Population biology of figs: Applications for conservation

Summary The flowering phenology typical of at least monoecious figs-intra-tree synchrony and inter-tree asynchrony-poses problems for persistence of the pollinator population, and hence of the fig population itself, when fig population size is small. Establishment and maintenance of a population of the short-lived, species-specific wasp pollinator require that the fig population include a critical minimum number of trees (critical population size: CPS). Below CPS, temporal gaps between flowering trees occur that are unbridgeable by the pollinator, leading to its local extinction. This has implications for conservation in two contexts: human-aided invasions of introduced fig/wasp pairs, in which initial populations of figs and/or wasps may be small, and the persistence of figs and wasps in fragmented forest, in which initially large populations may be drastically reduced. Long-distance range extension by fig/wasp pairs is problematical for two reasons: 1) the fig species must first attain CPS, most likely through repeated seed dispersal events, before the wasp can establish; and 2) long-distance transit should be difficult for the tiny, short-lived wasp pollinators. I review the biology of natural and human-aided range extension by figs and fig wasps, and show that in human-aided range extensions these two difficult steps are circumvented. Once introduced into an area where hosts are abundant, fig wasps should readily establish from a small number of initial colonists, since they mate before dispersal and are highly tolerant of inbreeding. They are thus less subject than many insects to the genetic and demographic hazards of small population size. Of 5–6 fig/wasp pairs that have performed human-aided long-distance range extensions, one Asian pair,Ficus microcarpa and its pollinatorParapristina verticillata, is established in numerous areas in the northern neotropics, and the plant may become a serious weed. In tropical forests, figs may provide keystone resources for frugivores, providing fruit during seasons when other resources are scarce. Figs pose difficult problems for conservation biology, since minimum viable populations appear to be large, and since many species of tropical rainforests occur at low densities. This means that minimum areas required for persistence of a fig population- and for those of other species that would be affected were figs to be removed from the system-may often be large.     

A mutualism at the edge of its range

Summary Comparing populations that differ in access to mutualists can suggest how traits associated with these interactions have evolved. I discuss geographical and seasonal variation in the success of a primarily tropical mutualism (the fig/pollinator interaction), and evaluate some possible adaptations allowing it to persist at the edge of its range. Pollinators probably have difficulty in seasonal sites because 1) fig trees flower rarely in winter and 2) trees that do flower are less detectable and more difficult to reach. Fig biologists believe that seasonality must have selected for adaptations allowing pollinators to survive winter. However, geographical comparisons do not support two current ideas, the synchrony-breakdown hypothesis and the specificity-breakdown hypothesis. I pose two alternatives: plasticity of fruit and wasp developmental time, and adaptations of free-living fig wasps. I also distinguish between the impact of seasonality on monoecious versus dioecious figs; the latter group appear better adapted to reproduce in cool climates. A combination of comparative, observational, and experimental approaches has great potential for advancing our understanding of mutualisms.     

Classification and distribution ofFicus

Summary The classification of the genusFicus has changed considerably in the course of time and is still the subject of further research and discussion. The main subdivisions in the most recent classification by Corner¹² are presented together with the genera of pollinating fig wasps (Agaonidae) associated with them. These subdivisions are discussed and grouped according to morphological and functional traits, in particular in connection with the unique pollination system. Two main groups are recognized: one with only monoecious species and the other with predominantly (gyno)dioecious species. The former comprises two subgroups (Pharmacosycea andUrostigma) and the latter three, more profoundly different subgroups (Ficus, Sycidium andSycomorus). The neotropical representatives of the genus are discussed in somewhat more detail. In addition, the distribution of the genus is summarized for the three main regions of distribution; Africa, America, and Asia-Australasia. Finally the concordance between subdivisions ofFicus and those of the Agaonidae is briefly discussed.     

