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     

Plasticity and diversity of the phenology of dioecious Ficus species in Taiwan

While Ficus present a series of traits often associated with dioecy, the prevalence of dioecy in Ficus is atypical. In Asian floras, dioecious Ficus species generally outnumber monoecious ones. Further this is also true in relatively northerly locations for Ficus such as the island of Taiwan. Ficus are pollinated by species-specific wasps that use fig flowers as breeding sites. In dioecious fig species, pollinators develop only in the inflorescences of male fig trees. In this study, we investigated the reproductive phenology of four dioecious Ficus species with distinct ecologies in several locations in northern and southern Taiwan. The two first species (Ficus erecta and Ficus septica) were investigated in four locations. Reproductive phenology was quite different among sites, even within a single species. For example, F. erecta presented well-defined crops at the population level in its usual high-elevation habitat but continuous fig production at low elevations, especially in South Taiwan. The two other fig species (Ficus pedunculosa var. mearnsii and Ficus tinctoria subsp. swinhoei), are shrubs growing together along seashores in exposed locations on coral reef remnants. These two species presented quite different traits allowing the survival of pollinating wasp populations. Ficus pedunculosa var. mearnsii produced figs continuously so that fresh receptive figs were always available for the pollinating wasps while F. tinctoria subsp. swinhoei extended the period of receptivity of its figs, so that receptive figs that had been waiting for pollinating wasps were almost always available. In summary, dioecious figs in Taiwan showed remarkable variation in their phenology, within species among locations or among species within location. Nevertheless, despite this variation, the phenology of the trees always allowed survival of pollinating wasp populations. Dioecious figs seem to have adopted a differentiated set of strategies which result in high resilience of pollinator populations. This resilience could help explain the atypical prevalence of dioecy in Ficus.     

The figs of winter: Seasonal importance of fruiting fig trees (Ficus: Moraceae) for urban birds

Birds and figs are conspicuous members of the tropical and subtropical ecosystems. Because they are easily observed and very speciose, their relationships have been well studied in many areas, and the figs are considered a keystone resource for many bird species which are efficient fig seed dispersers. Taiwan has a relatively high endemism rate for many taxa (17% of bird species) but because of its high human population density, most lowland habitats are heavily developed, of which much of it covered by dense urban habitation. To establish the importance of urban figs for birds, we focused our surveys mostly on three common urban fig species (Ficus caulocarpa, F. microcarpa and F. subpisocarpa). We observed trees with ripening figs from July 2013 to December 2016 in order to determine the composition of the fig-consuming bird community. In addition, we added all the information available in the scientific literature and birdwatchers' observations which we could find. In total, we observed 42 bird species consuming 18 fig species. The bird diversity in urban areas was non-negligible even during winter. Therefore, there are two reasons why figs are important for Taiwan's bird avifauna: in cities, the tree diversity is generally low so that figs provide a stable food resource; and since figs are fruiting all year-round, they are one of the few reliable resources available during winter when many migrant birds overwinter in Taiwan. Already crucial for many species in tropical and subtropical forests, fig trees may also be essential for urban birds in tropical and subtropical regions.     

Phenology of figs in Budongo Forest Uganda and its importance for the chimpanzee diet

This paper reports on the phenological patterns of figs in Budongo Forest, Uganda, and how it relates to chimpanzee food availability in different seasons. In addition, we analysed the dung of chimpanzees to understand the composition of fruits in their diet. The aim of our study was to assess Ficus phenology and how it affects chimpanzee diet. Fifteen species of figs were monitored for fruit (syconium) and leaf phenology between June 2000 and 2001. Ficus fruit production varied significantly between and within species, and also with tree trunk and crown diameters. Fig fruit production was asynchronous and individual fig trees produced crops from one to five times in a year. In addition to fruits, chimpanzees fed on young leaves of some Ficus species. Shedding of old Ficus leaves coincided with the dry season, followed by appearance of young leaves. The dry season in Budongo is a period of general fruit scarcity. The combination of fig fruits and young leaves make up the most important food in the diet of chimpanzees. From the chimpanzee dung, more than 78% of seeds comprised fig ‘seeds’ (nutlets) and the rest of the diaspores were from other tree species. Our findings suggest that chimpanzees disperse large number of diaspores in their dung, thereby serving as important agents of natural forest regeneration.     

