Substrate water potential constraints on germination of the strangler fig Ficus aurea (Moraceae)

Palms are the most common support hosts for strangler fig (Ficus aurea) in the tropical dry forest. At Hummingbird Cay Tropical Field Station in the Bahamas, viable F. aurea seeds scattered on open soil germinate within 6 d in the laboratory at 27C in light or darkness. To test the hypothesis that water relations between seed and substrate can restrict the site of F. aurea establishment, fig seeds were imbibed in mannitol solutions with water potential (psi) from 0 MPa to -2.0 MPa. At psi > -1.0 MPa, germination ranges from 70 to 90%. Below -1.0 MPa, germination drops under 4%. Seedling growth rate slows linearly with decreasing psi, due to reduced cell enlargement in the radicle. Sensitivity of F. aurea seeds to psi stress is similar to that of mesophytic crops with "threshold' psi of -1.2 to -1.5 MPa, below which germination and/or seedling growth stops. F. aurea shows no evidence of xerophytic adaptation in germination physiology. In the field, psi of humus in palm leaf bases does not drop below -1.0 to -1.1 MPa, while humus in terrestrial sites reaches psi F. aurea germination and seedling growth is met by humus in palm leaf bases, not by terrestrial sites. In dry forests, this restricts F. aurea establishment to the crown of palm trees.     

Characterization of microsatellite loci in the African fig Ficus sycomorus L. (Moraceae)

Microsatellite loci were characterized in the African fig tree Ficus sycomorus in order to investigate patterns of pollination and gene flow in this species. The loci characterized included new loci isolated from F. sycomorus and a single locus originally developed in Ficus carica. In total 12 loci were polymorphic when tested in between eight and 79 Namibian F. sycomorus individuals. Three of the new F. sycomorus loci were found to be polymorphic in cultivars of the edible fig F. carica suggesting a selection of these loci will be useful for population studies in other fig species.     

Anonymous and EST-based microsatellite DNA markers that transfer broadly across the fig tree genus (Ficus, Moraceae)

? Premise of the study: We developed a set of microsatellite markers for broad utility across the species-rich pantropical tree genus Ficus (fig trees). The markers were developed to study population structure, hybridization, and gene flow in neotropical species. ? Methods and Results: We developed seven novel primer sets from expressed sequence tag (EST) libraries of F. citrifolia and F. popenoei (subgen. Urostigma sect. Americana) and optimized five previously developed anonymous loci for cross-species amplification. The markers were successfully tested on four species from the basal subgenus Pharmacosycea sect. Pharmacosycea (F. insipida, F. maxima, F. tonduzii, and F. yoponensis) and seven species of the derived subgenus Urostigma (F. citrifolia, F. colubrinae, F. costaricana, F. nymphaeifolia, F. obtusifolia, F. pertusa, and F. popenoei). The 12 markers amplified consistently and displayed polymorphism in all the species. ? Conclusions: This set of microsatellite markers is transferable across the phylogenetic breadth of Ficus, and should therefore be useful for studies of population structure and gene flow in approximately 750 fig species worldwide.     

Phylogeny, biogeography, and ecology of Ficus section Malvanthera (Moraceae)

We conducted the first molecular phylogenetic study of Ficus section Malvanthera (Moraceae; subgenus Urostigma) based on 32 Malvanthera accessions and seven outgroups representing other sections of Ficus subgenus Urostigma. We used DNA sequences from the nuclear ribosomal internal and external transcribed spacers (ITS and ETS), and the glyceraldehyde-3-phosphate dehydrogenase (G3pdh) region. Phylogenetic analysis using maximum parsimony, maximum likelihood and Bayesian methods recovered a monophyletic section Malvanthera to the exclusion of the rubber fig, Ficus elastica. The results of the phylogenetic analyses do not conform to any previously proposed taxonomic subdivision of the section and characters used for previous classification are homoplasious. Geographic distribution, however, is highly conserved and Melanesian Malvanthera are monophyletic. A new subdivision of section Malvanthera reflecting phylogenetic relationships is presented. Section Malvanthera likely diversified during a period of isolation in Australia and subsequently colonized New Guinea. Two Australian series are consistent with a pattern of dispersal out of rainforest habitat into drier habitats accompanied by a reduction in plant height during the transition from hemi-epiphytic trees to lithophytic trees and shrubs. In contradiction with a previous study of Pleistodontes phylogeny suggesting multiple changes in pollination behaviour, reconstruction of changes in pollination behaviour on Malvanthera, suggests only one or a few gains of active pollination within the section.     

