Programme

4^th

                                    Arrival of participants

5^th        

            Morning:           Arrival of participants and registration

            2 pm                 Introduction to workshop – Rhett Harrison & Allen Herre

            2:30 pm             Brief introduction to Lambir Hills National Park and Research activities – Rhett

            3:00 pm             Guided walk to the Waterfall trail, recreation

            6:30 pm             Dinner

            7:30 pm             Session I: Figs in Lambir

                                    Guide to the figs of Lambir Hills National Park – Rhett Harrison

                                    Figs and dispersal guilds in Lambir Hills National Park – Mike Shanahan

                                    Phenology of F. schwarzii & F. cereicarpa and the fig wasp interaction –

Rhett Harrison

                                    Minimum population size – model for F. schwarzii – Norio Yamamura

6^th

            7 am                 Breakfast

            8 am                 Session II: Fig Research in BCI, Panama

                                    Overview of Research at BCI – Allen Herre

                                    Ecology and reproductive strategies of non pollinating fig wasps – Stuart West

                                    Molecular natural history of fig wasps – Carlos Machado

            12:30 pm           Lunch

            2 pm                 Tree towers, canopy walkway and 8 ha Research plot –

                                    Kenta Tanaka, Yoko Inui, Hideo Arakawa, Aya Hatada

                                    Hemi-epiphytes in canopy plot area – Rhett Harrison & Mike Shanahan

            6:30 pm             Dinner

            7:30 pm             Session III: Fig Research in Taiwan

                                    Flowering phenology of F. septica and its influence on associated wasps in

                                    southern Taiwan – Shu-Hsuan & I-Fang Sun

                                    Seasonal fluctuations of Ficus microcarpa L. and pollinators in Taiwan –

                                    Chen, Ying-Ru Chou, Lien-Siang. and Wu W.  J.

7^th

            7 am                 Breakfast

            8 am                 52 ha long term ecological dynamics plot – Hatori

                                    Geocarpic figs in the 52 ha plot – Rhett Harrison

            12:30                Lunch

            2 pm                 Lupong Aji, secondary forest figs – Rhett Harrison

            7:30 pm             Night walk – Mike and Rhett

8^th         6:30 am                         Depart for Niah National Park

            7:30 am                         Breakfast at Niah Junction

            8 am                 Plank walk and Limestone caves

            2:30 pm             Depart for Lambir

            6:30 pm             Dinner

            7:30 pm             Session III: Fig Research in Taiwan (continued)

                                    The mutualism between Wiebesia pumilae and Ficus pumila var. pumila –

                                    Species interactions within the fig wasp community of Ficus microcarpa L. –

                                    Chen, Ying-Ru, Wu, W. J. and Chou, Lien-Siang.

                                    Phenology of Ficus erecta var. beecheyana in a warm forest at Guandaushi –

                                    Shy-Yu Tzeng, Fu-Yuan Lu, and Chern-Hsiung Ou

                                    Fig wasp community of Ficus irisana Elm. –

                                    Yen-Ling Chen

9^th

            7 am                 Breakfast

            8 am                 Wasp ecology, collecting, identification and molecular Research –

Steven Compton, Carlos Machado, Stuart West, Jun Yokoyama

            12:30 pm           Lunch

            2 pm                 Temperature physiology and chemical ecology – Sandra Patino & Rhett Harrison

            4 pm                 Session IV: Research from around the region

                                    Colonisation of Krakatau by figs – Steven Compton

                                    Influence of temperature on emission of volitiles from figs of F. aurata –

Sandra Patino

            6:30 pm             Dinner

10^th

6 am                 Observations at a fruiting fig – Mike Shanahan

            7 am                 Breakfast

            8 am                 Leisure

            12:30                Lunch

2 pm                 Discussion: Figs in Asia, where to next

            5 pm                 Longhouse visit, including dinner

11^th

            7 am                 Breakfast

            8 am                 Participants depart

Participants

Abstracts

Session I: Figs in Lambir Hills National Park

A guide to the figs of Lambir Hills National Park

Rhett Harrison

Lambir Hills National Park (LHNP) is one of the few preserved areas of primary lowland mixed dipterocarp forest in the region. In a 52 ha plot established in 1992/3 over 1200 species of tree (>1 cm DBH) were identified making it the most species rich forest known. Figs (Ficus) are one of the most speciose tropical genera with over 750 species world wide. The centre of diversity is S. E. Asia with over 350 species in Malaysiana alone. With 80 species recorded to date LHNP harbours the most diverse fig community of any comparable area and perhaps the largest collection of conspecific plants in any one habitat. This guide, with a key and descriptions to all the species so far found in the park, will provide a convenient reference and also illustrates the diversity of life forms, characters and ecology which enables so many species to coexist.

