Summary1. Seed germination characteristics are often modified after seeds are ingested by frugivores. Factors that are intrinsic either to the plant or to the frugivore's digestive tract are responsible for the great variation observed in germination response. 2. Our objectives were to determine whether and how the seed germination patterns of five common western Mediterranean plant species are affected by seed passage through the guts of their major dispersers, and to elucidate the mechanism by which such patterns are changed. 3. We used captive birds ( Turdus merula and Sylvia melanocephala ) to obtain ingested seeds and compared their germination rate (speed) and germinability (final percent germination) with those of controls (uningested, pulp-removed seeds), controlling for seed age, size and source. Germination was monitored for 2 years in an experimental garden. We evaluated the possible changes in seed traits after ingestion by measuring weight and coat thickness, and by observing seed coat sculpture. 4. Rate of seed germination, but not germinability, changed in all species after gut treatment. The greatest effect was in Osyris , in which germination was much enhanced. A great acceleration of germination, which is likely to translate into a seedling size advantage, was also found in Asparagus . In the other three species tested, germination was slower for ingested than for control seeds. 5. For Rubus and Rubia seeds, we found a different germination response depending upon the frugivore species tested. A different degree of seed coat scarification caused by differences in gut retention time, chemical and/or mechanical abrasion probably account for such responses. 6. In three of the species ( Osyris , Rubia and Phillyrea ), seed weight decreased after gut treatment. Such weight loss was not caused by any change in coat thickness, but may have been because of the scarification and consequent alteration of the seed coat structure. 7. The five Mediterranean species studied germinate when rains are most likely to fall (mostly autumn and spring). The different speed of germination promoted by gut treatment within frugivores may increase the probability that seeds can recruit successfully at a given time and in a given place. 8. This study suggests that frugivores contribute to the heterogeneity in germination characteristics not only within plant populations but also within plant communities, each frugivore species having a particular effect on the seeds of each plant consumed.
Summary 1We elucidated the critical life stages and processes in the recruitment of Rhamnus ludovici-salvatoris , a dioecious shrub endemic to the Balearic Islands (western Mediterranean), populations of which have been shrinking in recent decades. We evaluated transition probabilities between all stages involved in regeneration from pollination to seedling recruitment, in order to determine the overall probability of an ovule in a flower becoming an established individual in a population. 2 We compared populations in an oak ( Quercus ilex ) forest and in a more disturbed habitat (an abandoned field). We studied seed rain and probabilities of predation, germination and survival in the main microhabitats within each community. 3 High levels of seed predation after dispersal and unfavourable conditions (water deficit) during germination and recruitment periods were the most critical factors limiting population growth in both habitats. 4 The greater fecundity of shrubs in the abandoned field did not compensate for the lower probability of ovules recruiting new individuals in that habitat compared to the forest. 5 Spatial patterns of recruitment in this species cannot be predicted from seed dispersal patterns produced by frugivorous birds. In both habitats, but especially in the forest, there was strong spatial discordance between seed rain (most seeds were deposited under conspecifics) and seedling establishment. In the forest, most established Rhamnus are found under oaks, whereas in the abandoned field, saplings and juveniles are more equally distributed across different microhabitats. 6 This empirical study supports the idea that habitat disturbance, in general, may considerably alter the patterns of plant recruitment. For endemic species with restricted and rather small populations, the loss of natural habitat is likely to be detrimental for species' persistence, as recruitment in disturbed sites is greatly reduced.
: The introduction of exotic species to an island can have significant effects on the population density and distribution of native species and on the ecological and evolutionary interactions among them (e.g., plant‐animal mutualisms). The disruption of these interactions can be dramatic, significantly reducing the reproductive success of the species and even leading to their extinction. On Menorca Island (Balearic Islands, western Mediterranean), we examined the consequences of the disruption of the mutualism between two endemic species: a perennial shrub, ( Daphne rodriguezii [Texidor]) and a frugivorous lizard (Podarcis lilfordi [Günther]). The lizard became extinct from this island (as well as from Mallorca) as a result of the introduction of carnivorous mammals, which has continued since Roman times. The relict mutualism between D. rodriguezii and the lizard currently persists only in an islet (60 ha) where P. lilfordi is still abundant. We hypothesized that the absence of this lizard from most Menorcan populations is the factor causing the regression of this plant, currently considered at risk of extinction. Through observation and experimentation in the field and laboratory, we found strong evidence that a lack of seed dispersal in Menorca is the main cause of the low seedling recruitment. First, the population with greatest seedling recruitment was that in the islet where lizards were abundant. Second, lizards appeared to be the only dispersers of D. rodriguezii. Lizards consumed large amounts of fruits, without affecting either germination or seedling growth, and moved seeds to sites suitable for plant establishment. Seedlings in Menorca, in contrast, recruited almost exclusively under the parent plants. Third, the effect of other factors that may influence plant population growth (a low fruit set and a high postdispersal seed predation) was similar between the islet and the Menorcan populations. To our knowledge, our results are the first that quantitatively show that a biological invasion can cause a disruption of a specialized plant‐vertebrate mutualism that sets the plant partner on the road to extinction.
Summary 1The fruit-colour polymorphism of Myrtus communis , a common Mediterranean shrub, is examined. We investigate whether frugivores affect the maintenance of the polymorphism, whether morphs differ in germination or seedling growth, and whether passage through frugivores' gut affects seed behaviour and seedling growth. 2 Blue berries are very similar in morphological traits and nutrient composition to the rarer white morph. Rates of fruit removal by birds, the main dispersers of this species, did not differ between morphs, suggesting that a colour preference is not involved in maintenance of the polymorphism. 3 Seeds of the two morphs showed the same germinability (final percentage germination) as well as similar rates of germination under controlled conditions (in growth chamber and greenhouse). Outdoors, seeds from blue berries tended to germinate slightly faster (which might give them an early advantage) but differences between morphs disappeared after several weeks of growth. 4 The germination of myrtle seeds is differently affected by passage through the digestive tract of birds, which appeared to significantly increase germinability, and of the carnivorous pine marten, Martes martes , which did not. The different seed retention time in the gut or the chemical composition of the food ingested along with the seeds may be responsible for such differences. 5 The detection of a positive effect of bird ingestion in outdoor conditions, but not in the growth chamber or in the greenhouse, demonstrates the importance of performing germination tests in the natural habitat. 6 Seed size, irrespective of morph, affects the response to passage through the bird gut. Large seeds, which are more rapidly ejected, and are thus less abraded, germinated at a similar speed to non-ingested controls. Variation in seed size within a single species should thus be considered in future studies.
The recruitment of Rhamnus alaternus appeared to be mainly limited by seed and seedling survival, regardless of the type of habitat in which the species is found.
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