Factors affecting seed rain beneath fleshy-fruited plants

We investigated factors affecting seed rain beneath nine fleshy-fruited fruiting plant species growing in a 1-ha plot of planted Pinus thunbergii in central Japan. We tested whether the numbers of seeds and seed species dropped by birds beneath fruiting plants were correlated with the number of fruits removed by birds from the plants. Most of fruiting plant species with high fruit removal had significantly high seed rain. Both the numbers of seeds and seed species dropped were significantly, positively correlated with the number of fruits removed across for all fruiting plant species. Therefore, fruit removal predicted the difference among heterospecific fruiting plants in seed rain. We also tested whether the number of fruits removed from fruiting plants by birds was related with fruit crop size, fruit size, and height of the plants, and the numbers of fruits and fruit species of neighboring plants near the plants. Most of fruiting plant species with high fruit crop size had significantly high fruit removal. The number of fruits removed was significantly, positively correlated with both the fruit crop size and the number of neighboring fruits across the nine fruiting plant species. However, the effect of the neighboring fruit density on fruit removal was lower remarkably than that of fruit crop size. Therefore, fruit crop size best predicted the differences among heterospecific fruiting plants in fruit removal. We suggest that fruiting plant species with high fruit crop size and high fruit removal contribute to intensive seed rain beneath them. This revised version was published online in June 2006 with corrections to the Cover Date.     

Crop Size and Fruit Neighborhood Effects on Bird Visitation to Fruiting Schefflera morototoni Trees in Puerto Rico1

Studies of zoochorous seed dispersal systems often consider crop size, yet seldom consider the kinds and amounts of fruits surrounding parent plants (the fruit neighborhood) when attempting to explain among‐plant variation in fruit removal. We studied avian frugivory at 24 Schefflera morototoni trees from February to May 1998 in central Puerto Rico. The number of fruits removed by avian seed dispersers per visit was similar among focal trees (typically 2–4). In contrast, visitation rate was highly variable (range: 0–71 visits per 4 h). We used multiple regression analyses to evaluate the relative roles of crop size (focal tree ripe fruit abundance) and fruit neighborhood variables (measured within 30 m of focal trees) in affecting visitation to focal trees by avian frugivores. Visitation rate was positively related to crop size (although this variable was only significant in one of four regression models considered) and negatively related to the presence or abundance of conspecific fruits, suggesting that trees competed intraspecifically for dispersers. Relationships between visitation and heterospecific fruits were mixed—some kinds of fruits appeared to enhance visitation to focal trees, while others seemed to reduce visitation. In most regression models, neighborhood variables had larger effects on visitation than focal tree fruit crop size. Our results highlight the important effects of local fruiting environments on the ability of individual plants to attract seed dispersers.     

Fruit production in Sorbus aucuparia L. (Rosaceae) and pre-dispersal seed predation by the apple fruit moth (Argyresthia conjugella Zell.)

Variation in fruit production and pre-dispersal seed predation by Argyresthia conjugella was studied in␣four populations of Sorbus aucuparia in northern Sweden.␣The number of infructescences, fruits per infructescence, consumed seeds and developed unattacked seeds per fruit were scored in marked trees from 1984 to 1990. The results showed that the number of fruits produced in each population determined the number of seed predators occurring in the host population, as the yearly number of seed predators was significantly and positively correlated with yearly number of fruits, in all but one population. The seed predators showed a delay in response to variation in number of fruits produced. This lag in response resulted in a large proportion of fruits being attacked and seeds consumed in a bad fruiting year that followed a good fruiting year, and vice versa. The proportion of fruits attacked and seeds consumed was largest in the population showing the greatest between-year variation in fruit production and lowest in the population showing the lowest between-year variation in fruit production. Furthermore, the individuals within the former population were synchronised, while they were not in the latter population. These results contradict one of the possible explanations of mast-seeding, where large synchronised between-year variation is supposed to reduce pre-dispersal seed predation. Instead, differences in attraction of the seed predator to differences in fruit crop size could explain the observed difference in seed predation between the two populations with opposite fruiting patterns. Within each population, irrespective of year, the proportion of fruits attacked and seeds consumed was independent of a tree's fruiting display. Therefore, trees with high fruit production, despite harbouring the largest number of seed predators, produced the largest number of developed seeds in absolute numbers, compared to trees that produced few fruits. Received: 25 February 1996 / Accepted: 30 November 1996     

