Finding hidden females in a crowd: Mate recognition in fig wasps

Multi-species mating aggregations are crowded environments within which mate recognition must occur. Mating aggregations of fig wasps can consist of thousands of individuals of many species that attain sexual maturity simultaneously and mate in the same microenvironment, i.e, in syntopy, within the close confines of an enclosed globular inflorescence called a syconium - a system that has many signalling constraints such as darkness and crowding. All wasps develop within individual galled flowers. Since mating mostly occurs when females are still confined within their galls,, male wasps have the additional burden of detecting conspecific females that are ``hidden'' behind barriers consisting of gall walls. In Ficus racemosa, we investigated signals used by pollinating fig wasp males to differentiate conspecific females from females of other syntopic fig wasp species. Male Ceratosolen fusciceps could detect conspecific females using cues from galls containing females, empty galls, as well as cues from gall volatiles and gall surface hydrocarbons. In many figs, syconia are pollinated by single foundress wasps, leading to high levels of wasp inbreeding due to sibmating. In F. racemosa, as most syconia contain many foundresses, we expected male pollinators to prefer non-sib females to female siblings to reduce inbreeding. We used galls containing females from non-natal figs as a proxy for non-sibs and those from natal figs as a proxy for sibling females. We found that males preferred galls of female pollinators from natal figs. However, males were undecided when given a choice between galls containing non-pollinator females from natal syconia and pollinator females from non-natal syconia, suggesting olfactory imprinting by the natal syconial environment. (C) 2013 Elsevier Masson SAS. All rights reserved.

Parasites exert conflicting selection pressures to affect reproductive asynchrony of their host plant in an obligate pollination mutualism

1. Plant reproductive phenology is generally viewed as an individual's strategy to maximize gamete exchange and propagule dispersal and is often considered largely dependent on patterns of floral initiation. Reproductive phenology, however, can be affected by proximate responses to pollinators, parasites and herbivores which could influence floral longevity or fruit development time. 2. We examined the influence of insect interactants on within-plant reproductive phenology in the fig-fig wasp nursery pollination mutualism in Ficus racemosa (Moraceae). Most figs support a wasp community comprised of a mutualistic pollinator, with several host-plant-specific non-pollinating herbivorous gallers and parasitoids. These wasps reproduce within enclosed inflorescences called syconia, which develop into fruit after pollination. While different wasp species oviposit into syconia at varying times during its ontogeny, all wasp progeny are constrained to exit syconia simultaneously just prior to fruit ripening. Developing larvae of early-ovipositing wasps may hasten syconium ontogeny through formation of earlier and larger nutrient sinks, whereas larvae of late-arriving parasites may lengthen syconium ontogeny to complete their development successfully. Seeds are also important nutrient sinks. The number of seeds and the type and number of developing wasps may therefore be expected to influence syconium development times, thereby affecting the reproductive synchrony of syconia on a plant. 3. Observations on naturally pollinated and parasitized syconia indicated that their seed and wasp content affected syconium development time. Experimental manipulations of syconia to produce only seeds or various combinations of wasps confirmed this finding. Early-ovipositing galler progeny reduced syconium development times, while gallers ovipositing concurrently with pollinators had no effect on syconium development. Late-ovipositing parasitoid progeny, the presence of only seeds within the syconium, or delayed pollination increased syconium development time. The differential development of syconia, which was influenced by mutualistic or parasitic progeny, accordingly contributed to within-tree reproductive asynchrony. 4. Synthesis. Individual reproductive units in fig trees called syconia, which also function as brood sites for pollinating and parasitic fig wasps, have plastic development durations dependent on pollination timing and species of wasps developing within them. Syconium development times are a likely compromise between conflicting demands from developing seeds and different wasp species.