粗叶榕（Focis hirta）繁殖系统的特征及其共生的榕小蜂

本文对粗叶榕(Focis hirta)的外部形态、花序结构以及寄主和传粉者的繁殖特性作了比较详细的观察和研究,讨论了它们互惠共生关系的一些基本特征。它的专性传粉者是榕小蜂科(Agaonidae)榕小蜂亚科(Agaoninae)的爪哇榕小蜂(Blastophaga javana Mayr)。观察发现粗叶榕还存在两种非授粉小蜂Sycoscapter sp．和Philotrypesis sp．,它们在榕果外产卵。     

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.     

粗叶榕(Ficus hirta)繁殖系统的特征及其共生的榕小蜂

本文对粗叶榕(Ficus hirta Vahl)的外部形态、花序结构以及寄主和传粉者的繁殖特性作了比较详细的观察和研究, 讨论了它们互惠共生关系的一些基本特征。它的专性传粉者是榕小蜂科(Agaonidae)榕小蜂亚科(Agaoninae)的爪哇榕小蜂(Blastophaga javana Mayr)。观察发现粗叶榕还存在两种非授粉小蜂Sycoscapter sp.和Philotrypesis sp.,它们在榕果外产卵。     

粗叶榕(Ficus hirta)繁殖系统的特征及其共生的榕小蜂

本文对粗叶榕(Ficus hirta Vahl)的外部形态、花序结构以及寄主和传粉者的繁殖特性作了比较详细的观察和研究,讨论了它们互惠共生关系的一些基本特征.它的专性传粉者是榕小蜂科(Agaonidae)榕小蜂亚科(Agaoninae)的爪哇榕小蜂(Blastophaga javana Mayr).观察发现粗叶榕还存在两种非授粉小蜂Sycoscapter sp.和Philotrypesis sp.,它们在榕果外产卵.     

Pollinator sharing in dioecious figs (Ficus: Moraceae)

As one of the most specialized pollination syndromes, the fig (Ficus)–fig wasp (Agaonidae) mutualism can shed light on how pollinator behaviour and specificity affect plant diversification through processes such as reproductive isolation and hybridization. Pollinator sharing among species has important implications for Ficus species delimitation and the evolutionary history of the mutualism. Although agaonid wasp pollinators are known to visit more than one host species in monoecious figs, pollinator sharing has yet to be documented in dioecious figs. The present study investigated the frequency of pollinator sharing among sympatric, closely‐related dioecious figs in Ficus sections Sycocarpus and Sycidium. Molecular and morphological species identification established the associations between pollinating agaonid wasp species and host fig species. Cytochrome oxidase I was sequenced from 372 Ceratosolen pollinators of Ficus section Sycocarpus and 210 Kradibia pollinators of Ficus section Sycidium. The association between fig species and morphologically distinct clades of pollinator haplotypes was predominantly one‐to‐one. In Ceratosolen, six of 372 pollinators (1.5%) visited fig species other than the predominant host. No pollinator sharing was detected between the two Sycidium host species, although a rare hybrid shared Kradibia pollinators with both parental species. These findings point to low rates of pollinator sharing among closely‐related dioecious fig species in sympatry, and perhaps lower rates than among monoecious figs. Such rare events could be evolutionarily important as mechanisms for gene flow among fig species. Differences in rates of pollinator sharing among fig lineages might explain the conflicting phylogenetic patterns inferred among monoecious figs, dioecious figs, and their respective pollinators. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 546–558.     

Host associations of the strangler fig Ficus watkinsiana in a subtropical Queensland rain forest

Abstract The distribution of epiphytic organisms is limited by the availability of, and dispersal to suitable hosts. We examined the distribution of a hemi‐epiphytic strangler fig, Ficus watkinsiana (Moraceae) in Cooloola National Park (Queensland, Australia), in order to determine whether this species exhibits a preference for certain host species and why. We assessed host bark roughness and flakiness, fruit type, and size to explain the observed distribution of F. watkinsiana. We surveyed over 1900 potential host trees of the 30 most common forest canopy species and found that host size measured by diameter at breast height accounted for most variation in fig prevalence (Binary Logistic Regression log‐likelihood = ?588.178, G = 314.494, d.f. = 1, P < 0.005). After controlling for host size, F. watkinsiana prevalence still differed significantly between host species (χ² = 54.612, d.f. = 24, P < 0.005), a difference that was only partly explained by variation in the bark roughness of host trunks. These results suggest that variation in the rate at which tree species host strangler figs are primarily related to individual tree size – figs may simply be more likely to colonize and thrive upon host species that grow larger.     

