' J Ii i S1'.\4BIOSIS (2008) 45, 113-116 ©2008 Balaban. Philadelphia/Rehovot I 1 \'I I \ (\;, • I � r Structure of a fig wasp community: Temporal segregatio!l19f ll !"\. I !' 1' ) \(\ oviposition and larval diets �- \ ",, \,,'" , •••)l * 2 Rui Wu Wang 1 and Qi Zheng '''· . Jl.J'l I)' : .U'lil) - -,1 i. I d I r 1 ' Key Laboratory of Cellu tar and Molecular Evolution, Kunming Institute of Zoology, Chinese>Academy_of s_ci_en_c_es� q, / �unming, Yunnan 650223, China, Tel. +86-0871-5199178, Fax. +86-0871-5191823, Email. [email protected]·a�,c�; . _ , i , -Graduate School of Kunmmg Medical College, Kunmmg, Yunnan 650031, Chma · I • I I ) ' � I I'· (Received February 20, 2007; Accepted May 16, 2007) Abstract Observations were made on six fig wasp species on Ficus racemosa growing in the Xishuangbanna Tropical Botanic Garden. Yunnan Province, China. The oviposition sequence was determined for Apocryptophagus restacea, Apocry ta �p2. ! p Apocryptophagus mayri, Ceratosolen fusciceps and Apocrypta westwood. An experiment was carried116ttt in-�\vl'i 1'cfVi:a ch' r.11 .v species of non-pollinating wasp was separately introduced into bagged figs which prevented otherrspeJ:.io(.fr001 O'f.i?t>Sitingc J-./ 1 This showed that the non-pollinators Apocryptopha!JUS testacea_ and A. ma_ri are foragers (�alrt rrw�r �} �f11tlJ.fof�rp�e )'.)'Jd . y , fl• flowers, and that the other species were paras1to1ds or mqu1hnes. Add1t1onal evidence from naturaj\y unpoilina d .. l inflorescences suggested that A. agraensis is most likely a parasitoid of the pollinator, Cerato.tblen1fa�!ct,h.r;1 !tflf · atfJJ El�>' 1 and sp. are the parasitoids of and c e� . lliei.bL Apocrypta westwood Apocrypta Apocryptophagus mayri A. tesl3lirea1rt.espd tiv 1 "· ' results indicate that the fig wasps inhabiting F racemosa figs either utilize different develOP(Tlllt;ltal . �ag� Q 'hl'ljrCi , S,il8lfrt f! r resources (female flowers) or utilize different food resources. : '·1£ J£f1 I ·J1u1t 1o � .Jd mwr ,,.:.:,·, Fig, fig wasp, mutualism, coexistence, niche segregation Keywords: �b:,� :JdJ no flJlfll t!JrlJsi d111i 11 11ri� X !iri) fll ,?.J:u·tl 8 [ £ k /'.l'/11 < Jins Ir Jl .n.;btS'! o[J ; ·b1L I. Introduction diet of the fig wasp larvae1 iatHtstbllr�b�rueluc>)tilat� 1the ecological relationships a�ng« sped�s.J \V..4£�a�oqtried to Figs support both pollinator and non-pollinator wasps discern whether there is 11iche1, � ati� gteg n .am,<1>Jilgr lb and are completely dependent upon the pollinators for the different fig wasp species, on,/'icu,f.tQ��mosa,b,,1in1ordt;r. tl,_ dispersal of pollen In racemosa, non-pollinating species understand how they co®�t; nesu·l·t . F J Th-k!-.11�0-v).d ,frrom1 provide no service for figs (Yang et al, 2000; Wang et al., differences in larval diet (lihw.kirl.Sl�'fl�do(\).oril>�tol,l, 1992..; 2005a) or the pollinator wasps (Weiblen, 2002; Kerdelhue Cook and Power, 1996), or, the.; .fig::Jv,'.!a.sp.S! may l.Jtj'l•t2iel1th.e Apocrypta same resources in spatially or � p !™ly d c ys ., ' et al., 2000). Previous studies have considered ,t m (l> istin .t1wai fh spp. as parasitoids, while Apocryptophagus spp. as gall complexity of the fig-wasp communitie J.�bscu1( makers (Kerdelhue and Raspius, 1996; Kerdelhue et al., relationships (West and Herre, 1994; Kerdelhue et al., 2000) . The non-pollinators are demonstrated as gall makers 2000). Hence, we both make observations of fig wasr of figs or parasitoid pollinators, which are detrimental to oviposition in nature, and used controlled introductions tt fig-fig wasp mutualism, and therefore may affect the identify the larval diets of cG-existio� lwa5,p ,sp.ep�§S lo this process of fig-fig pollinator co-evolution (West et al ., 1996; way we attempted to identify the il)lpl!<;t·of each/sp�cies:.�J Bronstein, 2001; Harrison and Yamamura, 2003). Diverse the fig-fig pollinator interact�on. iriJ . o "r assemblages of non-po linators co-exist within a fig • l 'l ,ll) ( inflorescence and how these communities are structured remains enigmatic (West and Herre, 1994; Kerdelhue et al., 2. Materials and Methods 11 01 ')IHI , 2000; Profitt et al., 2007). ,., uqi·1'J 1 Jn·,.- To address these questions, we set out to identify the Materials t.. ,10 , Jn·,, t<rn L The fig wasp community was studied OrJ;Jf.i racemosa . _ * rhe author to whom correspondence should be sent. trees at the Xishuangbanoa Tropical Boitanjq Garden, Presenred ar the 7rh lnternarional Fig Wasp Symposium. July 23-26, 2006. Yunnan, China 114 R.W. WANG AND Q. ZHENG Table I. The number of the galls and wasp offspring (means (SD)) per inflorescence for controlled introductions of each non-pollinator ;pecies. separately, to bagged figs. pec1es Number of treated fruits Mature fruits collected Galls Wasps Apcryptophagus testacea 25 14 248.8 (64.3) 220.5 (62.8) Apcryptophagus mayri 20 9 57.7 (33.6) 39.9 (24.9) Aprryptophagus agraensis 