Seasonality in the reproductive phenology ofFicus: Its evolution and consequences

Summary Ficus can only reproduce if they are pollinated by mutualistic wasps that breed within the figs. Pollen-loaded wasps enter the figs when the female flowers are receptive. Several weeks later, their offspring load pollen within the fig and then emerge. As individual trees typically produce crops of synchronous figs at long intervals, the shortlived wasps have to move to another, receptive, tree. The wasp population can only survive, and hence the fig population reproduce, if there are trees fruiting all over the year. When only few trees are present within a population gaps in the flowering sequence may lead to the extinction of the local pollinator population. Two models are presented. One investigates the number of trees necessary in order to sustain a local pollinator population when the tree population has a seasonal pattern of fruiting. The second model investigates how such a seasonal pattern may evolve within a fig population as a result of individual selection on the trees. It is shown that pollinator populations are fragilized under seasonal conditions. Hence, the breeding system ofFicus limits their expansion into highly seasonal habitats. Seasonal habitats may also lead to seasonal adjustment of male versus female investments and to the evolution of dioecy.     

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.     

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.     

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.     

Ancient DNA from Bronze Age bones of European rabbit (Oryctolagus cuniculus)

The European rabbit (Oryctolagus cuniculus) is now widely distributed throughout the world as a result of transportation by man. The original populations, however, were confined to southern France and Spain. In order to investigate the role of human intervention in determining the genetic diversity of rabbit populations, we are studying the origin of rabbits introduced onto a small Mediterranean island (Zembra) near Tunis over 1400 years ago, by examining ancient DNA extracted from rabbit bones found both on Zembra and on the European mainland. Ancient DNA was successfully extracted from rabbit bones found at two archaeological sites dated to at least the Early Bronze Age (more than 3500 years ago) in south-central France, and compared to that found in modern mainland and island populations using a small variable region of the cytochromeb gene. The results confirm that the Zembra Island population is descended from that present over 1400 years ago. The technical aspects of DNA extraction from bones and the implications of this type of research for determining the origin of introduced rabbit populations are discussed.     

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.     

Microtubule inhibitors block the morphological changes induced inDrosophila blood cells by a parasitoid wasp factor

Summary The shape change ofDrosophila melanogaster blood cells (lamellocytes) from discoidal to bipolar that is caused by a factor from the female parasitoidLeptopilina heterotoma is blocked by the tubulin inhibitors vinblastine and vincristine in vitro. The actin inhibitor, cytochalasin B, causes arborization ofDrosophila lamellocytes and acts synergistically with the wasp factor to alter lamellocyte morphology. Lamellocyte arborization induced by cytochalasin B is blocked by simultaneous treatment with vinblastine. These observations indicate that the changes in lamellocyte shape induced by both the wasp factor and cytochalasin B require microtubule assembly.     

Photoperiodic induction of diapause in an insect is vitamin A dependent

Summary A photoperiodic response was found to be absent in larvae of the parasitoid waspApanteles glomeratus when its host (caterpillars ofPieris brassicae) was reared on a low-carotenoid artificial diet. Addition of vitamin A to the host's diet restored the response to short-day photoperiods in the wasp larvae, thus showing that vitamin A is essential for photoperiodic induction of diapause. Possibly vitamin A or a derivative of vitamin A functions as the photoreceptor pigment for the photoperiodic reaction in this species of insect.     

A new chromosome number forBombina (Anura,Discoglossidae)

Summary The karyotype of a primitive discoglossid anuran,Bombina maxima, native to southwestern China, has 2n=28 chromosomes with 6 large and 8 small bi-armed homologous pairs. This is a higher chromosome number than described for otherBombina species, all of which have 2n=24.     

Macrobiotus pseudohufelandi Iharos as a model for cytotaxonomic study in populations of eutardigrades (Tardigrada)

Summary The morphotype, chromosome number and Feulgen-DNA content of bisexual and unisexual populations ofMacrobiotus pseudohufelandi were examined. Individuals of unisexual populations were triploid with ameiotic parthenogenesis. Their lowest Feulgen-DNA content is about three-fold that of sperm from a bisexual population. Egg shell shape also differs in the two types of population. However, the highest Feulgen-DNA content was the same (24 A.U.) in both diploid and triploid animals.Study funded by MPI grant.     