Phenological Adaptations in Ficus tikoua Exhibit Convergence with Unrelated Extra-Tropical Fig Trees

Flowering phenology is central to the ecology and evolution of most flowering plants. In highly-specific nursery pollination systems, such as that involving fig trees (Ficus species) and fig wasps (Agaonidae), any mismatch in timing has serious consequences because the plants must balance seed production with maintenance of their pollinator populations. Most fig trees are found in tropical or subtropical habitats, but the dioecious Chinese Ficus tikoua has a more northerly distribution. We monitored how its fruiting phenology has adapted in response to a highly seasonal environment. Male trees (where fig wasps reproduce) had one to three crops annually, whereas many seed-producing female trees produced only one fig crop. The timing of release of Ceratosolen fig wasps from male figs in late May and June was synchronized with the presence of receptive figs on female trees, at a time when there were few receptive figs on male trees, thereby ensuring seed set while allowing remnant pollinator populations to persist. F. tikoua phenology has converged with those of other (unrelated) northern Ficus species, but there are differences. Unlike F. carica in Europe, all F. tikoua male figs contain male flowers, and unlike F. pumila in China, but like F. carica, it is the second annual generation of adult wasps that pollinate female figs. The phenologies of all three temperate fig trees generate annual bottlenecks in the size of pollinator populations and for female F. tikoua also a shortage of fig wasps that results in many figs failing to be pollinated.     

The community structure of bird assemblages on urban strangler figs

Figs have been regarded as keystone plant resources that support diverse tropical vertebrate frugivore communities. Planting or conserving large fig trees, such as stranglers, have therefore been proposed for enhancing urban biodiversity. We compared the diversity and community structure of bird assemblages on strangler figs with non‐fig urban trees as well as between the fruiting and non‐fruiting fig trees in an urban setting in Singapore. The total bird abundance across all the fig trees when in fruit was 4.5‐fold higher than on non‐fig trees and 3.5‐fold higher than when the same fig trees were not fruiting, but only attracted two more species. On individual trees, after accounting for the presence of mistletoes, tree height, the area covered by buildings, road lane density, and the distance to natural vegetation, mean diversity was not different between non‐fig trees and fig trees when they were not in fruit. On the other hand, when fruiting, each fig tree on average had 1.4 more species, 3 more counts of non‐native birds, and 1.6 more counts of insectivorous birds than when not fruiting. There was significant compositional turnover between non‐fig trees and non‐fruiting fig trees, while community dispersion was significantly lower among fig trees in fruit. Our results demonstrate that fig trees provide fruit and non‐fruit resources for birds in an urban landscape but do not necessarily support more diverse total bird assemblages than non‐fig trees. Instead, bird communities on fruiting urban figs would be highly homogeneous and dominated by a few species. Abstract in Malay is available with online material.     

A comparison of pollinator fig wasp development in figs of Ficus montana and its hybrids with Ficus asperifolia

Figs (Moraceae) and pollinator fig wasps (Hymenoptera: Agaonidae) have a highly specific mutualistic relationship but fig wasps occasionally enter atypical hosts, and this can lead to hybrid fig trees and the potential for gene flow between species. Many fig trees are dioecious, with fig wasp offspring developing in galled ovules inside figs on male trees, whereas seeds develop only in figs on female trees. We generated experimental hybrids between the Asian Ficus montana Blume and a closely related African species Ficus asperifolia Miquel. Male F1s were sterile if entered by Kradibia tentacularis (Grandi) (Agaonidae), the pollinator of F. montana, because its offspring always failed to develop, without ovule enlargement. As with the F1s, figs on most male backcross plants [F. montana × (F. montana × F. asperifolia)] also aborted shortly after pollinator entry, resulting in a higher turnover of figs than with F. montana, although the times taken for the figs to reach receptivity were similar. Pollinator larvae nonetheless consistently managed to develop inside the figs of one backcross plant and also occasionally in a few figs from another backcross individual. In these figs, galled ovules developed as normal, whereas in figs that aborted the galled ovules failed to enlarge. The sex ratio of K. tentacularis progeny in the backcross figs was female biased and did not differ from that in F. montana figs. Sycoscapter spec. (Hymenoptera: Pteromalidae), a parasitoid of K. tentacularis, was able to lay eggs and developed normally inside male backcross figs where its host was present.     