Figs (Ficus,Moraceae) in Urban Hong Kong,South China1

Hong Kong (22°N) is on the northern margins of the Asian tropics and has a native fig flora of 24 species. A total of 3.4 km² of the urban area on Hong Kong Island was surveyed for spontaneous and planted fig plants of reproductive size. The 1124 individuals included 14 species in four subgenera: seven native species (F. fistulosa, F. hirta, F. hispida, F. microcarpa, F. pumila, F. subpisocarpa, and F. variegata), four of which were sometimes planted, one probably native (F. virens), one naturalizing species (F. religiosa), and five exotic species that occurred only in cultivation, two of which (F. altissima, F. rumphii) are pollinated and produce viable seeds. The native species in the two most common subgenera form distinct ecological guilds: those in subgenus Sycomorus bear large, many‐seeded, green or yellow figs and are bat‐dispersed pioneers on exposed soil; those in subgenus Urostigma bear small, few‐seeded, dark‐purple figs and are largely bird‐dispersed and epilithic. The density of potentially fruiting fig plants in the study area (2.3/ha) was within the range reported for tropical forests and between them they were visited by the entire urban frugivore fauna. This study shows the importance of the urban fig flora to urban wildlife and also highlights the risk of cultivated Ficus species becoming invasive, despite their obligate species‐specific pollinator mutualisms.     

Ficus (Moraceae) and Hymenoptera (Ghalcidoidea): Figs and their pollinators

Attempts are made to meet discrepancies between the botanical classification of Ficus and the zoological classification of its pollinators. They are discussed as Ficus elastica , subgen. Pharmacosycea , subsect. Palaeomorphe , dioecism in Ficus , the Ceratosolen complex, and sect. Conosycea. It seems from the botanical side that the evolution of Ficus into subgenera and sections preceded that of its insects into genera. On the whole there are no grounds for reclassification of Ficua , but leaf structure indicates that some rearrangement of the species of subgen. Urostigma sect. Conosycea may be needed to fit the entomology. Ficus asperiuscula poses a problem in sect. Sycidium.     

Biogeography and conservation of the genus Ficus (Moraceae) in Mexico

Aim The main objective of this study is to document the biogeographical patterns, endemism and degree of conservation of the species of Ficus (Moraceae) in Mexico. There are over 750 species of the genus Ficus distributed worldwide, and Mexico practically represents its northernmost limit in the American continent. Detailed studies at regional scales may help to understand the biogeography of large genera such as Ficus. Location Mexico. Methods The biogeographical patterns of Mexican Ficus were obtained from information of fig specimens available in two of the main herbaria of Mexico (2140 vouchers), collecting figs throughout this country, and revising the specialized literature. The presence of each species of Ficus was recorded for every one of Mexico's states and several tropical countries of America. Besides, the Mexican territory was divided into cells of 1° × 1° and the presence or absence of all species of the genus was recorded. Rarity of species was classified based on the width of geographic distribution, habitat specificity and population size. Results A total of 21 species of Ficus occur in Mexico, including six species (28.6%) that are endemic to this country. Five species are included in subgenus Pharmacosycea and 16 species are documented under subgenus Urostigma. Affinities of Ficus flora with other tropical countries in America generally decreased as geographical distances from Mexico increased. Mexican states and cells with highest values of Ficus species richness (both total and endemic species) were located. Ten species, including three endemics, presented a wide distribution. Five species, including two endemics, possess the three attributes of rarity (narrow geographical distribution, high habitat specificity and scarce local populations). Three species of Ficus, including the endemic and very rare Ficuslapathifolia (Liebm.) Miq., are not recorded in any protected area existing in Mexico. Main conclusions Most of the Mexican Ficus show a great morphological variation and occupy different habitats along their geographic distribution. The biogeographical patterns described here establish a fundamental scenario for ongoing studies on Ficus–pollinator interactions. However, many local populations are considered to be at risk, as there have been significant reductions in the number and size of local populations. Further studies are needed to understand the process of colonization, maintenance and persistence of fig–pollinator mutualism in species with different patterns of geographic distribution. Mexican Ficus require special policies for conservation due to their complex degree of rarity, particularly their geographic distribution in different types of vegetation, ranging from dry scrublands to tropical rain forests.     