Fig dispersal guilds in Lambir Hills National Park

Mike Shanahan

Figs form a major dietary component of more frugivore species (many of which act as seed dispersers) than the fruit of any other tropical genus. However, no animal in an area eats the fruit of all sympatric fig species nor is any fig dispersed by all frugivores present. The diversity of Ficus found at Lambir lends itself to the testing of the hypothesis that discrete dispersal guilds exist within the genus. Observations of frugivory at fruiting Ficus trees (conducted in 1997 and ongoing) suggest the existence of at least 4 such guilds. Preliminary data are presented and discussed with respect to the factors determining guild membership, differential seed dispersal, the implications of disperser absence and the conservation of biodiversity in an anthropogenically disturbed habitat.

The phenology of Ficus schwarzii and F. cereicarpa and their interactions with fig wasps

Rhett Harrison & Tamiji Inoue

The phenology and wasp dynamics of Ficus schwarzii (16 male, 14 female, 1 immature) and F. cereicarpa (5 male, 12 female, 6 immature) two dioecious, cauliflorous figs (section Sycocarpus) has been followed at the “Swamp site” for four years, since November 1994. Leaf production was highly variable between censuses but both species renewed leaf in small quantities all year round, while F. schwarzii had more tendency to produce occasional flushes of new leaf. No correlation of leaf production and fruiting was observed. Fruit production on F. cereicarpa was very asynchronous with larger individuals supporting all crop stages simultaneously, however, syconia within each bunch were synchronous. Pollinator limitation leading to abortion of unpollinated syconia was common. Female syconia could wait up to 3 months for pollinators. Pollinators caught on sticky traps only appeared in large numbers when males within the group were releasing wasps. The only non pollinator (Sycophaga sp.) was sporadic at the site. F. schwarzii produced frequent highly synchronous crops. The interfloral and fruiting/wasp releasing stages were 67+/-2.9 days in females and 40+/-0.9 days in males but crops on female trees were overlapped. The mean period of female crops was 98+/-4.5 days and of males was 85+/-2.1 days. The period’s frequency Distribution, however, was right skewed for females and left skewed for males reflecting the tendency of female trees to delay crop production when the previous crop showed low pollination success, whereas male trees quickly produced a small “wasp breeding” crop and followed this with another crop so wasps leaving the “wasp breeding” crop would pollinate the following crop. A short drought in June 1996 led to a synchronising of trees within the group and a consequent peaks and crashes in wasp production. When there were no wasps being released at the site few or no wasps were caught at receptive trees. One non pollinating wasp (Apocryptophagus sp.) was able to oviposite in and prevent abortion of unpollinated syconia. Hence, it was less affected by crashes of the pollinators. However, the other two non pollinators (Philotrypesis sp. (inqualine) and Apocrypta sp.(parasitoid)) suffered crashes simultaneously with the pollinator. Apocyptophagus sp. was very occasionally caught at female trees and has been observed to attempt oviposition though the ovules were aborted. However, Philotrypesis sp. and Apocrypta sp. were never caught at female trees. Captures at traps on the wrong fig species were also extremely rare.