High Temperatures Result in Smaller Nurseries which Lower Reproduction of Pollinators and Parasites in a Brood Site Pollination Mutualism

In a nursery pollination mutualism, we asked whether environmental factors affected reproduction of mutualistic pollinators, non-mutualistic parasites and seed production via seasonal changes in plant traits such as inflorescence size and within-tree reproductive phenology. We examined seasonal variation in reproduction in Ficus racemosa community members that utilise enclosed inflorescences called syconia as nurseries. Temperature, relative humidity and rainfall defined four seasons: winter; hot days, cold nights; summer and wet seasons. Syconium volumes were highest in winter and lowest in summer, and affected syconium contents positively across all seasons. Greater transpiration from the nurseries was possibly responsible for smaller syconia in summer. The 3-5 degrees C increase in mean temperatures between the cooler seasons and summer reduced fig wasp reproduction and increased seed production nearly two-fold. Yet, seed and pollinator progeny production were never negatively related in any season confirming the mutualistic fig-pollinator association across seasons. Non-pollinator parasites affected seed production negatively in some seasons, but had a surprisingly positive relationship with pollinators in most seasons. While within-tree reproductive phenology did not vary across seasons, its effect on syconium inhabitants varied with season. In all seasons, within-tree reproductive asynchrony affected parasite reproduction negatively, whereas it had a positive effect on pollinator reproduction in winter and a negative effect in summer. Seasonally variable syconium volumes probably caused the differential effect of within-tree reproductive phenology on pollinator reproduction. Within-tree reproductive asynchrony itself was positively affected by intra-tree variation in syconium contents and volume, creating a unique feedback loop which varied across seasons. Therefore, nursery size affected fig wasp reproduction, seed production and within-tree reproductive phenology via the feedback cycle in this system. Climatic factors affecting plant reproductive traits cause biotic relationships between plants, mutualists and parasites to vary seasonally and must be accorded greater attention, especially in the context of climate change.

A coat of many scents: Cuticular hydrocarbons in multitrophic interactions of fig wasps with ants

The fig fig wasp system of Ficus racemosa constitutes an assemblage of galler and parasitoid wasps in which tritrophic interactions occur. Since predatory ants (Oecophylla smaragdina and Technomyrmex albipes) or mostly trophobiont-tending ants (Myrmicaria brunnea) were previously shown to differentially use volatile organic compounds (VOCs) from figs as proximal cues for predation on fig wasps, we examined the response of these ants to the cuticular hydrocarbons (CHCs) of the wasps. CHC signatures of gallers were distinguished from those of parasitoids by the methyl-branched alkanes 5-methylpentacosane and 13-methylnonacosane which characterised trophic group membership. CHC profiles of wasp predator and wasp prey were congruent suggesting that parasitoids acquire CHCs from their prey; the CHC composition of the parasitoid Apocrypta sp 2 clustered with that of its galler host Apocryptophagus fusca, while the CHC profile of the parasitoid Apocryptophagus agraensis clustered with its galler prey, the fig pollinator Ceratosolen fusciceps. In behavioural assays with ants, parasitoid CHC extracts evoked greater response in all ant species compared to galler extracts, suggesting that parasitoid CHC extracts contain more elicitors of ant behaviour than those of plant feeders. CHCs of some wasp species did not elicit significant responses even in predatory ants, suggesting chemical camouflage. Contrary to earlier studies which demonstrated that predatory ants learned to associate wasp prey with specific fig VOCs, prior exposure to fig wasp CHCs did not affect the reaction of any ant species to these CHCs. (C) 2015 Elsevier Masson SAS. All rights reserved.