Morphological diversity and function of the stigma in Ficus species (Moraceae)

The stigma plays several roles such as pollen hydration and selection, and pollen tube nutrition. In the Ficus-fig wasp mutualism, stigmata have an additional, almost unknown, function by representing a physical interface for both plant and wasp reproduction. We used light and electron microscopy to compare the detailed morphology of the stigmata of nine Ficus species of different sections and with different pollination modes and sexual expressions. Figs were collected at the stage when the stigmata were receptive for pollination. Stigmata in actively pollinated monoecious species have well developed papillae concentrated on the adaxial surface exposed towards the fig cavity. Conversely, the passively pollinated monoecious species have the whole surface of the stigmata covered by somewhat smaller papillae. In both actively and passively pollinated monoecious species these features are consistent, irrespective of style length. In all actively pollinated gynodioecious species, the stigmata of pistillate flowers were tubular or infundibuliform whereas in almost all actively pollinated monoecious species (except F. racemosa) the stigmata were filiform, with one branch or two asymmetric branches. In gynodioecious species the short-styled flowers in “male” figs show a limited receptive surface with small papillae, while the stigmata of long-styled flowers in “female” figs are covered by papillae that extend down the sides of the style, increasing the stigmatic surface. In actively pollinated species, stigmata are cohesive, forming a common surface for pollen tube germination (= synstigma). The synstigma arrangement was quite variable: lax, cohesive or very cohesive, with entanglement by stigmatic papillae and stylar trichomes. Entanglement by stylar trichomes is common in gynodioecious species. The synstigma arrangement did not correlate with phylogeny or breeding system. This study is the first to report a very loose synstigma in actively pollinated monoecious Ficus species. Our analyses revealed that, in Ficus, the synstigma is functionally analogous to an extra-gynoecial compitum. Comparative studies will be required to test further hypotheses about the evolutionary determinants of such variation.     

Chemotaxonomic significance of leaf alkanes in species of Ficus (Moraceae)

The alkane pattern of leaf waxes from 24 tropical Ficus species in Nigeria was determined by gas chromatography and gas chromatography–mass spectrometry. Of the 12 alkanes identified, hentriacontane and tritriacontane were the major components in all the species studied. The chemosystematic significance of leaf alkanes is discussed in Ficus for the first time. There is an indication that alkane occurrence in Ficus could provide useful information to the understanding of species variability.     

Different ontogenetic processes promote dicliny in Ficus L. (Moraceae)

The absence of reproductive organs in flowers may ontogenetically arise from inception or by abortion during development. Ficus L., a species-rich genus of angiosperms, is an interesting model for floral developmental studies because of the diversity of sexual systems it contains. This study compares the floral morphology of Ficus citrifolia (monoecious), Ficus religiosa (monoecious), Ficus racemosa (secondarily monoecious), and Ficus hispida (gynodioecious) across development to establish the ontogenetic pathways that result in diclinous flowers. Figs were collected at various developmental stages and were prepared for surface (scanning electron microscopy) and histological (light microscopy) analyses. Dicliny in Ficus is defined by stamen absence from inception in pistillate flowers and either pistil absence from inception (F. citrifolia, F. racemosa and F. religiosa) or by abortion (F. hispida) in staminate flowers. The perianth is formed by a single whorl of sepals, as found in other families related to Moraceae. The gynoecium is tubular during development, a condition that may be related with pseudomonomery. The staminate and neutral flowers in F. hispida develop by similar mechanisms. The diversity in the sexual systems in Ficus results from combinations of different floral morphs (dicliny), which originate from both previously established ontogenetic mechanisms (loss of reproductive organ function by abortion or from inception). These mechanisms act independently of phylogenetic proximity or mechanisms of sex system evolution in Ficus. Other aspects of floral development observed in Ficus are discussed in relation to their systematic position and reproductive biology.     

Flavonoids and other bioactive constituents from Ficus thonningii Blume (Moraceae)

Continued interest in the chemistry of Ficus spp. led to the investigation of the figs and the roots of Ficus thonningii Blume. Two new flavonoids, thonningiol (1) and thonningiisoflavone (2) along with nineteen known compounds were isolated. β-Isoluteone (13) was isolated here for the first time from a natural source. Their structures were elucidated on the basis of spectroscopic evidence. Interestingly, thonningiisoflavone (2) and hydroxyalpinumisoflavone (21) showed strong DPPH radical scavenging activity with IC₅₀ = 65.50 μM and 68.20 μM respectively compared to the standard BHA with IC₅₀ = 44.20 μM. The methanolic extract of figs, taxifolin (14), conrauiflavonol (17) and shuterin (19) exhibited moderate antimicrobial activity against six micro-organisms with MIC below 1.5 mg/mL.