Effects of ingestion by neotropical bats on germination parameters of native free-standing and strangler figs (Ficus sp., Moraceae)

Fruit-eating animals can influence the germination success of seeds through transportation and handling. We experimentally tested the contribution of ingestion by the common fruit-eating bat, Artibeus jamaicensis (Phyllostomidae, Chiroptera), to the percentage and rate of seed germination of figs (Ficus, Moraceae), which are considered keystone species for many frugivores. We collected fruits from three species of native free-standing figs (subgenus Pharmacosycea: F. insipida, F. maxima and F. yoponensis) and three species of native strangler figs (subgenus Urostigma: F. nymphiifolia, F. obtusifolia and F. popenoei) on Barro Colorado Island, Panama. The germination success of seeds removed from fruit pulp either manually or by ingestion was very high (>92%), while seeds that were not removed from fruit pulp were destroyed by fast-growing fungi within a few days. The dynamics of seed germination were not influenced by ingestion, but differed between the two subgenera of figs. In free-standing figs, germination started significantly earlier (5.3 ± 0.7 days) than in strangler figs (8.6 ± 1.4 days). Furthermore, strangler seeds were covered with a sticky coating and their seedlings developed cotyledons faster than fine roots, in contrast to free-standing figs that showed the opposite pattern. Our results demonstrate that the germination of fig seeds is positively influenced by passage through the gut of A. jamaicensis. Furthermore, free-standing and strangler figs revealed differences in germination parameters that might be adaptive with respect to the suitability of microsites such as tree fall gaps or host trees for establishment.     

Geographic and taxonomic distribution of a positive interaction: ant-tended homopterans indirectly benefit figs across southern Africa

Although species pairs and assemblages often occur across geographic regions, ecologists know very little about the outcome of their interactions on such large spatial scales. Here, we assess the geographic distribution and taxonomic diversity of a positive interaction involving ant-tended homopterans and fig trees in the genus Ficus. Previous experimental studies at a few locations in South Africa indicated that Ficus sur indirectly benefited from the presence of a homopteran (Hilda patruelis) because it attracted ants (primarily Pheidole megacephala) that reduced the effects of both pre-dispersal ovule gallers and parasitoids of pollinating wasps. Based on this work, we evaluated three conditions that must be met in order to support the hypothesis that this indirect interaction involves many fig species and occurs throughout much of southern Africa and Madagascar. Data on 429 trees distributed among five countries indicated that 20 of 38 Ficus species, and 46% of all trees sampled, had ants on their figs. Members of the Sycomorus subgenus were significantly more likely to attract ants than those in the Urostigma subgenus, and ant-colonization levels on these species were significantly greater than for Urostigma species. On average, each ant-occupied F.sur tree had 37% of its fig crop colonized by ants, whereas the value was 24% for other Ficus species. H. patruelis was the most common source for attracting ants, although figs were also attacked by a range of other ant-tended homopterans. P. megacephala was significantly more common on figs than other ant species, being present on 58% of sampled trees. Ant densities commonly exceeded 4.5 per fig, which a field experiment indicated was sufficient to provide protection from ovule gallers and parasitoids of pollinators. Forty-nine percent of all colonized F. sur trees sampled had ant densities equal to or greater than 4.5 per fig, whereas this value was 23% for other Ficus species. We conclude that there is considerable evidence to suggest that this indirect interaction occurs across four southern African countries and Madagascar, and involves many Ficus species. Received: 11 December 1997 / Accepted: 6 April 1998     

