Host-specificity and coevolution among pollinating and nonpollinating New World fig wasps

Figs (Ficus spp., Moraceae) and their pollinating wasps (Hymenoptera, Agaonidae, Chalcidoidea) constitute a classic example of an obligate plant-pollinator mutualism, and have become an ideal system for addressing questions on coevolution, speciation, and the maintenance of mutualisms. In addition to pollinating wasps, figs host several types of nonpollinating, parasitic wasps from a diverse array of Chalcid subfamilies with varied natural histories and ecological strategies (e.g. competitors, gallers, and parasitoids). Although a few recent studies have addressed the question of codivergence between specific genera of pollinating and nonpollinating fig wasps, no study has addressed the history of divergence of a fig wasp community comprised of multiple genera of wasps associated with a large number of sympatric fig hosts. Here, we conduct phylogenetic analyses of mitochondrial DNA sequences (COI) using 411 individuals from 69 pollinating and nonpollinating fig wasp species to assess relationships within and between five genera of fig wasps (Pegoscapus, Idarnes, Heterandrium, Aepocerus, Physothorax) associated with 17 species of New World Urostigma figs from section Americana. We show that host-switching and multiple wasp species per host are ubiquitous across Neotropical nonpollinating wasp genera. In spite of these findings, cophylogenetic analyses (TREEMAP 1.0, TREEMAP 2.02beta, and parafit) reveal evidence of codivergence among fig wasps from different ecological guilds. Our findings further challenge the classical notion of strict-sense coevolution between figs and their associated wasps, and mirror conclusions from detailed molecular studies of other mutualisms that have revealed common patterns of diffuse coevolution and asymmetric specialization among the participants.     

Complex patterns of host switching in New World arenaviruses

We empirically tested the long-standing hypothesis of codivergence of New World arenaviruses (NWA) with their hosts. We constructed phylogenies for NWA and all known hosts and used them in reconciliation analyses. We also constructed a phylogenetic tree of all Sigmodontinae and Neotominae rodents and tested whether viral-host associations were phylogenetically clustered. We determined host geographical overlap to determine to what extent opportunity to switch hosts was limited by host relatedness or physical proximity. With the exception of viruses from North America, no phylogenetically codivergent pattern between NWA and their hosts was found. We found that different virus clades were clustered differently and that Clade B with members pathogenic to humans was randomly distributed across the rodent phylogeny. Furthermore, viral relatedness within Clade B was significantly explained by the geographic overlap of their hosts' ranges rather than host relatedness, indicating that they are capable of host switching opportunistically. This has important bearings on their potential to become panzootic. Together, these analyses suggest that NWA have not codiverged with their hosts and instead have evolved predominantly via host switching.     

Hantavirus evolution in relation to its rodent and insectivore hosts: no evidence for codivergence

Hantaviruses are considered one of the best examples of a long-termassociation between RNA viruses and their hosts. Based on theappearance of strong host specificity, it has been suggestedthat hantaviruses cospeciated with the rodents and insectivoresthey infect since these mammals last shared a common ancestor,approximately 100 million years ago. We tested this hypothesisof host–virus codivergence in two ways: 1) we used cophylogeneticreconciliation analysis to assess the fit of the virus treeonto that of the host and 2) we estimated the evolutionary ratesand divergence times for the Hantavirus genus using a BayesianMarkov Chain Monte Carlo method and similarly compared thesewith those of their hosts. Our reconciliation analysis providedno evidence for a history of codivergence between hantavirusesand their hosts. Further, the divergence times for the Hantavirusgenus were many orders of magnitude too recent to correspondwith the timescale of their hosts' speciation. We thereforepropose that apparent similarities between the phylogenies ofhantaviruses and their mammalian hosts are the result of a morerecent history of preferential host switching and local adaptation.Based on the presence of clade-defining amino acids in all genomicsegments, we propose that the patterns of amino acid replacementin these viruses are also compatible with a history of host-specificadaptation.     

Phylogenetic congruence of mealybugs and their primary endosymbionts

Tight interactions between unrelated organisms such as is seen in plant-insect, host-parasite, or host-symbiont associations may lead to speciation of the smaller partners when their hosts speciate. Totally congruent phylogenies of interacting taxa have not been observed often but a number of studies have provided evidence that various hemipteran insect taxa and their primary bacterial endosymbionts share phylogenetic histories. Like other hemipterans, mealybugs (Pseudococcidae) harbour multiple intracellular bacterial symbionts, which are thought to be strictly vertically inherited, implying codivergence of hosts and symbionts. Here, robust estimates of phylogeny were generated from four fragments of three nuclear genes for mealybugs of the subfamily Pseudococcinae, and a substantial fragment of the 16S-23S rDNA of their P-endosymbionts. Phylogenetic congruence was highly significant, with 75% of nodes on the two trees identical, and significant correlation of branch lengths indicated coincident timing of cladogenesis. It is suggested that the low level of observed incongruence was influenced by uncertainty in phylogenetic estimation, but evolutionary outcomes other than congruence, including host shifts, could not be rejected.     