Plant reproductive traits mediate tritrophic feedback effects within an obligate brood-site pollination mutualism

Plants, herbivores and parasitoids affect each other directly and indirectly; however, feedback effects mediated by host plant traits have rarely been demonstrated in these tritrophic interactions. Brood-site pollination mutualisms (e.g. those involving figs and fig wasps) represent specialised tritrophic communities where the progeny of mutualistic pollinators and of non-mutualistic gallers (both herbivores) together with that of their parasitoids develop within enclosed inflorescences called syconia (hence termed brood-sites or microcosms). Plant reproductive phenology (which affects temporal brood-site availability) and inflorescence size (representing brood-site size) are plant traits that could affect reproductive resources, and hence relationships between trees, pollinators and non-pollinating wasps. Analysing wasp and seed contents of syconia, we examined direct, indirect, trophic and non-trophic relationships within the interaction web of the fig-fig wasp community of Ficus racemosa in the context of brood site size and availability. We demonstrate that in addition to direct resource competition and predator-prey (host-parasitoid) interactions, these communities display exploitative or apparent competition and trait-mediated indirect interactions. Inflorescence size and plant reproductive phenology impacted plant-herbivore and plant-parasitoid associations. These plant traits also influenced herbivore-herbivore and herbivore-parasitoid relationships via indirect effects. Most importantly, we found a reciprocal effect between within-tree reproductive asynchrony and fig wasp progeny abundances per syconium that drives a positive feedback cycle within the system. The impact of a multitrophic feedback cycle within a community built around a mutualistic core highlights the need for a holistic view of plant-herbivore-parasitoid interactions in the community ecology of mutualisms.

Estado actual del bosque de galería de la parte alta del rio Santa Elena, sector norte de la Universidad Nacional Agraria, Managua

~vii~ RESUMEN La presente investigación se realizó en el bosque de galería de la parte alta del río Santa Elena, de la Universidad Nacional Agraria, donde se identificó el estado actual y la estructura horizontal y vertical, la composición florística en el estrato arbóreo del río, además, se determinó la diversidad a través del índice ecológico de Shannon-Weiner, por lo cual se realizó un inventario al cien por ciento (100 %). En cuanto a la estructura horizontal, se encontró el mayor número de individuos correspondiendo a la categoría 10 a 19.9 cm, la que presento menores cantidades de árboles fue la categoría mayor de 90cm, la dominancia más alta fue para Ficus trigonata (Chilamate). En cuanto al índice de valor de importancia (IVI), los valores más altos, le corresponden a las especies, Ficus trigonata (Chilamate), seguido por Guazuma ulmifolia (Guácimo de ternero) y Pithecellobium dulce (Espino de playa), con respecto a la abundancia específica, Ficus trigonata (Chilamate) y Guazuma ulmifolia (Guácimo de ternero), son las más abundantes, obteniéndose el mayor valor en la ribera 4, similares resultados para la ribera 2, ribera 3, y los que presentaron menor diversidad fueron ribera 1 e Isla, existiendo las mayores abundancias para la ribera 4 y la ribera 2, sobresaliendo las especies de: Ficus trigonata (Chilamate), Inga vera (Guavillo) y Terminalia catappa (Almendra), las cuales se encuentran en todas las riberas, y otras presentes en tres y dos riberas, no obstante, las actividades que se realizan en el área y los en sayos de agricultura han permitido que disminuya la cobertura vegetal y dando espacio a nuevas especies arvense que se desarrollen llegando incluso a las orillas del río, por esta razón se proponen soluciones para conservar el recurso bosque de galería.

Comportamiento de nopal (Opuntia ficusindica L.) en asocio con seis cultivos prehispánicos, Ecolotes Ave Maria, Las Esquinas, Carazo 2013

