a. Fig and fig wasps
The study of the Ficus genus in Taiwan started in the beginning of the 20th century on Ficus pumila var. awkeotsang (高尾與一郎, 1917). Then during the 80s, numerous studies on this species have been done, mostly on the biochemistry of the jelly that can be produced from the dried fig of this species (Huang and Chen, 1979;
Huang et al., 1980; Lin et al., 1989; Liu et al., 1989; 1990) which is traditionally used for beverage. The reproductive system of this species has been also well studied in this period with the first studies having an ecological point of view and considering the mutualism between fig tree and wasp (Hu et al., 1986; Ho, 1987). The taxonomical work on Moraceae family and especially the fig tree species have been done at the end of the 80s (Liao, 1989). Recently the taxonomy has been reviewed by listing and establishing all the Taiwan Ficus species (Tzeng, 2004), this exhaustive work permits to have a clear view of the Ficus flora and their distribution on the island. More recently F. fistulosa var. benguetensis has been put as species: F.
benguetensis (Berg, 2011).
However, the ecological approach on the interspecific mutualism between figs and fig wasps did not get attention until early 1990s. Then number of studies on mutualistic pairs increased quickly in the last two decades. The studied species were:
F. ampelas (Chang, 2003), F. aurantiacea var. parvifolia (Chou and Yeh, 1995), F.
38
erecta var. beecheyana (Wu, 1996; Tzeng et al., 2003), F. formosana (Tseng, 1999), F. irisana (Chen, 1998), F. microcarpa (Hsieh, 1992; Chen, 1994; Chen et al., 2004;
Yang, 2011), F. pumila (Ho, 1991; 1997; Yao, 1998), F. septica (Ho, 2009) and F.
tinctoria subsp. swinhoei (Huang, 2007). Most of these works are master thesis which had not been published in peer reviews. Nevertheless they give a good basis for further studies. Among these studies, the data on the monoecious F. microcarpa is strongly congruent in the phenological pattern: The Taipei fig tree population bore figs almost constantly along the year but with one short period with no fig in late fall (Chen et al., 2004) and few figs during the winter season. This type of phenology was also the one of F. ampelas in Chiayi city: the trees bore figs all along the year but for this dioecious species the male fig production was slightly earlier than the female fig production (Chang, 2003). On the other hand, F. erecta was studied in the Taiwan mountain areas between 500 and 800m altitude (e.g. Yangmingshan at northern Taiwan by Wu, 1996; Huesun forest of Central Taiwan by Tzeng et al., 2003; 2006).
Both studies showed that this species was growing figs with similar pattern, indeed two productions peaks per year have been observed with a very low level of fig presence between the peaks. Separated by about 130km, the climate showed some differences between these two sites, Yangmingshan was colder than the Huesun forest:
the winter minimum monthly temperatures were below 10°C in Yangmingshan and only below 15°C at the Huesun forest (Wu, 1996; Tzeng et al., 2003; 2006).
Furthermore Ficus formosana, also studied at the Huesun forest, had very similar phenology with two peaks of fig production (Tseng, 1999). F. erecta and F.
formosana belong both to the subgenus Ficus in the Ficus classification and live in the same mountainous zone, but staying in different forest types. F. formosana is
39
more linked to the understorey environment in gymnosperm forests and F. erecta is often observed at the edge of the vegetation (Tzeng, 2004).
Nevertheless the phenologies of these two species were clearly distinct from the growth pattern of F. microcarpa. Interestingly, the phenology of F. irisana at the Huesun forest was intermediate between the pattern of F. microcarpa and the one of F. erecta and F. formosana (Chen, 1998) but closer to the closely related species, F.
ampelas. Indeed the trees are producing figs all along the year but pollinated figs were not observed all along the year indicating a probable low pollination rate. For this species, male and female trees seemed to have more similar phenologies than the previous species. Finally the two remaining species, F. pumila and F. tinctoria showed continuous phenology at the population level (Ho, 1991; Yao, 1998; Huang, 2007) and very strong asynchrony between individuals in the case of F. pumila (Ho, 1991; Yao, 1998). Similarly, F. septica in Taichung showed strong asynchrony between trees (Ho, 2009). This study showed that the trees in the central part of Taiwan were bearing figs along the year: indeed they started crops when the previous one was close to the end of the fig maturation. Finally, the phenological studies from Taiwan covered a great panel of environments and showed the diversity of the fig phenology through the different groups of the Ficus classification and climatic affinities.