Fruit removal of Coprosma quadrifida (Rubiaceae) by birds in south-eastern Australia

Abstract Removal of the fleshy fruits of Coprosma quadrifida by birds was examined in relation to fruit crop size, neighbouring fruit crop size and time in the season in montane wet sclerophyll forest in south-eastern Australia. On average, 84% of individual fruit crops were removed. Frugivores removed fruit in proportion to its abundance, indicating that they continually assess availability rather than respond to a critical abundance. The size of neighbouring fruit crops did not influence fruit removal rates, suggesting that there is little competition among neighbours for dispersers. The ripe fruits of C. quadrifida were eaten by 14 bird species, comprising 50% of the bird species trapped. Silvereyes (Zosterops lateralis) were the most abundant consumers of fruits. With the exception of the parrot, Platycercus elegans, all birds defecated seeds intact and had no influence on seed viability.     

Latitudinal and elevational variation in fruiting phenology among western European bird-dispersed plants

I try to test the prediction that bird-dispersed plants should produce fruits when fruit-eating birds are most abundant by reviewing some phenological data of fleshy fruit production in western Europe The prediction that fruit ripening dates in populations of the same species should occur later at lower latitudes and elevations, to coincide with the maximum abundance of fruit-eating birds, is not supported by the data The patterns of seasonal variation in the total number and biomass of fruits, but not in the proportion of species in fruit, in communities at different latitudes and elevations do coincide with patterns of seasonal abundances of avian frugivores 1 suggest that this coincidence is due to the greater relative abundance (and contribution to total fruit production) in each locality of species that fruit at times of the year when birds are most abundant These species may have achieved a demographic advantage by getting more seeds dispersed than species that ripen fruits in other seasons     

Fruit removal efficiency and success: influence of crop size in a neotropical treelet

The reproductive success of animal-dispersed plants is closely linked to the number of seeds that they are able to disperse. The fruit crop size hypothesis states that a plant with large fruit crop size will attract more dispersers than a plant with a smaller fruit crop, which may result in more seeds being dispersed from the foremost. In this study, we experimentally examined the effect of crop size and other factors on primary seed dispersal in a neotropical shrub/tree, Casearia corymbosa (Flacourtiaceae). We used two predictive variables of reproductive success, which produce an accurate picture of seed dispersal ratio: fruit removal efficiency (proportion of a fruit crop removed by frugivores) and fruit removal success (relative contribution of each individual tree to the number of fruits removed in the population). We established two levels of fruit crop size at the C. corymbosa individuals, using plants with large (150 fruits) and small crops (50 fruits). We found that individual plants with larger crops had significantly higher values of fruit removal efficiency (92.6%) and success (5%) than plants with smaller crops (69.3% and 1.3%, respectively). Fruit removal efficiency was related to vegetation type, plant height and fruit width, but the variance explained by these variables was low ( < 8%). Fruit removal success was significantly related to crop size ( > 90% of the variance explained). These results suggest that fruit removal efficiency and success are strongly related to fruit crop size of C. corymbosa plants.     

Fruit size in a tropical tree species: variation,preference by birds,and heritability