El nopal (Opuntia ficus-indica L.), familia de las Cactáceas, es originario de América. Es una planta que se adapta muy bien en diferentes condiciones climáticas, se desarrolla exitosamente en zonas donde llueve muy poco; también tiene una gran importancia por sus diferentes usos que se le conocen. Para conocer de su comportamiento como cultivo asociado, se estableció un ensayo en la finca Ecolote Ave María, ubicada en Las Esquinas, Carazo, en el km 37 ½ carretera Managua – San Marcos, Carazo, en la época de primera 2013. Se utilizó un diseño de bloques completos al azar (BCA), con cuatro repeticiones y seis tratamientos: pipián cuarenteño (Cucúrbita angyrosperma J. C. Huber), amaranto (Amaranthus caudatus L.), chan (Hyptis suaveolens L.), frijol blanco (Phaseolus acutifolius, Gray), yuca (Manihot esculenta Crantz) y batata (Ipomoea batata L.), con el objetivo de conocer con cuál de estos cultivos en asocio se obtienen mejores rendimientos de cladodios de nopal. El análisis de los datos obtenidos del estudio reflejan que se obtuvo una supervivencia del 100 % en todo los tratamientos, el mejor número de brotes por planta fue de 2, el mejor ancho de cladodios (3.63 cm) y el mejor rendimiento fue de (2 386.5 kg/ha), en asocio con amaranto; el mejor largo de cladodios (7.65 cm) fue con el cultivo de Yuca, respecto al comportamiento de los cultivos prehispánicos, batata con una altura de 172.8 cm ha sido el más exitoso, seguido por el chan con 131.42 cm de largo de guía y la yuca con 111.3 cm de altura, los cultivos de amaranto, frijol blanco y pipián cuarenteño no lograron adaptarse a las condiciones climáticas del ensayo.

Composição florística e estrutura da comunidade arbustivo-arbórea de um trecho de floresta submontana na vertente sudeste do Parque Estadual da Ilha Grande, Angra dos Reis / RJ

O estudo objetivou avaliar a composição florística e estrutural dos componentes arbustivo-arbóreo da Floresta Ombrófila Densa submontana em diferentes estágios de regeneração natural, na vertente sudeste do Parque Estadual da Ilha Grande/RJ. Para o inventário florístico foram realizadas coletas assistemáticas em diferentes trechos nessa vertente. A complementação da lista de espécies foi feita a partir, da consulta às exsicatas dos herbários do Rio de Janeiro (FCAB, GUA, HB, HRJ, R, RB, RBR, RFA, RFFP e RUSU) e do inventário fitossociológico. Foi verificado o status de conservação das espécies inventariadas para a Flora Brasileira. Para o inventário fitossociológico foram estabelecidas 34 parcelas amostrais, totalizando 1,02 ha de área amostrada. Todos os indivíduos arbustivo-arbóreos com DAP ≥ 5 cm foram registrados e, após identificação, foram depositados no Herbário da Universidade do Estado do Rio de Janeiro (HRJ). O pacote estatístico FITOPAC 2.1. foi utilizado para a análise dos dados. A similaridade entre o remanescente investigado neste estudo e as outras quatorze áreas distintas do Rio de Janeiro, da própria Ilha Grande ou não, foi avaliada, utilizando-se o coeficiente de Similaridade de Sorensen; pelo critério de agrupamento por ligação média não ponderada (UPGMA) e pelo método de autorreamostragem para a estrutura de grupos; utilizados os programas PAST v1.34 e Multiv 2.4. A partir do levantamento em herbários e dos inventários florístico e fitossociológico realizados neste trabalho, foram analisados 3.470 registros, sendo 1.778 do levantamento de herbários, 1.536 do levantamento fitossociológico e 156 do inventário florístico. Esses registros corresponderam a 606 espécies ou morfo-espécies de Angiospermas e uma de Pteridófita. Os resultados obtidos revelaram a existência de 22 espécies ameaçadas de extinção para a Flora do Brasil. Dentre, as quais, sete são exclusivas da amostragem fitossociológica: Abarema cochliacarpos (Gomes) Barneby & J.W. Grimes, Chrysophyllum flexuosum Mart., Ficus pulchella Schott ex Spreng., Macrotorus utriculatus Perkins, Myrceugenia myrcioides (Cambess.) O.Berg, Rudgea interrupta Benth e Urbanodendron bahiense (Meisn.) Rohwer. No estudo fitossociológico, inventariou-se 1.536 indivíduos de 217 espécies, subordinadas a 53 famílias. O índice de diversidade de Shannon (H) calculado foi de 4,702 nats/ind e equabilidade (J) de 0,874. As 10 famílias com maior riqueza foram: Myrtaceae (31 spp.), Rubiaceae (21), Fabaceae (17), Lauraceae (12), Euphorbiaceae (11), Monimiaceae (8), Melastomataceae (7), Sapindaceae (7), Sapotaceae (6) e Annonaceae (6). Os 10 maiores Valores de Importância das espécies foram para Chrysophyllum flexuosum (3,43%), Lamanonia ternata Vell. (3,40%), Hyeronima alchorneoides Allemão (2,83%), Actinostemon verticillatus (Klotzsch) Baill. (2,55%), Psychotria brasiliensis Vell. (2,55%), Eriotheca pentaphylla (Vell.) A. Robyns (2,28%), Guatteria australis A. St.-Hil. (2,12%), Mabea brasiliensis Müll. Arg. (2,04%), Miconia prasina (Sw.) DC. (1,89%) e Rustia formosa (Cham. & Schltdl. ex DC.) Klotzsch (1,82%). Amostraram-se 27% de espécies representadas por apenas um indivíduo. As análises florísticas avaliadas a partir do Índice de Similaridade de Sorensen indicaram como principais variáveis para a formação dos blocos, os diferentes valores de diversidade para as áreas e a distribuição fitogeográfica das espécies. Os resultados obtidos junto aos dados dos grupos ecológicos, para os indivíduos da fitossociologia, indicaram maior percentual de indivíduos secundários tardios amostrados. Conclui-se que a área de estudo é uma floresta secundária em estágio intermediário de regeneração, com grande riqueza de espécies, muitas das quais de relevante importância ecológica.