Fruiting Phenology and the Conservation of the Great Pied Hornbill (Buceros bicornis) in the Western Ghats of Southern India1

The phenology of principal fruits consumed by the endangered Great Pied Hornbill (Buceros bicornis) was monitored for two years in a wet forest habitat in southern India. Lipid–rich fruits, produced by several interior forest trees mainly of the family Lauraceae, were highly seasonal in their availability, and their production in the dry, hot season coincided with the breeding of the hornbill. Sugary fruits, produced mainly by several species of Ficus, were available year–round due to aseasonal fruiting patterns. Because Ficus fruited even at times of low fruit resource availability, and was heavily utilized by hornbills and other frugivores, it played a keystone role in the maintenance of the avian frugivore community. Overall fruit production was scarce between July and January during the southwest and northeast monsoon seasons. Vitex altissima produced berries abundantly during much of this time (September–December) and thus was another important fruit resource for avian frugivores. To safeguard the fruit resource base for the Great Pied Hornbill, we recommend: (1) The protection of Ficus and Vitex trees from overexploitation, and (2) the conservation of forest integrity to maintain compositions and densities of the lipid–rich fruit tree species utilized by the hornbill.     

西双版纳地区4种榕树果实的资源分配及其种子萌发特性

为了解榕树果实的资源分配和种子萌发特征,对西双版纳地区的垂叶榕(Ficus benjamina)、高山榕(F.altissima)、聚果榕(F.racemosa)和无柄雅榕(F.concinna)的果实营养、繁殖资源分配,以及种子萌发特性进行了研究。结果表明,4种榕果的营养分配存在差异,榕果重的种类果肉亦多,榕果大者果腔较大,且种子数量较多,但雄花和榕小蜂数量不一定多。榕果分配给雌性功能及雄性功能繁殖的小花比例也不相同,聚果榕在雌性功能上的分配显著大于雄性功能,其它3种榕果则分配于雄性功能的资源比雌性功能的多,但程度不同。此外,4种榕树的种子大小及重量存在显著的种间差异,以至于影响到种子的萌发率及根茎生长,果实最大的聚果榕种子最多,萌发率高,但由于种子最小,其根茎生长缓慢且短;果实适中的垂叶榕和高山榕种子较大,不但萌发率较高,其根茎生长也较快;果实和种子都小的无柄雅榕,种子萌发率低,根茎生长慢。生长在水热环境好的榕树比生长在相对贫瘠环境的榕树,由于极少面临水热光照等限制,更容易进化出较小的种子,萌发能力和速度也相对较差。这说明不同生境中榕树进化出了不同的繁殖分配机制及资源利用策略来适应环境变化。     

Fruiting phenology of fig trees in Kalinzu Forest,Uganda

The study determined the abundance and species composition of fig trees that fruited in the different forest types of Kalinzu Forest Reserve (KFR), Uganda. It also assessed the seasonal variations in abundance and species composition of the fig trees, the relationships between the fruiting patterns and rainfall and the figs’ inter‐ and intraspecific patterns of fruiting episodes. Sixteen fig species represented by 515 individuals were monitored monthly from December 2007 to January 2010. Most individuals and species that fruited were in the secondary forest types (the Musanga‐ and Parinari‐dominated secondary forests) and abundances of individuals of the different species were significantly associated with particular forest types. One colonizing species (Ficus sur) was the most abundant species that fruited and was mostly recorded in the secondary forests. Species composition and abundances of trees that fruited varied seasonally, and only the abundances of two canopy species (Ficus lingua and Ficus sansibarica) were significantly related with monthly rainfall. Most species experienced at least four fruiting phases, and F. sur displayed the longest episode covering 22 months. The results suggest that the past intensive logging in KFR promoted the regeneration of a diversity of fig species, and most species generally experience community‐wide asynchronous fruiting.     