Ocotea tenera (Lauraceae), an understory bird-dispersed tree, produces single-seeded fruits that vary in diameter from 1.4 to 2.4 cm. Much of the variation within a population at Monteverde, Costa Rica occurred within individual trees. The relative size of fruits produced by different trees remained generally constant over an 11-year period despite slight differences between years in the average size of fruits produced by a given tree.Fruit-eating birds could thus express their preferences for particular fruit size characteristics by choosing among trees that have distinct distributions of fruit diameters, and between individually variable fruits within trees. In a field study of individually marked fruits, birds removed 46.2% of fruits; the rest of the fruits were destroyed by invertebrate (25.3%) and vertebrate (4.3%) pulp-feeders or aborted by the plant after remaining ripe but uneaten for as long as 100 days (24.2%). The four major avian seed dispersers of O. tenera each have gape widths exceeding all but the largest fruits. Birds preferred plants with greater-than-average-sized fruits; within trees, they favored larger fruits, apparently because net pulp mass increases with fruit diameter. Fruits that ripened early in the season were more likely to be removed and were removed more quickly than late-ripening fruits.Based on mother-offspring regressions of mean fruit size, the phenotypic variation in fruit diameter in O. tenera is highly heritable, indicating the potential for an evolutionary response to selection by birds. Nonetheless, directional selection on fruit size or shape is likely to be inconsistent, constrained by genetic correlations, and weak compared to selection on traits like fecundity or phenology.     

Determinants of fruit removal in <Emphasis Type="Italic">Geonoma pauciflora</Emphasis>, an understory palm of neotropical forests

Competition for seed dispersers is supposedly a selective force that drives the evolution of plant reproductive traits. In the understory of tropical forests, such competition should be especially severe among bird-dispersed plant species because (i) the production of copious fruit crops is limited by low light availability; (ii) there usually is a high density of fleshy fruited plants, and (iii) understory frugivorous birds are not abundant. In this paper, we took advantage of a high-density population of Geonoma pauciflora, a bird-dispersed palm species growing in the understory of the Brazilian Atlantic forest, to investigate the influence of plant traits and its immediate neighborhood on fruit removal. Intrinsic (crop and fruit sizes) and extrinsic traits (related to light availability) affected the fruit removal of G. pauciflora. Crop size had a greater influence than fruit size on the absolute number of fruits removed, whereas none of the investigated traits influenced decisively fruit removal efficiency (i.e., the proportion of an individual’s crop removed). The influence of light availability was mostly indirect, through its positive influence on fruit production. A significant positive spatial autocorrelation in removal efficiency occurred among neighboring plants within a 7-m radius, which is indicative of facilitation among neighboring individuals. The consequence of such positive spatial autocorrelation in removal efficiency for clonal plants such as G. pauciflora is that by attracting a frugivore a given ramet may promote the removal of fruits of other ramets, thus enhancing the reproductive output of the genet as a whole.     

Fruit characteristics and factors affecting fruit removal in a Panamanian community of strangler figs

We describe fruiting characteristics for 12 species in a community of strangler figs (Moraceae: Urostigma) studied in Panama. We quantify diurnal and nocturnal removal rates and proportions of fruits removed, and relate them to the activities of the main dispersers of the figs: bats and birds. These results combined with previous studies show that there are clear differences between fig species with fruit that ripen red and those with fruit that remain green(ish). In the red-fruited species, the fruit are small, ripen asynchronously over relatively long periods, produce little scent, and are mainly taken during the day by birds. In contrast, in the green(ish)-fruited species, the fruits are larger, span a range of sizes, ripen relatively synchronously, produce very distinctive aromas, and are mainly taken at night by bats. This dichotomy in fruiting characteristics suggests coadaptive links between groups of dispersers and different species within the genus Ficus. All fig species produce a range of fruit crop sizes (10–155 fuits/m² canopy area) of which a high proportion were removed by seed dispersers (>80%). Removal rates (fruit removed per day) were positively correlated with crop size, suggesting that trees with large crop size attract more frugivores. Removal rates of green-fruited figs were significantly lower and persistence and abortion of ripe fruit were significant higher around full moon, apparently due to the reduced activity of bats. We further estimate the number of bats that are sustained by a tree fruit crop and account for the observed fruit removal. We then discuss the evidence for coadaptation between different groups of figs and their seed dispersers, Finally, we consider the conservation implications for figs as keystone resources in tropical forests. Received: 26 April 1999 / Accepted: 10 January 2000     

Why do some,but not all,tropical birds migrate? A comparative study of diet breadth and fruit preference