榕属与榕小蜂科的分类研究进展

简述了榕树与其传粉者的共生体系,讨论了榕属植物与榕小蜂科的分类历史和现状,指出了二者分 类研究中存在的分歧,并对榕属与榕小蜂科的分类不匹配问题进行了评述。结合我国榕属与榕小蜂科分类的 研究现状,展望今后研究的方向及前景。     

传粉榕小蜂与非传粉小蜂间寄主识别行为的趋同进化

在高度专性传粉的榕树-榕小蜂互惠共生系统中普遍存在着一些非传粉小蜂,它们中的一些种类进入果腔后也能为榕树传粉,且在形态和物候上已与传粉榕小蜂发生了趋同进化。但其寄主识别行为是否也与传粉榕小蜂发生了趋同进化还不得而知。我们在西双版纳选择了钝叶榕(Ficuscurtipes)及其3种进果繁殖小蜂开展了相关的行为实验。3种小蜂中,1种是钝叶榕的专性传粉榕小蜂(Eupristina sp.),另外2种是寄居性非传粉小蜂(杨氏金小蜂Diaziellayangi和Lipothymus sp.),这2种非传粉小蜂进入果腔后也像传粉榕小蜂那样为钝叶榕传粉。我们以钝叶榕不同发育时期的榕果及这3种小蜂为材料,采用Y型嗅觉仪观察了这3种小蜂对各发育时期榕果和信息化学物质6-甲基-5-庚烯-2-醇、6-甲基-5-庚烯-2-酮及这2种化合物的混合物的选择行为。结果表明,当提供雌花期榕果与其他发育时期榕果和空气对照供这3种小蜂选择时,它们均显著地偏向于选择雌花期榕果;当提供雄花期榕果与其他发育时期榕果和空气对照供这3种小蜂选择时,它们均显著地偏向于选择其他发育时期榕果和空气对照,即都会避开雄花期榕果;此外,这3种小蜂均对钝叶榕雌花期果释放的一种主要化合物6-甲基-5-庚烯-2-醇的同一剂量(1μL)表现出显著的偏好。这一结果为传粉榕小蜂与非传粉小蜂间的寄主识别行为趋同进化的假说提供了证据。     

A cophylogenetic perspective of RNA-virus evolution

The extent to which viruses and their hosts codiverge remains an open question, given that numerous cases of both "cospeciation" and horizontal switching have recently been documented. DNA viruses that form persistent infections are thought to be the most likely candidates for phylogenetic congruence. Phylogenetic reconciliation analysis was used to compare established phylogenies for four RNA viruses and their hosts. The analysis employs a cophylogeny mapping technique, implemented in TreeMap v2.0, to find the most parsimonious combinations of evolutionary events able to reconcile any incongruence. This technique is guaranteed to recover all potentially optimal solutions to the reconciled tree and specifically tests the null hypothesis that an associate phylogeny is no more congruent with a host phylogeny than would be a random tree with the same taxon set. Phylogenies for Hantavirus, Spumavirus, and avian sarcoma leukosis virus were found to be significantly similar to their host trees, whereas Lyssavirus and Arenavirus displayed no significant congruence. These results demonstrate that RNA viruses are able to form stable associations with their hosts over evolutionary time scales and that the details of such associations are consistent with persistent infection being a necessary but not sufficient precondition.     

Ability to gall: the ultimate basis of host specificity in fig wasps?

1. Fig trees (Ficus spp.) and their host‐specific pollinator fig wasps (Agaonidae) are partners in an obligate mutualism. Receptive phase figs release specific volatiles to attract their pollinators, and this is generally effective in preventing pollinator species from entering figs of the wrong hosts. 2. If entry is attempted into atypical host figs, then ostiole size and shape and style length may also prevent reproduction. In spite of these barriers, there is increasing evidence that fig wasps enter atypical hosts, and that this can result in hybrid seed and fig wasp offspring. 3. This study examines the basis of pollinator specificity in two dioecious fig species from different geographical areas. Kradibia tentacularis pollinates Ficus montana in Asia. Ficus asperifolia from East Africa is closely related but is pollinated by a different species of Kradibia. 4. In glasshouses, K. tentacularis was attracted to its normal host, F1s and backcrosses, but only rarely entered figs of F. asperifolia. Foundresses were able to lay eggs in hybrids, backcrosses, and F. asperifolia, although flower occupancy was lowest in F. asperifolia figs and intermediate in hybrids. 5. The fig wasp failed to reproduce in female F. montana, male F. asperifolia, and male F1s, and most but not all backcrosses to F. montana. This was a result of the failure to initiate gall production. 6. Host specificity in this fig wasp is strongly influenced by host volatiles, but the ability to gall may be the ultimate determinant of whether it can reproduce.     

Degradation of aberrant proteins by larval tobacco hornworm,Manduca sexta L. (Sphingidae)

The tobacco hornworm Manduca sexta (Sphingidae) readily incorporates L-canavanine, the L-2-amino-4-(guanidinooxy)butyric acid structural analog of L-arginine, into newly synthesized proteins. As a result, the developing fifth-instar larva produces structurally aberrant canavanyl proteins that can exhibit severely impaired function. This situation is exacerbated by canavanine's ability to stimulate de novo protein synthesis. M. sexta larvae can respond to anomalous protein production by degrading canavanyl proteins nearly five times faster than normal proteins. The proteases of this insect can distinguish between normal and anomalous proteins and thereby avoid destruction of essential macromolecules. Aberrant protein degradative activity is not dependent upon de novo protein synthesis induced by canavanyl proteins. The fat body appears to be the source of proteases that degrade aberrant proteins; degradation is curtailed in the presence of sulfhydryl protease inhibitors as well as inhibitors of trypsin-like activity.     

Development of 17 microsatellite markers for Ceratosolen constrictus, the pollinating fig wasp of Ficus fistulosa

The obligate mutualism between figs (Ficus) and fig pollinating wasps (Agaonidae) is regarded as a classic example of mutualism. Seventeen polymorphic microsatellite loci were developed for Ceratosolen constrictus, the pollinating wasp of the dioecious fig Ficus fistulosa. The number of alleles per locus ranged from two to 15 and the observed and expected heterozygosities ranged from 0.040 to 0.846 and from 0.040 to 0.916, respectively. These microsatellite loci offer a powerful tool for evolutionary and population genetic studies in C. constrictus, and gene flow of F. fistulosa.