Ficus seed shadows in a Bornean rain forest

Due to their copious seed production and numerous dispersers, rain forest fig trees have been assumed to produce extensive and dense seed shadows. To test this idea, patterns of seed dispersal of two species of large hemiepiphytic fig tree were measured in a Bornean rain forest. The sample included four Ficus stupenda and three F. subtecta trees with crop sizes ranging from 2,000 to 40,000 figs (400,000 to 13,000,000 seeds). Seed rain out to a distance of 60 m from each study tree was quantified using arrays of seed traps deployed in the understory. These trees showed a strongly leptokurtic pattern of dispersal, as expected, but all individuals had measurable seed rain at 60 m, ranging from 0.2 to 5.0 seeds/m². A regression of In-transformed seed rain density against distance gave a significant fit to all seven trees' dispersal patterns, indicating that the data could be fitted to the negative exponential distribution most commonly fitted to seed shadows. However, for six of seven trees, an improved fit was obtained for regressions in which distance was also In-transformed. This transformation corresponds to an inverse power distribution, indicating that for vertebrate-dispersed Ficus seeds, the tail of the seed rain distribution does not drop off as rapidly as in the exponential distribution typically associated with wind dispersed seed shadows. Over 50% of the seed crop was estimated to fall below each fig tree's crown. Up to 22% of the seed crop was dispersed beyond the crown edge, but within 60 m of the tree. Estimates of the maximum numbers of seeds which could have been transported beyond 60 m were 45% for the two largest crops of figs, but were under 24% for the trees with smaller crops. Seed traps positioned where they had an upper canopy layer above them were associated with higher probabilities of being hit by seeds, suggesting that vertebrate dispersal agents are likely to perch or travel through forest layers at the same level as the fig crown and could concentrate seeds in such areas to some degree. The probability of a safe site at 60 m from the fig tree being hit by seeds is calculated to be on the order of 0.01 per fruiting episode. Fig trees do not appear to saturate safe sites with seeds despite their large seed crops. If we in addition consider the rarity of quality establishment sites and post-dispersal factors reducing successful seedling establishment, hemiepiphytic fig trees appear to face severe obstacles to seedling recruitment.     

Some pollinators are more equal than others: Factors influencing pollen loads and seed set capacity of two actively and passively pollinating fig wasps

The nursery pollination system of fig trees (Ficus) results in the plants providing resources for pollinator fig wasp larvae as part of their male reproductive investment, with selection determining relative investment into pollinating wasps and the pollen they carry. The small size of Ficus pollen suggests that the quantities of pollen transported by individual wasps often limits male reproductive success. We assessed variation in fig wasp pollen loads and its influence on seed production in actively pollinated (Ficus montana) and passively pollinated (Ficus carica) dioecious fig trees.The ratios of number of male flowers on number of female flowers in a glasshouse-maintained F. montana population were highly variable. When fig wasps were introduced into receptive female figs, the resulting seed numbers were strongly linked to the numbers of pollinators that had been seeking access to pollen, relative to the number of anthers in their natal figs. In F. carica estimates of the amounts of pollen produced per fig and the quantities of pollen carried by emerging fig wasps suggest that less than 10% of the pollen is transported. Pollinators of F. carica that emerged earlier from figs carried more pollen, and also generated more seeds when introduced into receptive female figs.We show here that all pollinators are not equally valuable and producing more pollinators is not necessarily a good option in terms of Ficus male fitness. Previous results on F. montana figs showed that only around half of the flowers where pollinators lay eggs produced adult offspring. The amount of pollen collected by young female fig wasps may be a major determinant of their reproductive success.     