Annual migrations of birds profoundly influence terrestrial communities. However, few empirical studies examine why birds migrate, in part due to the difficulty of testing causal hypotheses in long-distance migration systems. Short-distance altitudinal migrations provide relatively tractable systems in which to test explanations for migration. Many past studies explain tropical altitudinal migration as a response to spatial and temporal variation in fruit availability. Yet this hypothesis fails to explain why some coexisting, closely-related frugivorous birds remain resident year-round. We take a mechanistic approach by proposing and evaluating two hypotheses (one based on competitive exclusion and the other based on differences in dietary specialization) to explain why some, but not all, tropical frugivores migrate. We tested predictions of these hypotheses by comparing diets, fruit preferences, and the relationships between diet and preference in closely-related pairs of migrant and resident species. Fecal samples and experimental choice trials revealed that sympatric migrants and residents differed in both their diets and fruit preferences. Migrants consumed a greater diversity of fruits and fewer arthropods than did their resident counterparts. Migrants also tended to have slightly stronger fruit preferences than residents. Most critically, diets of migrants more closely matched their preferences than did the diets of residents. These results suggest that migrants may be competitively superior foragers for fruit compared to residents (rather than vice versa), implying that current competitive interactions are unlikely to explain variation in migratory behavior among coexisting frugivores. We found some support for the dietary specialization hypothesis, propose refinements to the mechanism underlying this hypothesis, and discuss how dietary specialization might ultimately reflect past interspecific competition. We recommend that future studies quantify variation in nutritional content of tropical fruits, and determine whether frugivory is a consequence or a cause of migratory behaviour.     

Crop size,plant aggregation,and microhabitat type affect fruit removal by birds from individual melastome plants in the Upper Amazon

We studied the efficiency (proportion of the crop removed) and quantitative effectiveness (number of fruits removed) of dispersal of Miconia fosteri and M. serrulata (Melastomataceae) seeds by birds in lowland tropical wet forest of Ecuador. Specifically, we examined variation in fruit removal in order to reveal the spatial scale at which crop size influences seed dispersal outcome of individual plants, and to evaluate how the effect of crop size on plant dispersal success may be affected by conspecific fruit abundance and by the spatial distribution of frugivore abundance. We established two 9-ha plots in undisturbed terra-firme understory, where six manakin species (Pipridae) disperse most seeds of these two plant species. Mean levels of fruit removal were low for both species, with high variability among plants. In general, plants with larger crop sizes experienced greater efficiency and effectiveness of fruit removal than plants with smaller crops. Fruit removal, however, was also influenced by microhabitat, such as local topography and local neighborhood. Fruit-rich and disperser-rich patches overlapped spatially for M. fosteri but not M. serrulata, nonetheless fruit removal of M. serrulata was still much greater in fruit-rich patches. Fruit removal from individual plants did not decrease in patches with many fruiting conspecifics and, in fact, removal effectiveness was enhanced for M. fosteri with small crop sizes when such plants were in patches with more conspecifics. These results suggest that benefits of attracting dispersers to a patch balanced or outweighed the costs of competition for dispersers. Spatial pattern of fruit removal, a measure of plant fitness, depended on a complex interaction among plant traits, spatial patterns of plant distribution, and disperser behavior.     

Fruit production,migrant bird visitation,and seed dispersal of Guarea glabra in Panama

Summary The relationship between bird visitation and the size of the available fruit crop was studied at an understory tree (Guarea glabra Vahl, Meliaceae) in the tropical wet forest of Barro Colorado Island, Panama Canal Zone. Twelve resident species and seven North American migrant species fed on the Bright orange arilloids (seeds with arils), which were not depleted during the normal fruiting period. The number of individual visitors, the number of visiting species, and the number of seeds removed increased linearly with the size of the available fruit crop. The proportion of seeds removed did not increase with the size of the available fruit crop, indicating that dispersal is a function of the number of fruit available and not a disproportionate function of large fruit displays. Four species of North American migrants (Myiarchus crinitus, Catharus ustulatus, Vireo olivaceus, and Vermivora peregrina) accounted for 70% of the visits and 60% of the seeds removed from the trees. No resident species visited Guarea as frequently as any one of these migrants. There was no indication that any single visitor was dependent on this tree for nutrition, nor that the tree was dependent upon any single species for dispersal. We hypothesize that the fruiting season of G. glabra is adaptively synchronized with northward migration of opportunistically frugivorous North American birds.     