Differences in diet of Atlantic bluefin tuna (Thunnus thynnus) at five seasonal feeding grounds on the New England continental shelf

The stomachs of 819 Atlantic bluefin tuna (Thunnus thynnus) sampled from 1988 to 1992 were analyzed to compare dietary differences among five feeding grounds on the New England continental shelf (Jeffreys Ledge, Stellwagen Bank, Cape Cod Bay, Great South Channel, and South of Martha’s Vineyard) where a majority of the U.S. Atlantic commercial catch occurs. Spatial variation in prey was expected to be a primary influence on bluefin tuna distribution during seasonal feeding migrations. Sand lance (Ammodytes spp.), Atlantic herring (Clupea harengus), Atlantic mackerel (Scomber scombrus), squid (Cephalopoda), and bluefish (Pomatomus saltatrix) were the top prey in terms of frequency of occurrence and percent prey weight for all areas combined. Prey composition was uncorrelated between study areas, with the exception of a significant association between Stellwagen Bank and Great South Channel, where sand lance and Atlantic herring occurred most frequently. Mean stomach-contents biomass varied significantly for all study areas, except for Great South Channel and Cape Cod Bay. Jeffreys Ledge had the highest mean stomach-contents biomass (2.0 kg) among the four Gulf of Maine areas and Cape Cod Bay had the lowest (0.4 kg). Diet at four of the five areas was dominated by one or two small pelagic prey and several other pelagic prey made minor contributions. In contrast, half of the prey species found in the Cape Cod Bay diet were demersal species, including the frequent occurrence of the sessile fig sponge (Suberites ficus). Prey size selection was consistent over a wide range of bluefin length. Age 2–4 sand lance and Atlantic herring and age 0–1 squid and Atlantic mackerel were common prey for all sizes of bluefin tuna. This is the first study to compare diet composition of western Atlantic bluefin tuna among discrete feeding grounds during their seasonal migration to the New England continental shelf and to evaluate predator-prey size relationships. Previous studies have not found a common occurrence of demersal species or a pre-dominance of Atlantic herring in the diet of bluefin tuna.