Furthermore the last phonological work on F. microcarpa in Taipei provided important data on the pollinating wasp population size (Yang, 2011). During this study, the number of female pollinating wasps inside the figs (foundress) has been counted. The vast majority has one or two foundresses inside and once 12 wasps have been observed in one fig. This data has been used to infer the total population of female wasps living around the studied group of Ficus microcarpa trees in Taipei
40
(Yang, 2011). Indeed three calculations may be used to have a clearer view of the wasp population level. Firstly the occupation rate, the crowdedness index, the average foundress number and the estimated pollinator population size (Yang, 2011). Thus, the highest population level is generated by the spring emergences, every year the low population level in winter is quickly recovered. The weekly survey on the fig and wasp populations permitted to infer that the population level is strongly fluctuating within a year (Yang, 2011).
These studies established good foundation for further research on aspects of the fig tree biology such as the mutualism with the biology and the identification of the pollinating Agaonidae wasps (sensu Cruaud et al., 2010) or for the fig wasp population survival. From the aspects of fig pollinator wasps (Hymenoptera:
Agaonidae), revision and the description of wasp species of eight genera occurred in Taiwan have been done in 1997 (Chen and Chou). Besides 17 species recorded, another seven species have been newly described: Blastophaga esquirolianae (pollinator of Ficus esquiroliana), B. taiwanensis (F. formosana), B. pedunculosae (F.
pedunculosa), B. tannoensis (F. tannoensis), Ceratosolen wui (F. benguetensis) and Platyscapa hsui (F. caulocarpa). Moreover on Ficus septica, two pollinator species have been observed in the South of the Taiwan Island (Lin et al., 2011). One of the agaonid wasp species is strictly limited to the extreme south of the island with an extension to the Southwest plain. The southern species is particularly abundant in Orchid Island (Lin et al., 2011). Furthermore, the genetic study on these pollinators has shown that the differentiation of the different fig wasp populations on the Taiwan Island was weak leading to conclude that the gene flow is strong (Lin et al., 2008). On the other, the genetic analysis on Ficus pumila and F. pumila var. awkeotsang showed that the fig trees from the Taiwan Island are weakly differentiation (Lee, 2008).
41
Moreover the two varieties were significantly less differentiated than their common pollinator Wiebesia pumilae. Anatomical differences have been recently highlighted on the pollinating wasps occurring in the two varieties of F. pumila (Jiang, 2011).
And this species was genetically indistinguishable from the pollinator of F.
sarmentosa, W. callida (Lee, 2008).
Besides pollinating fig wasp, some figs host non-pollinating fig wasps (NPFW).
These parasites are mostly not described for the Taiwan species nevertheless the studies showed the number of NPFW species occurring for each fig species (Table 1).
Most of the species from these studies had been identified at the genus level without been described. On the other hand, the hymenopteran parasitic community of F.
microcarpa, which is a monoecious species, has been described (Chen et al., 1999).
The NPFW community of F. microcarpa gathers more species than the others dioecious studied species. Ten species have been described for F. microcarpa and the community seems to be similar with the monoecious F. subpisocarpa (Bain, Pers.
Obs.). Moreover experiments, consisting in sealing the ostiole of the F. microcarpa figs (Chen et al., 2001), showed that two species were undoubtedly gallmakers (Odontofroggatia sp. and Walkerella kurandensis).
42
Table 1: Ficus species and their associated wasp species. The pollinating wasp species references correspond to their description while the non-pollinating wasp species references are about the observations of these species. When a species has not been observed or described from Taiwan, the area/country has been added to the name (Area abbreviations: Au, Australia; Ch, China; J, Japan; Mel, Melanesia; Ph, Philippines; Sol, Solomon Islands and W for worldwide). The wasp species from the genus Blastophaga subgenus Valisia have been raised at the genus level (Valisia gen.
nov.) as described in Cruaud et al., 2010. Furthermore the wasp species of the genus Liporrhopalum have been put in synonymy with the genus Krabidia (Cruaud et al., 2010).
Ficus Pollinating wasp Non-pollinating wasp References
Subgenus Urostigma
Platyscapa hsui Chen and Chou, 1997
Platyscapa fischeri (J, Ph) Yokohama and
Iwatsuki, 1998 Otitesella clarae (J, Ph)
Camarothorax sp2 (J)
F . m ic ro c a rp a
Eupristina verticillata Waterston, 1921
Bruchophagus sensoriae Chen et al., 1999 Sycophila petiolata
Walkerella yashiroi (J, USA) Yokohama and Iwatsuki, 1998;
Beardsley, 1998 F. benjamina
var. bracteata
Eupristina koninsbergeri Grandi, 1916
Philotrypesis distallatoria Chou and Wong, 1997
43 Table 1: continued
Ficus Pollinating wasp Non-pollinating wasp References
F. pubinervis Dolichoris valentine Grandi, 1916
Parasitic fauna unknown
F. nervosa Dolichoris nervosae nervosae Hill, 1967 Parasitic fauna unknown
Subgenus Ficus F. pedunculosa var.
mearnsii
Blastophaga peducunlosae Chen and Chou, 1997 Parasitic fauna unknown
F. erecta var.
beecheyana
Blastophaga nipponica Grandi, 1921
Sycoscapter inubiae Tzeng, 1997; Tzeng et al., 2006
Sycoscapter sp.