Gel-electrophoretic description of European populations ofTerellia virens (Loew) (Diptera,Tephritidae); implications for its use as an agent for the biological control ofCentaurea spp. (Asteraceae) in North America

Allozyme frequencies of 15 enzyme loci, 14 of which were polymorphic, were used to characterize sevenTerellia virens populations originating from three allopatrically distributedCentaurea species. The two populations whose origins were geographically furthest apart, from Israel (onC. iberica) and from Switzerland (onC. vallesiaca), showed relatively high values of genetic distance from the 5 populations sampled in Austria and Hungary (onC. maculosa) (Nei's D>0.07). The latter five displayed a high degree of genetic similarity. No diagnostic (fixed) allelic differences were observed between these three groups ofT. virens populations, but they could be well characterized by significant differences in allelic frequencies at 9 enzyme loci. Independently of this study, the populations from Switzerland (C. vallesiaca) and eastern Austria (C. maculosa) were selected as potential source populations for future introductions into North America for the biological control of introducedC. maculosa andC. diffusa. Based on the observed genetic differences and results from field experiments on the host specificity of these two potential source populations, it is argued that host specificity screening tests should be conducted separately for local (host plant) populations, as such populations might accept a different set of hosts. Biotype mismatch and the risk of spill-overs to native species could thus possibly be reduced.     

Soluble proteins of chemical communication in the social wasp <Emphasis Type="Italic">Polistes dominulus</Emphasis>

Members of the odorant-binding protein (OBP) and chemosensory protein (CSP) families were identified and characterised in the sensory tissues of the social wasp Polistes dominulus (Hymenoptera: Vespidae). Unlike most insects so far investigated, OBPs were detected in antennae, legs and wings, while CSPs appeared to be preferentially expressed in the antennae. The OBP is very different from the homologous proteins of other Hymenopteran species, with around 20% of identical residues, while the CSP appears to be much better conserved. Both OBP and CSP, not showing other post-translational modifications apart from disulphide bridges, were expressed with high yields in a bacterial system. Cysteine pairing in the recombinant and native proteins follows the classical arrangements described for other members of these classes of proteins. OBPs isolated from the wings were found to be associated with a number of long-chain aliphatic amides and other small organic molecules. Binding of these ligands and other related compounds was measured for both recombinant OBP and CSP.Received 14 May 2003; received after revision 8 June 2003; accepted 12 June 2003     

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.     

Heterochromatin characterization and distribution in the chromosomes of two populations ofIdotea baltica basteri Audouin, 1826 (Isopoda,Valvifera)

Two mediterranean populations ofIdotea baltica basteri from Messina and Naples showed a set of chromosomes composed by 58 all-biarmed chromosomes. The heterochromatin was located in the pericentromeric region of the chromosomes, and its composition appeared heterogeneous. In fact, not all the homologs showed heterochromatin resistant to digestion with three restriction enzymes (Alu I, Hae III and Sau 3A). Moreover, the two populations showed polymorphism in a band of G+C-rich telomeric heterochromatin, which was present only in the population from Messina. It is hypothesized that chromosomal polymorphism might reflect the geographical isolation of the two populations. It is also suggested that a process of diversification is taking place.     

Immunological comparison between albumins of three species of mice (genusMus)

Summary Three closely related species of short-tailed mice (Mus musculus musculus, M. spretoides andM. spicilegus) were tentatively discriminated using immunological techniques based on albumin cross-reactivity. Different fractionations of crude albumin antisera allowed the recovery of antibody populations specific to theM. m. musculus albumin, whereas antibody population differences do not seem to exist betweenM. spicilegus andM. spretoides. Moreover, immunoreactivities tested with native and S-carboxymethylated albumins revealed that species-specific antibodies correspond to antigenic determinants depending on the amino acid sequence (sequential determinants). The observed immunological differences are related to species divergence and albumin sequences.     

Population genetics of a colonizing lizard: Loss of variability in introduced populations ofPodarcis sicula

Allozyme electrophoresis was used to study the genetic variability (proportion of polymorphic loci and heterozygosity) in insular populations (Corsica, Elba, Montecristo, Marettimo, Pantelleria) of the lacertid lizardPodarcis sicula. These populations were presumed to have originated from episodes of accidental anthropogenic introduction. In order to test the hypothesis of a man-aided colonization and to provide comparative data, heterozygosity and polymorphism were also estimated in autochthonous populations ofP. sicula from the Italian peninsula and Sicily. In each case, the presumed introduced population showed levels of genetic variability significantly lower than those detected in the autochthonous ones. Very little genetic differentiation was found among native and presumed colonist populations, Nei's standard genetic distances ranging from 0.001 to 0.009. These results strongly support the hypothesis thatP. sicula was only recently introduced to the studied islands, and provide additional evidence of reduced genetic variability due to founder effect in insular populations originating from episodes of human transportation.     

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.     

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.     

Asiatic cobras: Systematics and snakebite

Summary The population affinities of the Asiatic cobras of the genusNaja are investigated, using multivariate analysis of a range of morphological characters. This complex, which was formerly thought to be monospecific, consists of at least eight full species. In some cases, species whose bites require different antivenoms occur sympatrically. The new understanding of the systematics of the Asiatic cobra complex calls for a reappraisal of cobra antivenom use in Asia, and for more research into venom composition.