Structure and function of the fig

Summary An overview is given of the functional anatomy of the fig. The fig is the site of pollination and oviposition, it produces wasps, seeds and pollen, and it is the unit of dispersal. Therefore the fig acts as both a flower and a fruit. When a flower is both fertilized and receives a wasp egg, it eventually produces a wasp. Fertilization in flowers that do not recieve a wasp egg initiates seed production. Wide variation exists in the structural features of figs among different subgenera and sections. Monoecious species and gynodioecious species have different strategies for maintaining the symbiosis with the pollinating fig wasps. Monoecious species produce wasps and seeds in a single fig type and show imperfect heterostyly. Gynodioecious species have specialized figs for wasp and for seed production and are characterized by perfect heterostyly.     

Ecological and evolutionary dynamics of fig communities

Summary I review the status of five topics in fig research: pollen-vector versus seed production, flowering phenology and wasp population dynamics, monoecy versus dioecy, parasite pressure, and fig wasp behavior. I raise several new questions based on recent research on two components of fig reproduction: pollen-donation (male) and seed-production (female) success. I focus on how these two components of reproductive success depend on the flowering phenology of the figs and the population dynamics of the pollinator wasps.     

Phenology of fruit and leaf production by ‘strangler’ figs on Barro Colorado Island,Panamá

Summary Fruit and leaf initiation by 26 trees representing five stranglerFicus species in the subgenusUrostigma were monitored for 5–8 years in a seasonal lowland forest of central Panamá. Individual trees of each species initiated fruit in synchronized crops. High variation in the number of crops, intervals between crops and dates of crop initiation indicate that these species, like species in the subgenusPharmacosycea, initiate fruit crops the year around. Nevertheless, mean crop initiation dates for four of five species fell within the four-month dry season. Similarly, all species produced new leaf flushes throughout the year, however, mean leaf flush dates of all species fell within the first three months of the dry season.     

Preliminary results on the action of a plant growth regulator (Ethrel) in reducing the attack ofPrays oleae Bern. on olive fruits

Summary Preliminary experiments carried out in the Granada province of Southern spain over three years and in three different habitats have shown that a single application of Ethrel (2 chloro-ethyl phosphonic acid) to the olive tree at the beginning of fruit formation is sufficient to reduce significantly attack byPrays oleae Bern. and subsequent damage to the fruit. At the same time the treatment did not cause any detrimental change to the trees and had no measurable effect on the activity of beneficial insects such as Chrysopids (Neuroptera), which are known to be importantP. oleae egg predators in the study area.     

Pollen-collecting by stingless bees on cacao flowers

Summary Stingless bees (Trigona jaty) routinely visit the flowers ofTheobroma cacao (Sterculiaceae) in the Atlantic lowlands of Costa Rica. The bees collect pollen and behave as pollen thieves in flowers well exposed to direct sunlight in cacao plantations, and avoid flowers in heavy shade. Pollination rates are maximized, however, in heavy shade due to the high abundance of the small-bodied pollinating midges (Ceratopogonidae and Cecidomyiidae) found in such places. Pollen-thieving by stingless bees, therefore, may only impact on fruit set in cacao trees in direct sunlight, with only minimal to no impact in areas of cacao where natural pollinator activity is high.This research was funded by grants from The American Cocoa Research Institute of the United States of America. I thank T. Noeske for statistical assistance.     

Odour preference of a parasitic wasp depends on order of learning

FemaleLeptopilina boulardi wasps, larval parasites ofDrosophila melanogaster, can learn to respond to more than one odour by associating these odours with oviposition experience. These wasps can memorise and respond to at least two different odours, and prefer the last one learnt.     

Enemy-induced dispersal in a parasitic wasp

Females of the parasitic waspAphidius uzbekistanicus, a specialist aphid primary parasitoid, react to the presence of their specialist hyperparasitoid enemyAlloxysta victrix by leaving the area. The wasps leave to increase their reproductive success, because only their offspring, but not the wasps themselves, are threatened by enemy attack. Furthermore, this dispersal is elicited through the action of volatile chemical cues produced by the hyperparasitoids. We provide evidence for 6-methyl-5-hepten-2-one being one of the volatiles eliciting dispersal. The tendency of the wasps to leave areas which are being colonized by their enemies reduces their efficiency as aphid antagonists.     