F. formosana
Blastophaga taiwanensis Chen and Chou, 1997 Sycoscapter spA Tzeng et al., 2008 Sycoscapter spB
F. tannoensis Blastophaga tannoensis Chen and Chou, 1997 Parasitic fauna unknown
F. vaccinoides Blastophaga yeni Chen and Chou, 1997
Parasitic fauna unknown F. esquiroliana
Valisia esquirolianae Chen and Chou, 1997;
Cruaud et al., 2010 Parasitic fauna unknown
F. ruficaulis
Valisia filippina Wiebes, 1993; Cruaud et al.,
2010 Parasitic fauna unknown Subgenus Synoecia
F. sarmentosa Wiebesia callida Grandi, 1927a
Parasitic fauna unknown F. pumila var.
pumila
Wiebesia pumilae Hill, 1967
Wiebesia sp. Lee, 2008; Jiang, 2011
No parasitic wasp in Taiwan F. pumila var.
awkeotsang
Wiebesia pumilae Hill, 1967
Wiebesia sp. Lee, 2008; Jiang, 2011
No parasitic wasp in Taiwan
F. trichocarpa Wiebesia vechti Wiebes, 1993
Parasitic fauna unknown F. aurantiacea var.
parvifolia
Wiebesia contubernalis Grandi, 1927b
Sycoscapter sp. Chou and Yeh, 1995 Subgenus Sycidium
F. cumingii Krabidia panchoi Wiebes, 1993
Parasitic fauna unknown
44 Table 1: continued
Ficus Pollinating wasp Non-pollinating wasp References
F. irisana
Krabidia gibbosae Hill, 1967; Cruaud et al., 2010
Krabidia philippinensis Hill, 1969; Cruaud et al., 2010
F. benguetensis Ceratosolen wui Chen and Chou, 1997
Parasitic fauna unknown
F. septica
Ceratosolen bisulcatus Mayr, 1885
Ceratosolen bisulcatusjucundus Grandi, 1927b
Ceratosolen sp. Lin et al., 2011
Philotrypesis sp1. Ho, 2009 Philotrypesis sp2.
Sycoscapter sp.
Unknown genus
45 b. Fig eaters
In the world, the figs are consumed by an important number of vertebrate species (Shanahan et al., 2001). In Taiwan, most of the documented observations of fig consumption has been done in the South of the island on five species. F. ampelas, F.
caulocarpa and F. irisana are consumed by numerous birds, the Formosan macaque and the Formosan squirrel (Tamura et al., 1989; Song, 2006), F. septica was also consumed in southern Taiwan by the Formosan squirrel (Tamura et al., 1989). This fig species is mainly dispersed by fruit bat species (see appendix of Shanahan et al., 2001) which one species is present in the South of the island (Hsu, 1997). The consumed fig species have been observed frequently in the studied sites in the South of Taiwan and Orchid Island (Chao et al., 2007; 2010). In the northeastern part of the Taiwan Island, the Formosan rock macaques have been observed eating numerous figs species (Su and Lee, 2001). In the same zone, the diet of the Indian giant flying squirrel has been studied and about 15% of their diet came from the figs of F.
subpisocarpa (Kuo and Lee, 2003). Finally, unidentified figs were the only vegetal part of three carnivore species (Chuang and Lee, 1997).
c. Non-Hymenopteran parasites
Moth and butterfly larvae (Lepidoptera) have been described feeding on Ficus leaves of figs (Wang and Emmel, 1990; Lin, 2005). Another group of insects from the Thripidae family (Thysanoptera) has been documented parasiting leaves of F. septica (Chen and Lu, 1994). Moreover, numerous homopteran species have been observed feeding on branches and aerial roots of numerous fig species (Bain, pers. obs).
46 d. Conclusion
This review presents the biological and ecological studies on Ficus in Taiwan. These works present a very good definition of the fig tree communities for numerous Ficus species and at different levels: from the pollination mutualism to the seed dispersion.
Although many of them are mainly descriptive these fruitful results have built the basis for further investigations to understand more deeply such as the population dynamics and genetic fluxes (Lin et al., 2008; 2011). Inferences in population genetics cannot be done without a good coverage of the biological features of the studied species. The former studies added to the work done on this thesis provided a solid basis to further studies on evolutionary mechanisms involved on Taiwan Island in the fig-fig wasp mutualism.
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