Persistence of fig wasp population and evolution of dioecy in fig: A simulation study

Tomoyuki Kameyama, Rhett Harrison and Norio Yamamura

The relationship between figs and their pollinator wasps is one of the most well known examples of the species-specific obligate mutualism. Because figs as a population bear fruits almost all the year round on which birds and mammals feed when other resources are scarce, they are thought to be keystone species in the tropic rain forest ecosystem. For figs to be available resources wasps have to pollinate figs, and this means wasps have to persist in a fig population for at least a certain period of time. In this paper we modelled this mutualistic system and examined the condition for the stable persistence of a wasp population. Major results are as follows. (1) When the average duration of the flowering interval of fig trees is short, even a small fig population can sustain a wasp population successfully. For example, a population whose average period of flowering cycle have halved can sustain a wasp population with the number of trees less than half of the original population. (2) The probability of successful survival of a wasp population is small when the variation of the interval periods of fig flowering is large. (3) The survival rate of a wasp population is large when the fig population size is large. (4) The wasp survival rate is large when figs can prolong their receptivity to wait for wasp if no wasps are available. (5) The wasp survival rate is predictable from the average pollination rate (the average proportion of that figs were entered by wasps) of fig population. (6) The persistence period of a wasp population increases exponentially with the number of fig trees. Based on the above results we propose a new hypothesis on the evolution of dioecy from monoecy in Ficus. When monoecious figs became adapted to the understorey environment rather than the canopy, the number of fig trees that are pollinated by a wasp population would have decreased because of the reduction in the wasp dispersal range. This led to the reduction in flowering cycles, favoring for evolution of dioecy because of the different lengths of the development of seed and wasps.

Session II: Fig studies in Panama

An over-view of fig studies in Panama

Edward Allen Herre

            I will discuss the advantages and often unexpected synergism resulting from collaborative studies, especially where work in apparently unrelated fields suddenly leads to new insights. This lesson cannot be over emphasised. I will discuss the work of others at BCI including John Nason’s work on pollen dispersal distances, Elizabeth Kalko and Carmi Korine’s work on dispersers and others. Finally, I will discuss a little about our work on the stability of the mutualism.

Studies on the non pollinating wasps on BCI

Stuart West

            In addition to their species specific pollinating wasp each fig harbours from one to thirty non pollinating species. The study of these non pollinating species is important both for (1) understanding their impact on the mutualism and (2) studying more general questions in community ecology and the evolution of reproductive strategies. We describe the life history of several New World non pollinating fig wasps, and examine their effect on the production of pollinator wasps and viable seeds. We use data from these species, and also species from Africa and Australia, to carry out comparative studies on how the sex ratio (proportion of males), male mating strategy (winged/wingless/dimorphic) and levels of virginity evolve in response to population structure.

Molecular Natural History of fig wasps

Carlos Machado

I will give an overview of molecular studies carried out in Panama. I will deal with four main subjects: 1. Global phylogeny of fig associated wasps (pollinators and non-pollinators): I will focus on the agreement with current morphology-based taxonomy and on the evolution of different ecological strategies. 2. Phylogeny of 15 genera of pollinating wasps: I will focus on the agreement of the molecular phylogenies with morphological phylogenies and with the classification of figs. 3. Molecular population genetics of one mitochondrial gene from Panamanian pollinating and non-pollinating wasps. 4. Tests of factors affecting rates of molecular evolution using fig wasps and nematodes.

Session III: Fig studies in Taiwan

Flowering phenology of Ficus septica and its influence on associated wasps in southern Taiwan