聚果榕两种非传粉小蜂产卵与果实脱落的关系

下载PDF阅读器在聚果榕与其传粉榕小蜂Ceratosolen fusciceps组成的互利共生系统中,与传粉榕小蜂共存的还有榕树的寄生性小蜂Platyneura testacea和Platyneura mayri.这些寄生性小蜂由于不能给榕树带来任何收益而只是利用榕树的种子或与传粉小蜂竞争植物的雌花资源,因而可能导致榕树与榕小蜂之间合作系统的崩溃.植物果实的脱落机制普遍被认为是维持系统稳定的关键因素之一.然而,定量实验和野外观测发现P. mayri产卵并不会引起果实脱落,只有P. testacea产卵会使果实大量脱落.通过对3株样树进行比较发现:当产卵时的P. testacea数量越多时,它产生的瘿花数就越多,榕果发生脱落的比例越高.P. testacea的过度产卵是导致榕果选择性脱落的主要原因.结果表明:脱落机制并不能完全阻止非传粉小蜂的寄生,榕树只能选择性地脱落掉先于传粉小蜂产卵的榕果.这也同时表明维持榕树与榕小蜂互利共存的机制不仅仅只有榕果的脱落机制,可能还存在其它未被发现的机制.

榕小蜂食性分化与榕树-榕小蜂系统稳定性

在榕树与其传粉小蜂组成的互利阿共生系统中,理解传粉小蜂与各种非传粉小蜂如何共存是解决这一系统稳定性维持机制问题的关键之一.生态位分化被普遍认为是传粉小蜂与各种非传粉小蜂共存的最主要动力.而作为生态位分化中最基础的食性分化在这一系统中如何具体实现尚小清楚.2006年12月至2007年6月.我们以聚果榕(Ficus racemosa)为材料,通过对果内6种榕小蜂进行独立放蜂及两两组合定最放蜂,并对传粉小蜂分别进行不携带花粉和不能产卵的技术处理,研究了寄生在聚果榕果内的5种非传粉小蜂的食性及相互关系,分析了在不同季节下寄生蜂与寄主间的相关系数.研究结果表明:在5种非传粉小蜂中,Platyneura testacea和P mayri是造瘿者,能独立刺激子房发育成瘿花,并使果实发育成熟;而 Apocrypta sp、A.westwoodi和P.agraensis只能寄生于某些已发育的虫瘿,为拟寄生者,它们各自分别与P.testacea,P.mayri和传粉小蜂Ceratosolen fusciceps存在着一对一的寄生关系.拟寄生者与寄主间的相关性在不同季节下会显示出不同的结果,这表明过去文献中用物种间的相关系数推理而确定的食性关系可能是不可靠的.对自然采集榕果内的小蜂群落分析表明,传粉小蜂处于优势地位,这说明在自然情况下非传粉小蜂的种群维持在一个较低水平,对榕树-传粉小蜂系统稳定性影响较小,故能与之长期共存.

非传粉小蜂的食性及其对榕树- 传粉小蜂系统稳定性的影响

榕树与其传粉小蜂形成了高度专一的互惠共生系统。非传粉小蜂则是该系统的资源掠夺者, 但它与 该系统共存的机制仍不清楚。于2003 年12 月—2004 年4 月在西双版纳以聚果榕( Ficus racemosa L. ) 为材料, 研究了寄生在聚果榕榕果内的5 种非传粉小蜂的食性及相互关系, 以探讨非传粉小蜂与榕树- 传粉小蜂系统共 存的机制。结果表明: 寄生在聚果榕榕果内的5 种非传粉小蜂中, 仅Platyneura testacea Motschulsky 和Platyneu2 ra mayri Rasplus 能刺激子房发育成瘿花, 是造瘿者; Apocrypta sp . , Apocrypta westwoodi Grandi 和Platyneura a2 graensis Joseph 不能刺激子房发育成瘿花, 是拟寄生者。传粉小蜂的拟寄生者和造瘿者对传粉小蜂有负的影响, 但在蚂蚁和造瘿者的拟寄生蜂作用下, 这种负面影响并不显著, 而且它们对榕树繁殖没有显著影响。对小蜂自 然种群的分析表明, 传粉小蜂处于优势地位。说明在自然情况下传粉小蜂的拟寄生者和造瘿者的种群维持在一 个较低水平, 对榕树- 传粉小蜂系统稳定性影响较小, 故能与之长期共存。

