Argentine ant invasion does not appear to have greatly determined the insectivorous bird assemblage of the forests evaluated.

Pre-dispersal seed removal of fleshy fruits and Quercus coccifera (Fagaceae) acorns in a recently burned Mediterranean habitat

The seed fate in early successional habitats can determine plant composition and regeneration capacity after disturbance. Predispersalseed removal has been poorly studied in Mediterranean habitats, especially in burned and logged habitats. We assessed it for two years in pine forests with experiments excluding vertebrates from fleshy fruits (infructescences of Smilax aspera and Rubia peregrina) and acorns (branches of Quercus coccifera). We compared one unburned and one burned area (control). Acorn removal was nil in the burned area while in the unburned habitat seed removal occurred from the beginning of the experiments. It is suggested that the greater vegetation cover in the unburned area shelter rodents from predators and increased their activity. In contrast, reduced cover in the burned area and the occurrence of gnawed acorns on the ground suggest acorn removalby rodents mainly in the post-dispersive stage. Smilax aspera seed removal was slower, and total loss of fruits due to senescence higher,in the burned area. Seed removal appears to be conditioned by interannualvariations related to the activity and density of granivores and frugivores, the availability of fleshy fruits, and the maturation of fruits.In post-fire managed areas the role of granivores and frugivores in the regeneration process should be taken into account.