Population and individual level of masting in a fleshy-fruited tree

Masting is usually considered as a population phenomenon but it results from individuals?? reproductive patterns. Studies of individual patterns of seed production and their synchrony are essential to an understanding of the mechanisms of masting. The aim of this study was to find the relationship between population and individual levels of masting. We examined individuals?? contribution to masting, considering their endogenous cycles, interannual variability and associated weather cues, as well as inter-individual synchrony of fruit production. We studied masting of Sorbus aucuparia L., which in Europe is one of the most common trees bearing fleshy fruits and is strongly affected by a specialized seed predator. The data are 11-year measurements of fruit production of 250 individuals distributed on a 27-ha area of subalpine forest in the Western Carpathians (Poland). Population- and individual-level interannual variability of fruit production was moderate. Synchrony among individuals was relatively high for all years, but the trees were much less synchronized in heavy crop years than in years of low fruit production. Weak synchrony among trees for heavy production years suggests that the predator satiation hypothesis does not explain the observed masting behavior. Fruit production, both at individual and at population level, was highly correlated with weather conditions. However, the presence of masting cannot be fully explained by the resource-matching hypothesis either. We suggest that adverse weather conditions effectively limit fruit production, causing high inter-individual synchrony in low crop years, whereas the unsynchronized heavy crop years seem to have been affected by individually available resources.     

The role of termites in the decomposition of wood and leaf litter in the Southern Guinea savanna of Nigeria

Summary The number of flowers produced by inflorescences of Yucca whipplei (Agavaceae) consistently exceeds the number of fruits produced by about one order of magnitude. To determine the factors responsible for low fruit set, the relation between pollinator availability, the amount of resources spent on reproduction (as indicated by inflorescence size), and the number of fruits matured was studied during 1978 and 1979 at 18 locations in chaparral, coastal sage scrub, and desert scrub communities of southern California.The following results support the conclusion that pollinators do not usually limit fruit production in Yucca whipplei. Rather, fruit production is limited by the amount of resources available to support developing fruits. (1) Fruit production is positively correlated with inflorescence size both within and between populations. The average size of inflorescence for a population is an excellent predictor of mean fruit production. Furthermore, 54% of the total variance in fruit production of individual plants can be explained by inflorescence size. (2) In contrast, although fruit production within most populations is positively correlated with an index of the number of pollinator visits to an inflorescence, the relative abundance of pollinators for a population is a poor predictor of mean fruit production, and only 9% of the total variance in fruit production can be explained by the visitation index. Furthermore, at four sites studied for two years, there was little change in average inflorescence size or fruit production from 1978 to 1979, despite large differences in relative abundance of pollinators at each of the sites. (3) Based on geographic proximity, and physiographic and vegetational similarities, study sites were grouped into regional clusters. Both inflorescence size and fruit production varied considerably between regions. Of the total variation in fruit production, 27% can be attributed to differences between regions. Most of this variation is the result of regional differences in inflorescence size, which in turn influence fruit production.Why does Yucca whipplei produce such large inflorescences if so few fruits can be supported? Two relevant hypotheses are discussed: (1) the floral display is the result of selection for pollen dissemination at the expense of fruit set; and (2) the floral display is the result of selection for a bet-hedging strategy either to increase the probability of adequate pollination when pollinators are unusually rare, or to allow individuals to support more fruits when resources are unusually abundant.     

Signal convergence in fruits: a result of selection by frugivores?