15 0 0 0 Apocrypta sp. 21 0 0 0 Apocrypta westwoodi 23 0 0 0 Table 2. Comparison of bagged inflorescences (only oviposited by pollinators) (n=l 5) and natural inflorescences collected from the field (n=l 4) (means (SD)). Variable Treated fruits Natural fruits t value Diameter of fruits 2.91 (0.15) 2.81 (0.16) 1.872 ns Foundress 11.87 (9.43) 8.64 (3.97) 1.213 ns Galls 2444.87 (368.21) 3503.43 (384.20) 7.576** Seeds 1694.4 7 ( 452.84) 1794.64 (404.42) 0.627** **P<0.01. NS, P<0.05. Yunnan Province, China (21°41'N, 101°25'E) from introduced every two days, so that approximately 30 wasps November 2003 to December 2004. Ficus racemosa of each species were introduced per bag. We collected (section Sycomorus) is a monoecious fig distributed from aborted inflorescences and examined them for gall India to Australia (Corner, 1965). It forms a large tree that development. After the inflorescences matured, we can reach 30 m in height and grows in moist valleys or collected them, allowed the adult wasps to exit naturally, along rivers (Yang et al., 2000). This species bears very and preserved the wasps in 75% alcohol. We counted all of large numbers of fruits that are produced on branches and the galls and adult wasps from each fruit. Galls containing trunks rather than on the ends of twigs (caulif lorous). A dead wasps were opened to identify the wasps. survey of 2 l 8 fruits, in the Xishuangbanna Tropical Botanic To determine the impact of non-pollinators on the Garden Botanic garden, revealed that mature fruits of fig-fig pollinator interaction, we bagged 15 inflorescences F racemosa range in size from 2.92 cm to 5.4 cm. At this before the oviposition of any non-pollinating wasps. At the site, the pollinator of F racemosa is Ceratosolen fusciceps receptive stage we opened the bags to allow the pollinators (Agaonidae). In addition, there are five species of enter naturally. We then re-bagged them again unti I the figs non-pollinators, namely, Apocryptophagus testacea, matured. The mature inflorescences were collected and the A. mayri, A. agraensis (Caliimomidae), and Apocrypta contents were counted. We simultaneously collected 14 westwood and Apocrypta sp. (undescribed) (Pteromalidae) untreated inf lorescences from the same trees. (Weiblen, 2002; Wang et al., 2005a,b). The non-pollinator wasps insert their ovipositors through the wall of the Data analysis inflorescence. We used the Student's t test to test hypotheses relating Methods oviposition of non-pollinators to viable seed and pollinator offspring production, or other quantitative characteristics of We conducted direct observations of oviposttt0n the fruits of F racemosa. Correlation analysis (Pearson) behaviour. We selected 22 fruits on 2 trees for observation was used to analysis the quantitative relationship between and made observations three times per day (09:00-10:00, different fig wasps contained in the mature fruits of 14:00-15:00, and 17:00-18:00). Observations started from F racemosa. the first day that the non-pollinator wasps began to oviposit and continued for 30 days at which time no more wasps were observed ovipositing. 3. Results To identify the diet of non-pollinating fig wasps of F racemosa, we made controlled introductions. We enclosed Ovipos it ion sequence of non-pollinating wasps receptive fruits with organdy bags, and then separately Non-pollinating wasps of racemosa o iposited in the introduced the non-pollinator wasps in the same sequence F v we observed in the field. Twenty-five branches were treated, sequence: Apocryptophagus testacea, Apocrypta sp, and each bag contained 4-6 inflorescences. Wasps were Apocryptophagus mayri, pollinator wasp (C. fusciceps). FIG WASP OYIPOSITION AND LARVAL DIET \ 115 · P. testacea P.- mayn t -- 9 -- • - -- •• - I A. f\ (o I c. - '(51 C\/ · ..() .?.u -- I �I - - - - - I _ _ ,_.- -0 65.. I C. fusclceps-' ---+----------------seeds · . ) - · -·i··- . - ·- '"O "' ·' . ..:._ �·I :?5•· I · - Apocrypts sp P. agraensis I A. westwoodi 10 15 20 25 30 i ____ i i i Days potent al effect � potent al co ropet t on - on······ on ······ Figure I. The oviposition frequency and temporal sequence of fig �··�� potential parasitis111 wasps on Ficus racemosa at Xishuangbanna Tropical Botanic on ······ Garden. China over a period of 30 days. ( - • - Apocryptophagus testacea - · · \7 · - ··· Apocrypta Figure 2. Results of correlation analyses among populations of six sp2 ··· ·· 0 ··· ·· Apocryptophagus mayri II C. fusciceps - - fig wasp species, and the relationships between the fig wasp • - - Apocrypta westwoodi - - T - - Apocryptophagus species and viable seeds in the inflorescences of F racemosa at agraensis). Xishuangbanna Tropical Botanic Garden, China. The data of adult wasps is collected over one year (N=133) and the data of viable seeds and wasp offspring (galls) of pollinators is collected in November (N=26).