The isolation,identification and synthesis of the alarm pheromone ofVespula squamosa (Drury) (Hymenoptera: Vespidae) and associated behavior

Summary A material that elicits alarm and attack behavior byVespula squamosa (Drury) workers was isolated from venom extracts and identified by spectroscopic methods as N-3-methylbutylacetamide. This compound elicited attack responses from worker wasps identical to those responses observed when venom was applied at the same dosage. This is the first behavioral role reported for this compound.The authors thank E. Adamak, R. Murphy and F. Takken for technical assistance. This article reports the results of research only. Mention of a proprietary product does not constitute an endorsement or the recommendation for its use by USDA.     

Body size variability and water balance: A comparison between mainland and island populations ofMastomys huberti (Rodentia: Muridae) in Senegal

Generally rodents are found to be larger on islands than on the mainland. However, there are some exceptions to this rule, and the aim of this paper is to examine one of them. On the mainland of Senegal,Mastomys huberti occupies humid habitats. However, it occurs also on dry and sandy islands (Saloum delta), where its representatives are dwarf. Since water availability appeared to be the limiting factor in these islands when compared to the mainland, we studied water turnover characteristics in relation to body size, in mainland and island populations at the end of the dry season, under both field and laboratory conditions. All populations were found to be water balanced in their natural habitats. They presented similar rates of water turnover, even though island animals were subjected to stronger constraints than mainland ones. Laboratory experiments suggested that the physiological plasticity of one of the island populations may be reduced. Island populations have a higher kidney size to body weight ratio than those from the mainland. We propose that smaller size in the islands allows the maintenance of water balance with a smaller amount of water, and that a higher ratio of kidney filtration surface to body size may helpMastomys huberti to survive in dry islands. We discuss the factors responsible for body size variability and variation in water exchange characteristics and conclude that different factors could explain body size variation among island populations, depending on the species considered and the ecological constraints met within the islands.     

Experimential evidence for endogenously programmed differential migration in the blackcap (Sylvia atricapilla)

Female hand-raised blackcaps (Sylvia atricapilla) held in constant conditions (except for simulated seasonal changes in photoperiod) showed significantly more, and significantly longer, autumnal migratory activity as well as significantly later initiation of spring migratory activity than males from the same population (including siblings) held under identical conditions.This research was supported by the Alexander von Humboldt Foundation (Terrill) and the Deutsche Forschungsgemeinschaft (Berthold).     

Cortinarins inCortinarius speciosissimus? A critical revision

Summary Fluorescent cyclic decapeptides, the so-called cortinarins, have been claimed to cause the toxicity of the deadly poisonous mushroomCortinarius speciosissimus. As the structure elucidation described in the literature is doubtful in decisive parts, the proposed structures must be disputed. The present study demonstrates that the fluorescence is mainly due to ergosta-4,6,8 (14),22-tetraen-3-one (8) and to photodecomposition products of the nephrotoxin orellanine (1). Moreover, we were not able to reproduce the isolation of fluorescent or any other cyclic peptides related to cortinarins fromC. speciosissimus, by various methods.     

The evolutionary history ofDrosophila buzzatii. V. Differential survivorship onOpuntia betweenD. buzzatii andD. serido

Summary Survival time ofDrosophila buzzatii adults on anOpuntia (prickly pear) medium was significantly longer than that of its nearest relativeD. serido. A significant difference was also found betweenD. buzzatii adults from two experimental populations, one of them fed onOpuntia rots for more than two years and another one kept on standardDrosophila medium for the same period of time. These results suggest that adult selection may be taking place in cactiphilicDrosophila in their natural habitats and could be responsible for the niche differentiation betweenD. buzzatii andD. serido.This work has been supported by the Comisión Asesora de investigación Científica y Técnica, Spain, by funds to project 4514/79 awarded to the third author.     