Shu-Hsuan Hu & I-Fang Sun

Ficus septica is a functional dioecious fig, which distributed commonly throughout lowland Taiwan. Pilot study indicated that there were one species of pollinating wasps (Ceratosolen jucundus) and three species of non-pollinating wasps (Philotrypesis sp., Apocryptophagus sp., Sycoryctes sp.) associated with F. septica. This project is trying to understand the intricate relationships among F. septica and associated pollinating as well as non-pollinating wasps. We especially addressed the following questions: 1) To what extent did flowering phenology of F. septica affect the dynamics of wasps populations? 2) How do female figs escape from being parasitized by pollinating and non-pollinating wasps? 3) The effects of foundress number on the sex ratio adjustment of pollinating wasps. 4) Do all three non-pollinating wasps depend on pollinating wasps for the survival of their species? The study site is located at the Kenting uplifted coral reef forest in southern Taiwan. Flowering phenology of twenty female and seventeen male trees we remonitored monthly from July 1996 to February 1998. Figs from different developmental stages were collected throughout the year, the numbers of pollinating wasps, non-pollinating wasps as well as sex ratios of each fig were counted and calculated. We also counted the number of seeds in each fig, measured the ovipositor length of pollinating and non-pollinating wasps. In addition, style length of both male and female flowers, thickness of B and C phases figs wall were also measured. The results showed: 1) F. septica flowered one to four crops a year, and have year-round flowering and fruiting patterns. Trees usually flowered synchronously within individuals but flowered asynchronously among individuals. In addition, when the study area have D phase trees existed, both female and male figs (syconia) have higher percentage of continuing development from B phase to C phase than being aborted. 2) The average ovipositor length of pollinating wasps is longer than the average style length of male flowers, but shorter than that of female flowers. Furthermore, the average ovipositor lengths of all three non-pollinating wasps are longer than the average thickness of fig walls. 3) In pollinator wasps, single foundress has the highest percentage in both male and female figs. However, average seed set of female fig is not significantly correlated with foundress number. The brood sex ratios of the pollinating and non-pollinating wasps are all female-biased. The sex ratio of these wasps fluctuated among seasons, and the magnitude of fluctuation was greater in the non-pollinating wasps than that of the pollinating wasps. 4) Our data suggested that one non-pollinating wasps, Apocryptophagus sp. can exist without the presence of pollinating wasps. However, the other two non-pollinating wasps are depending on pollinating wasps to maintain their populations.

Seasonal fluctuation of Ficus microcarpa L. and pollinators in Taiwan

Chen Ying-Ru and Wen-Jer Wu

            The most important problem between figs and pollinators is the time match between the fig crops and pollinators population. In order to understand the relationship of phenological matching, 35 Ficus microcarpa L. trees were investigated weekly on the campus of National Taiwan University in Taiwan during August 1992 to October 1997. The Research work includes phenological investigation and pollinator population dynamics. There were two flowering seasons (summerfall and winterspring) divided by two distinctive resting periods. There were few, to no syconia on the trees during these resting periods. The November-January resting period last more than a month. The April-May period was shorter. Crops in winterspring were longer than in summerfall.

            Eupristina verticillata the pollinator of Ficus microcarpa L. when leaving D-phase figs must immediately find B-phase figs bcause of their very short lifespan. The relative population size of pollinators was estimated by the fig entering rate (wings counted) and the number of pollinators in B-phase figs. Fig entering rates were low in winterspring but high in summerfall. The number of pollinators in one fig varied from one to six. The population dynamics of the wasps fluctuated dramatically within a year. However, it was quite steady between years. This implied a high mortality rate of the pollinator population.

The mutualism between Wiebesia pumilae (Hill) and Ficus pumila var. pumila

Jo-Chien Yao & Lien-Siang Chou

Wiebesia pumilae (Hill)(Hymenoptera: Agaonidae), a species-specific pollinator of Ficus pumila var. pumila L. (Moraceae), is the only wasp species associated with this scandent, gynodioecious fig. Several male syconia and galls were dissected once a week to observe the morphology and development of the immature wasps. The larva has 3 instars, and are grub-like, without eyes, legs, and other conspicuous structure. Their sex cannot be distinguished by external features until the late pupal stage. Investigating the phenology of fig trees showed that male and female syconia are not receptive at the same time. This may result in the female wasps could not develop the ability to distinguish the sex of syconia. The female wasps with and without pollen were introduced into receptive inflorescences respectively. On female tree, the syconia introduced with wasps without pollen were all aborted. But the syconium on male tree seemed to go on developing and did contain larva in each gall. This phenomenon may help to explore the importance of pollen in the development of wasp larvae, and the possible co-evolution process.

Species interactions within the fig wasp community on Ficus microcarpa L.