The Dispersal Syndrome hypothesis remains contentious, stating that apparently nonrandom associations of fruit characteristics result from selection by seed dispersers. We examine a key assumption under this hypothesis, i.e. that fruit traits can be used as reliable signals by frugivores. We first test this assumption by looking at whether fruit colour allows birds and primates to distinguish between fruits commonly dispersed by birds or primates. Second, we test whether the colours of fruits dispersed by primates are more contrasting to primates than the colours of bird‐dispersed fruits, expected if fruit colour is an adaptation to facilitate the detection by seed dispersers. Third, we test whether fruit colour has converged in unrelated plant species dispersed by similar frugivores. We use vision models based on peak sensitivities of birds’ and primates’ cone cells. We base our analyses on the visual systems of two types of birds (violet and ultraviolet based) and three types of primates (trichromatic primates from the Old and the New Worlds, and a dichromatic New World monkey). Using a Discriminant Function Analysis, we find that all frugivore groups can reliably discriminate between bird‐ and primate‐dispersed fruits. Fruit colour can be a reliable signal to different seed dispersers. However, the colours of primate‐dispersed fruits are less contrasting to primates than those of bird‐dispersed fruits. Fruit colour convergence in unrelated plants is independent of phylogeny and can be better explained by disperser type, which supports the hypothesis that frugivores are important in fruit evolution. We discuss adaptive and nonadaptive hypotheses that can potentially explain the pattern we found.     

Reproductive ecology of Thymelaea velutina (Thymelaeaceae)- Factors contributing to the maintenance of heterocarpy

Thymelaea velutina (Thymelaeaceae) is a dioecious shrub that presents a unique type of heterocarpy which consists of the simultaneous production of dry and fleshy fruits. It is endemic to the Balearic Islands (Western Mediterranean) and is found both in dunes and mountain areas. The goal of this study was to identify which factors influence the production of both fruit types, examining the variation of their effects at a spatio-temporal scale (comparing two localities in different years). Specifically, we investigated (1) whether pollen limitation influences the type of fruit produced, and (2) the possible differences in seed size, mass, dispersal capacity, seed predation, germination patterns and seedling survival between fruit types. We also examined if the production of fleshy fruits was modified with the application of gibberellins to reproductive branches. Although fleshy fruits were consistently more abundant than dry ones at both populations, their proportion was significantly higher at the site with greater precipitation. The addition of either pollen or gibberellins did not affect the proportion of each fruit type. Seeds in fleshy fruits are consistently larger, heavier and more likely to be dispersed than seeds in dry fruits, but germinability, germination rate and seedling survival was similar among fruit types. Heterocarpy in this species is currently maintained as there is no apparent factor that exerts any strong selective pressure on either fruit type. The two fruit types might even have different `functions', one serving especially for dispersal and population expansion and the other for producing a seed bank that ensures an eventual germination.     

Are large frugivorous birds better seed dispersers than medium‐ and small‐sized ones? Effect of body mass on seed dispersal effectiveness

Frugivorous birds vary in seed dispersal effectiveness (SDE) depending on their body mass. It has been suggested that large birds are more effective dispersers than small ones because they consume a large number of fruits, disperse seeds of distinct sizes, and transport seeds over long distances. Yet, few studies have evaluated the impact of body mass on SDE of birds. In this study, we compiled one database for the quantity (i.e., frequency of visits to plants and number of seeds removed per visit) and quality (i.e., germination of seeds after gut passage and gut retention time of seeds) of seed dispersal by frugivorous birds to evaluate the impact of body mass on SDE. In addition, we compiled data on plant characteristics such as life‐form, fruit type, number of seeds per fruit, and size of seed to evaluate their influence on the quantity and quality of seed dispersal. Data were analyzed with linear mixed effects models and quantile regressions to evaluate the relationship between body mass of birds and quantity, quality, and SDE, in addition to the influence of plant characteristics on SDE. The body mass of birds was negatively related to the frequency of visits to plants. Furthermore, it was positively related to the number of seeds removed per visit, although negatively related to seed size. The life‐form of plants was the only factor explaining the germination of seeds after gut passage. Yet, the body mass of birds was positively related to the gut retention time of seeds. Small and medium birds have a relatively higher SDE than large birds. These results differ from the assertion that large birds are more effective dispersers of plants. Small and medium birds are also effective dispersers of plants that should be preserved and protected from the impact of human activities.     