The Importance of Ficus (Moraceae) Trees for Tropical Forest Restoration

Forest restoration is an increasingly important tool to offset and indeed reverse global deforestation rates. One low cost strategy to accelerate forest recovery is conserving scattered native trees that persist across disturbed landscapes and which may act as seedling recruitment foci. Ficus trees, which are considered to be critically important components of tropical ecosystems, may be particularly attractive to seed dispersers in that they produce large and nutritionally rewarding fruit crops. Here, we evaluate the effectiveness of remnant Ficus trees in inducing forest recovery compared to other common trees. We studied the sapling communities growing under 207 scattered trees, and collected data on seed rain for 55 trees in a modified landscape in Assam, India. We found that Ficus trees have more sapling species around them (species richness = 140.1 ± 9.9) than non‐Ficus trees (79.5 ± 12.9), and significantly more saplings of shrub and large tree species. Sapling densities were twice as high under Ficus trees (median = 0.06/m²) compared to non‐Ficus (0.03/m²), and seed rain densities of non‐parent trees were significantly higher under Ficus trees (mean = 12.73 ± 3/m²/wk) than other fruit or non‐zoochorous trees (2.19 ± 0.97/m²/wk). However, our regression model found that canopy area, used as a proxy for tree size, was the primary predictor of sapling density, followed by remnant tree type. These results suggest that large trees, and in particular large Ficus trees, may be more effective forest restoration agents than other remnant trees in disturbed landscapes, and therefore the conservation of these trees should be prioritized.     

Sexual specialization in two tropical dioecious figs

Ficus species (figs) and their species-specific pollinator wasps are involved in an intimate mutualism in which wasps lay eggs in some ovaries of the closed inflorescences (syconia), and mature, inseminated offspring carry pollen from mature syconia to fertilize receptive inflorescences. In monoecious species, each syconium produces seeds and wasps. In functionally dioecious fig species, making up approximately half the figs worldwide, male and female functions are separated; hermaphrodite (functionally male) trees produce wasps and pollen only, while female trees produce seeds only. This sexual separation allows selection to act independently on the reproductive biology of each sex. Examining sexual specialization in a tight mutualism allows us to determine aspects of the mutualism that are flexible and those that are canalized. In this study, we quantified the phenology of two species of dioecious figs, F. exasperata and F. hispida, for 2 years by following the fates of several thousand syconia over time. In studying each of these species in a dry and a wet site in south India, we tested specific predictions of how dioecious figs might optimize sexual function. On female trees of both species, more inflorescences matured during the wet (monsoon) season than in any other season; this fruiting period enabled seeds to be produced during the season most suitable for germination. In F. exasperata, functionally male trees released most wasps from mature syconia in the dry season, during peak production of receptive female syconia, and thus maximized successful pollination. In F. hispida, “male” trees produced more syconia in the dry and monsoon seasons than in the post-monsoon season. In both species, male and female trees abscised more unpollinated, young inflorescences than pollinated inflorescences, but abscission appeared to be more likely due to resource- rather than pollinator- limitation. The phenology of F. exasperata requires that male inflorescences wait in receptive phase for scarce pollinators to arrive. As expected, male inflorescences of this species had a longer receptive phase than female inflorescences. In F. hispida, where pollinators are rarely scarce, duration of receptive phase was the same for both sexes. Duration of developing phase was longer in female syconia of both species than in male syconia, most likely because they need a longer period of investment in a fleshy fruit. Variation in developing phase of female syconia in one species (F. exasperata) was also greater than that in male syconia, and enabled female trees to sample a variety of germination environments in time. The strong sexual differences in both fig species support the hypothesis that selection for sexual specialization has strongly influenced the reproductive biology of these species. Received: 28 May 1997 / Accepted: 2 February 1998     

Moths boring into Ficus syconia on Iriomote Island, south-western Japan

Herbivory in the syconia of six Ficus (Moraceae) species (F. superba, F. varieagata, F. virgata, F. irisana, F. bengutensis and F. septica) was examined in March 2002 on Iriomote Island, south‐western Japan. Larvae of two lepidopteran species, Pachybotys spissalis (Guenée) (Pyralidae: Pyraustinae) and Stathmopoda sp. (Stathmopodidae) were observed to bore into the Ficus syconia. The attack rate by the moths varied from 0 to 38.5% across Ficus trees. The interiors of the syconia were heavily grazed by the moth larvae. Because figs (syconia) can be regarded as galls and seeds, according to sex and developmental stage, the moth larvae could be considered as gall or seed herbivores, and predators of fig wasps. Moth attack in the Ficus syconia could cause the destruction of fig wasp populations, as fig wasps develop in the syconia.     