 Chen, Y. R.*, W. J. Wu* and L. S. Chou**

Although as a small size, figs of Ficus microcarpa L. hosted 6 genera, 18 species of non-pollinators, which oviposited by puncturing fig wall from outside. In order to investigate the relationships among this complicated community, we approach it by observing the time sequence of oviposition, dissecting D-phase figs and recording ovipositing floret types. Based on the observation of 35 trees from 1993-1997, we found Philotrypesis emeryi and Walkerella came to oviposit before pollinators, Sycoscapter, Sycoryctes, Philotrypesis sp. and Sycophila oviposited after pollinators. Analyzing the D-phase figs, pollinators occupied the long- pedicel florets and the seeds occupied the short-pedicel ones. We found a number of non-pollinators occupied the short-pedicel florets and affected the seed production, but Philotrypesis, Sycoryctes and Sycoscapter which have long ovipositors occupied the long-pedicel florets were the same ovipositing sites of pollinators. The result is inferred that the offsprings of Sycoscapter, Sycoryctes and Philotrypesis depend on pollinators, even eat the pollinators. Some D-phase figs only included Odontofroggattia or Walkerella. These genera could stimulate figs growing and staying on trees. Thus, they might not depend on pollinators. Now we are trying to bag figs before the entering of various wasps and hope to prove the effect among each species of wasps.

Phenology of Ficus erecta var. beecheyana in a warm forest at Guandaushi central Taiwan

Shy-Yu Tzeng, Fu-Yuan Lu, and Chern-Hsiung Ou

            This study carried out from Nov. 1995 to Apr. 1997 at Guandaushi forest station was aimed at the phenology of 71 mature trees of the dioecious fig species, Ficus erecta var. beecheyana. Trees were annually deciduous with a pronounced seasonal leaf drop and new growth. Male trees reached a peak level of syconia production 2 months consequent to onset of female syconium production, and individual trees bore 1 – 5 crops per year. The flowering cycle was evidently significantly affected by the fluctuation in weather. Female trees responded to more weather factors than males, which only responded to rainfall. The majority of D-phase male syconium and B-phase female and male syconium avoided the heaviest rainfall. Therefore, it maybe that this helps increase the survival of the very fragile pollinators, Blastophaga nipponica, and thereby aids pollination and maintenance of the family line. The flowering cycle of female syconium appears to be adapted to the seasonal climate and may improve the chance of good germination and seedling establishment at Guandaushi forest station.

The wasp community of Ficus irisana Elm.

Yen-Ling Chen

Ficus irisana Elm. had three species of fig wasps, Kradibia commuta, Philotrypesis sp. and Sycoscapter sp., which constantly appeared during the main flowering season that was during late June to December. K. commuta was the pollinator and could produce 199.0 offspring on average, but the number decreased when non pollinators were present. Based on field observation, bagging experiment and statistical analysis Philotrypesis sp. oviposited at interfloral phase after the pollinator and was an inquiline or parasitoid of the pollinator. However, Sycoscapter sp. was the latest to oviposit, also during the interfloral period and was a parasitoid of Philotrypesis sp.

Session IV: Fig studies from around the region

Influence of temperature on emission of volatiles from figs of Ficus aurata

Martine Hossaert-McKey, Laure Grison, Sandra Patino, Alice Edwards & John Grace

Volatile compounds emitted by receptive figs function as long and short distance attractants for their mutualistic pollinators. Receptive fig fragrance has been demonstrated to be a complex mixture of aromatic compounds. Variation of the quantity and composition of the emission was studied for F. aurata, a dioecious fig species of secondary forests in Brunei. Using the headspace technique, we collected volatiles emitted by receptive figs under different temperature regimes and performed a comparative analysis of these volatiles. Temperature has a strong influence on the intensity of the emission as well as on the relative abundances of the different compounds in the floral fragrance. The significance of our results will be discussed in relation to the ecology and reproductive biology of F. aurata.

Colonization of the Krakatau Islands by fig trees

Steve Compton

Life on the Krakatau islands was entirely destroyed by volcanic activity in 1883. The return of plants and animals to the islands has been well documented, and this talk will describe the history of fig tree colonization and its possible wider implications thanks to their putative 'keystone' role. The problems that might be generated by fig trees' specialist pollination syndrome do not appear to have materialised, and the talk will discuss why.

Back to Ficus home page       Back to figs and fig wasp home page     Back to fig wasp home page

Web authors Simon van Noort (Iziko South African Museum)

and Jean-Yves Rasplus (INRA, France)

Citation: van Noort, S. & Rasplus, JY. 2024. Figweb: figs and fig wasps of the world. URL: www.figweb.org(Accessed on <day-month-year>).

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