The elusive engine in <Emphasis Type="Italic">Myxococcus xanthus</Emphasis> gliding motility

Bacterial motility is essential for chemotaxis, virulence and complex social interactions leading to biofilm and fruiting body formation. Although bacterial swimming in liquids with a flagellum is well understood, little is known regarding bacterial movements across solid surfaces. Gliding motility, one such mode of locomotion, has remained largely mysterious because cells move smoothly along their long axis in the absence of any visible organelle. In this review, I discuss recent evidence that focal adhesion systems mediate gliding motility in the social bacterium Myxococcus xanthus and combine this evidence with previous work to suggest a new working hypothesis inspired from knowledge in apicomplexan parasites. I then propose experimental directions to test the model and compare it to other pre-existing models. Finally, evidence on gliding mechanisms of selected organisms are presented to ask whether some features of the model have precedents in other bacteria and whether this complex biological process could be explained by a single mechanism or involves multiple distinct mechanisms. Received 12 April 2007; received after revision 8 June 2007; accepted 27 June 2007     

(S)-(+)-4,4,4-Trifluoro-3-(indole-3-)butyric acid,a novel fluorinated plant growth regulator

Racemic 4,4,4-trifluoro-3-(indole-3-)butyric acid (TFIBA) has been synthesized and shown to inhibitAvena coleoptile elongation. (S)-(+)-TFIBA (fig. 1), which was prepared by an enzymatic method and markedly promotes root growth of Chinese cabbage, lettuce and rice plants, is a novel fluorinated plant growth regulator. Activity of the (S)-(+)-enantiomer of TFIBA was 10-fold greater than that of the (R)-(–)-enantiomer in the first two plant species and 5-fold greater in rice.     

Loss of FrmA leads to increased cell-cell adhesion and impaired multi-cellular development of Dictyostelium cells

Cell-cell adhesion is a critical property of all multi-cellular organisms and its correct regulation is critical during development, differentiation, tissue building and maintenance, and many immune responses. The multi-talin-like FERM domain containing protein, FrmA, is required during starvation-induced multi-cellular development of Dictyostelium cells. Loss of FrmA leads to increased cell-cell adhesion and results in impaired multi-cellular development, slug migration and fruiting bodies. Further, mixing experiments show that FrmA null cells are excluded from the apex of wild-type mounds, to which cells that normally form the organising centre known as the tip sort. These data suggest a critical role for FrmA in regulating cell-cell adhesion, multi-cellular development and, in particular, the formation of the organising centre known as the tip. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Received 28 August 2008; received after revision 10 October 2008; accepted 21 October 2008     

Two classes of metabolites fromTheonella swinhoei are localized in distinct populations of bacterial symbionts

The marine spongeTheonella swinhoei (lithistid Family Theonellidae, Order Astrophorida) has yielded many important, bioactive natural products, most of which share structural features with bacterial natural products. The presence of microbial symbionts inT. swinhoei has been reported, and it was originally suggested that the cytotoxic macrolide swinholide A and many of the bioactive cyclic peptides fromT. swinhoei were all produced by symbiotic cyanobacteria. By transmission electron microscopy, we found four distinct cell populations to be consistently present inT. swinhoei: eukaryotic sponge cells, unicellular heterotrophic bacteria, unicellular cyanobacteria and filamentous heterotrophic bacteria. Purification and chemical analyses of each cell type showed the macrolide swinholide A to be limited to the mixed population of unicellular heterotrophic bacteria, and an anti-fungal cyclic peptide occurred only in the filamentous heterotrophic bacteria. Contrary to prior speculation, no major metabolites were located in the cyanobacteria or sponge cells.     

Double cone mosaic pattern in the retina of larval and adult piranha,Serrasalmus spilopleura

The mosaic pattern of double cones of retinas in larval and adult piranha,Serrasalmus spilopleura, was examined. Samples from two habitats in the same reservoir, characterized by different light conditions, were examined. The same square pattern was found in all sampled retinas indicating that, for this species, there is no apparent relation between the patterns of double cone arrangement and environmental luminosity.