Multiple parapatric pollinators have radiated across a continental fig tree displaying clinal genetic variation

The ways that plant‐feeding insects have diversified are central to our understanding of terrestrial ecosystems. Obligate nursery pollination mutualisms provide highly relevant model systems of how plants and their insect associates have diversified and the over 800 species of fig trees (Ficus) allow comparative studies. Fig trees can have one or more pollinating fig wasp species (Agaonidae) that breed within their figs, but factors influencing their number remain to be established. In some widely distributed fig trees, the plants form populations isolated by large swathes of sea, and the different populations are pollinated by different wasp species. Other Ficus species with continuous distributions may present genetic signatures of isolation by distance, suggesting more limited pollinator dispersal, which may also facilitate pollinator speciation. We tested the hypothesis that Ficus hirta, a species for which preliminary data showed genetic isolation by distance, would support numerous pollinator species across its range. Our results show that across its range F. hirta displays clinal genetic variation and is pollinated by nine parapatric species of Valisia. This is the highest number of pollinators reported to date for any Ficus species, and it is the first demonstration of the occurrence of parapatric pollinator species on a fig host displaying continuous genetic structure. Future comparative studies across Ficus species should be able to establish the plant traits that have driven the evolution of pollinator dispersal behaviour, pollinator speciation and host plant spatial genetic structure.     

The incidence and pattern of copollinator diversification in dioecious and monoecious figs

Differences in breeding system are associated with correlated ecological and morphological changes in plants. In Ficus, dioecy and monoecy are strongly associated with different suites of traits (tree height, population density, fruiting frequency, pollinator dispersal ecology). Although approximately 30% of fig species are pollinated by multiple species of fig‐pollinating wasps, it has been suggested that copollinators are rare in dioecious figs. Here, we test whether there is a connection between the fig breeding system and copollinator incidence and diversification by conducting a meta‐analysis of molecular data from pollinators of 119 fig species that includes new data from 15 Asian fig species. We find that the incidence of copollinators is not significantly different between monoecious and dioecious Ficus. Surprisingly, while all copollinators in dioecious figs are sister taxa, only 32.1% in monoecious figs are sister taxa. We present hypotheses to explain those patterns and discuss their consequences on the evolution of this mutualism.     

Frugivore assemblage of <Emphasis Type="Italic">Ficus superba</Emphasis> in a warm-temperate forest in Yakushima,Japan

Understanding fig consumption patterns is important because figs are regarded as a keystone resource for many frugivorous species in the tropics. While much work on fig consumption has been conducted in tropical regions, temperate forests are particularly interesting for study owing to pronounced seasonal variations in temperature and community-level fruiting phenology. We studied frugivore consumption of Ficus superba (Miq.) Miq. var. japonica Miq syconia in a warm-temperate forest in Yakushima, southern Japan. We conducted 141 4-h focal observations of fruiting F. superba trees over 12 months. We aimed to assess the relative quantitative contribution of each species of frugivore to F. superba consumption over a year as well as factors affecting seasonal variation in consumption. Japanese macaques were by far the most important F. superba syconia consumer (87.6 %), followed by brown-eared bulbuls (5.0 %), and varied tits (4.2 %). Japanese macaques increased their F. superba consumption when the temperature was high and fruit availability (F. superba and other species) was low. Macaques seemed to avoid searching for rare F. superba trees during winter and used F. superba syconia as a fallback food during fruit scarcity. Birds showed the opposite pattern: they increased F. superba syconia consumption when the temperature was low and fruit availability was high. This was probably because birds eat insects as their main food in the summer and switch to fruit as autumn turns to winter.