Influence of between-year variation in the density of Rhus <Emphasis Type="Italic">trichocarpa</Emphasis> fruits on the removal of fruit by birds

The fruiting phenology and fruit removal patterns of Rhus trichocarpa Miq. (Anacardiaceae) were investigated in a warm-temperate secondary forest in Japan. Mature fruits of this species are dispersed by birds. Effects of fruit display size and canopy openness on fruit removal were investigated in years with different fruit densities (i.e., masting and non-masting years). Moreover, effects of increased canopy openness during winter on fruit removal were also investigated. Seasonal patterns of fruit removal were quite different between masting and non- masting years. In the non-mast year, fruits were removed by birds soon after maturation in the summer. In contrast, in the mast year, fruits were removed gradually by birds from summer to winter. Moreover, the rate of fruit removal was greater for trees with a larger display size in the non-mast year, whereas that was greater for trees with greater canopy openness in the mast year. Canopy openness increased in winter, and fruit removal in winter was enhanced in trees with a more open canopy only in the non-mast year. These observations strongly suggest that avian dispersers became satiated in the mast year, whereas fruit removal was enhanced in the non-mast year. In the mast year, although many fruits were not dispersed until winter, they were neither depredated nor rotten, and the long period of fruit removal by birds may have enhanced fruit dispersal. The large abundance of seedlings and saplings of this species in Japanese secondary forests suggests that this fruiting strategy is beneficial.     

The effect of unripe fruits on ripe fruit removal by birds in Pistacia terebinthus: flag or handicap?

The shrub Pistacia terebinthus produces crowded infructescences with up to several hundred fruits, which are bright red when unripe and turn green when ripe. Most fruits contain an empty seed and never reach maturity. More ripe fruits were removed by birds from experimental bicolored fruit displays (consisting of infructescences with ten ripe fruits and stripped of unripe fruits, paired with infructescences with only unripe fruits) than from monocolored ones (single infructescences with ten ripe fruits and stripped of unripe fruits). Thus, the presence of unripe fruits seems to increase the conspicuousness or attractiveness of fruit displays to fruit-eating birds. A second experiment compared ripe fruit removal from experimental infructescences having only ripe fruits, with that from control infructescences containing both ripe fruits and natural numbers of unripe fruits, all on P. terebinthus plants. Unlike the first experiment, each bicolored display in this case consisted of a single infructescence with both unripe and ripe fruits. A higher proportion of ripe fruits was removed by birds from infructescences free of unripe fruits. This result suggests that the presence of unripe fruits reduces the accessibility of ripe fruits for fruit-eating birds. This is further supported by field observations of bird foraging behavior.     

The fruits of selectivity: how birds forage on Goupia glabra fruits of different ripeness

Although many studies have been published on avian fruit selection, few have addressed the effects of fruit scarcity on the patterns of fruit choice. Here, we compared the consumption of seven bird species for six simultaneously present maturation stages of Goupia glabra fruits. Ripe G. glabra fruits contain more lipids, carbohydrates and energy, and fewer phenols, than unripe fruits. All bird species selected from among ripening stages and removed a higher proportion of ripe fruits than of intermediate or unripe fruits. Importantly, however, fruit choice was flexible in all species. Whether birds preferred or avoided fruits of intermediate ripeness depended on the overall fruit supply. When ripe fruits were scarce, birds showed a higher acceptance of fruits of intermediate ripeness, but still rejected the least ripe fruit stages. In a foraging bout, most birds fed on fruits of the same ripeness. By doing so, birds maximised instantaneous energy gain per time, because search time was longer for riper fruits while energy intake was lower for less ripe fruits. The results suggest that birds select fruits based on fine-scale differences in profitability, but accept less profitable fruits during low fruit abundance. If environmental factors such as overall fruit availability influence avian fruit choice, we suggest that the potential for directional selective pressures on fruit compounds is restricted.