Structure of a fig wasp community: Temporal segregation of oviposition and larval diets

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 testacea, Apocrypta sp2, Apocryptophagus mayri, Cera

Diffusive coevolution and mutualism maintenance mechanisms in a fig-fig wasp system

In reciprocal mutualism systems, the exploitation events by exploiters might disrupt the reciprocal mutualism, wherein one exploiter species might even exclude other coexisting exploiter species over an evolutionary time frame. What remains unclear is how such a community is maintained. Niche partitioning, or spatial heterogeneity among the mutualists and exploiters, is generally believed to enable stability within a mutualistic system. However, our examination of a reciprocal mutualism between a fig species (Ficus racemosa) and its pollinator wasp (Ceratosolen fusciceps) shows that spatial niche partitioning does not sufficiently prevent exploiters from overexploiting the common resource (i.e., the female flowers), because of the considerable niche overlap between the mutualists and exploiters. In response to an exploiter, our experiment shows that the fig can (1) abort syconia-containing flowers that have been galled by the exploiter, Apocryptophagus testacea, which oviposits before the pollinators do; and (2) retain syconia-containing flowers galled by Apocryptophagus mayri, which oviposit later than pollinators. However, as a result of (2), there is decreased development of adult non-pollinators or pollinator species in syconia that have not been sufficiently pollinated, but not aborted. Such discriminative abortion of figs or reduction in offspring development of exploiters while rewarding cooperative individuals with higher offspring development by the fig will increase the fitness of cooperative pollinating wasps, but decrease the fitness of exploiters. The fig fig wasp interactions are diffusively coevolved, a case in which fig wasps diversify their genotype, phenotype, or behavior as a result of competition between wasps, while figs diverge their strategies to facilitate the evolution of cooperative fig waps or lessen the detrimental behavior by associated fig wasps. In habitats or syconia that suffer overexploitation, discriminative abortion of figs or reduction in the offspring development of exploiters in syconia that are not or not sufficiently pollinated will decrease exploiter fitness and perhaps even drive the population of exploiters to local extinction, enabling the evolution and maintenance of cooperative pollinators through the movement between habitats or syconia (i.e., the metapopulations).

Seasonal changes in the trade-off among fig-supported wasps and viable seeds in figs and their evolutionary implications

What the real trade-off is among fig-supported wasps and the viable seeds of figs is heatedly debated in the studies of fig/fig wasp mutualism. In the present study, we collected wasp offspring (galls) and the viable seeds of premature fruits, and determined the foundress number in receptive fruits and all the types of wasps supported by Ficus racemosa L. during both the rainy and dry seasons in Xishuangbanna, China. The data show that the galls were positively correlated with viable seeds (n=32;r=0.74; P < 0.001) when the proportion of vacant female flowers (PVFF) was high, in April (68.0%), and were negatively correlated with viable seeds (n=48;r=-0.59; P < 0.05) when PVFF were limited (PVFF 42.6%) during a colder month (January). The mean foundress number per fruit during the colder months is significantly lower than during the warmer months (F-5,F-603 = 27.9; P < 0.001) and pollinator wasps can live longer during the colder months, During the colder months, the proportions of non-pollinators and wasp offspring are higher than those found during other months, whereas the proportion of viable seeds is not different compared with that of other months. Non-pollinator wasps tend to oviposit the female flowers that have been oviposited by pollinator wasps. The non-pollinators only negatively affect pollinator wasps and there is no obvious negative effect of non-pollinator wasps on viable seeds, so ovipositing by non-pollinator wasps will not result in the extinction of the figs during the process of evolution. The results of the present study indicate that figs can allow less foundresses to be in fruit cavities when PVFF are limited, which provides supporting evidence for the previous assumption that the plants have developed a mechanism to maintain a stable system because of the conflicts between the parties involved.