Chromosome numbers are not fixed in Agaonidae (Hymenoptera: Chalcidoidea)

Agaonidae (the pollinators of fig trees, Ficus spp., Moraceae) are a distinctive family of chalcid wasps with uncertain affinities within the Chalcidoidea. Chromosome numbers have only been described previously for one species (Blastophaga psenes, 2n = 12, the pollinator of F. carica, of subgenus Ficus). In this paper, we used a modified technique to analyze the karyology of three Ceratosolen species, Ceratosolen solmsi, C. graveli and C. emarginatus which are associated with figs of Ficus subgenus Sycomorus. Their karyotypes are 2n = 10. All the chromosomes are metacentric (NF = 20) with little difference in relative lengths, making them hard to distinguish. The variation in chromosome numbers that we have detected suggests that karyology can usefully complement molecular-based studies of the phylogeny of fig wasps.     

Laying the foundations of evolutionary and systematic studies in crickets (Insecta,Orthoptera): a multilocus phylogenetic analysis

Orthoptera have been used for decades for numerous evolutionary questions but several of its constituent groups, notably crickets, still suffer from a lack of a robust phylogenetic hypothesis. We propose the first phylogenetic hypothesis for the evolution of crickets sensu lato, based on analysis of 205 species, representing 88% of the subfamilies and 71% tribes currently listed in the database Orthoptera Species File (OSF). We reconstructed parsimony, maximum likelihood and Bayesian phylogenies using fragments of 18S, 28SA, 28SD, H3, 12S, 16S, and cytb (~3600 bp). Our results support the monophyly of the cricket clade, and its subdivision into two clades: mole crickets and ant‐loving crickets on the one hand, and all the other crickets on the other (i.e. crickets sensu stricto). Crickets sensu stricto form seven monophyletic clades, which support part of the OSF families, “subfamily groups”, or subfamilies: the mole crickets (OSF Gryllotalpidae), the scaly crickets (OSF Mogoplistidae), and the true crickets (OSF Gryllidae) are recovered as monophyletic. Among the 22 sampled subfamilies, only six are monophyletic: Gryllotalpinae, Trigonidiinae, Pteroplistinae, Euscyrtinae, Oecanthinae, and Phaloriinae. Most of the 37 tribes sampled are para‐ or polyphyletic. We propose the best‐supported clades as backbones for future definitions of familial groups, validating some taxonomic hypotheses proposed in the past. These clades fit variously with the morphological characters used today to identify crickets. Our study emphasizes the utility of a classificatory system that accommodates diagnostic characters and monophyletic units of evolution. Moreover, the phylogenetic hypotheses proposed by the present study open new perspectives for further evolutionary research, especially on acoustic communication and biogeography.     

A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera)

Chalcidoidea (Hymenoptera) is extremely diverse with an estimated 500 000 species. We present the first phylogenetic analysis of the superfamily based on both morphological and molecular data. A web‐based, systematics workbench mx was used to score 945 character states illustrated by 648 figures for 233 morphological characters for a total of 66 645 observations for 300 taxa. The matrix covers 22 chalcidoid families recognized herein and includes 268 genera within 78 of 83 subfamilies. Morphological data were analysed alone and in combination with molecular data from ribosomal 18S (2105 bp) and 28S D2–D5 expansion regions (1812 bp). Analyses were analysed alone and in combined datasets using implied‐weights parsimony and likelihood. Proposed changes in higher classification resulting from the analyses include: (i) recognition of Eriaporidae, revised status; (ii) recognition of Cynipencyrtidae, revised status; (iii) recognition of Azotidae, revised status; (iv) inclusion of Sycophaginae in Agaonidae, revised status; (v) reclassification of Aphelinidae to include Aphelininae, Calesinae, Coccophaginae, Eretmocerinae and Eriaphytinae; (vi) inclusion of Cratominae and Panstenoninae within Pteromalinae (Pteromalidae), new synonymy; (vii) inclusion of Epichrysomallinae in Pteromalidae, revised status. At a higher level, Chalcidoidea was monophyletic, with Mymaridae the sister group of Rotoitidae plus the remaining Chalcidoidea. A eulophid lineage was recovered that included Aphelinidae, Azotidae, Eulophidae, Signiphoridae, Tetracampidae and Trichogrammatidae. Eucharitidae and Perilampidae were monophyletic if Eutrichosomatinae (Pteromalidae) was included, and Eupelmidae was monophyletic if Oodera (Pteromalidae: Cleonyminae) was included. Likelihood recovered a clade of Eupelmidae + (Tanaostigmatidae + (Cynipencyrtus + Encyrtidae). Support for other lineages and their impact on the classification of Chalcidoidea is discussed. Several life‐history traits are mapped onto the new phylogeny.     

The structure of the female reproductiiye system in the Agaonidae (Chalcidoidea,Hymenoptera)

An account is given of the gaster, ovipositor mechanism and reproductive system in female Agaonidae. The structure and possible functions are compared with that of other chalcid families and the phylogenetic relationships of the group are discussed.     

Phylogeny and classification of the Signiphoridae (Hymenoptera: Chalcidoidea)

Abstract. A data set consisting of twenty-eight anatomical characters scored for twenty-eight terminal taxa representing the world fauna of Signiphoridae was analysed using parsimony and compatibility methods. The Coccophaginae (Aphelinidae) and the Azotinae (Aphelinidae) were used as outgroups to establish polarity of character state changes. Relationships of Signiphoridae to other Chalcidoidea are discussed. Several multistate characters were treated in the parsimony analyses either as unordered or as ordered into transformation series using additive binary coding, which in some cases drastically reduced the number of equally parsimonious solutions. Monophyly of Signiphoridae is supported by seven synapomorphies. Four genera, Chartocerus, Thysanus, Clytina and Signiphora , are recognized within Signiphoridae based on synapomorphies. Rozanoviella syn.n . and Kerrichiella syn.n . are synonymized under Signiphora. Species of Signiphora are further assigned to four species groups, three of which are demonstrably monophyletic. Nine species or subspecies are transferred to Chartocerus from Signiphora ( australicus comb.n. , australiensis comb.n. , australiensis orbiculatus comb.n., beethoveni comb.n. , corvinus comb.n. , funeralis comb.n. , reticulata comb.n. , ruskini comb.n. , thusanoides comb.n.) , one species to Thysanus from Signiphora (melancholicus comb.n.) , and one species to Signiphora from Kerrichiella ( coleoptratus comb.n.) . A key to genera of Signiphoridae and species groups of Signiphora is presented. A diagnosis, relevant nomenclatural history, and a list of included species are given for each genus and species group, and the biology and distribution of each is summarized.     

信息化合物对薜荔榕小蜂选择行为的影响

爱玉子和薜荔均为桑科榕属植物,与薜荔榕小蜂保持稳定的共生关系,信息化合物在维系榕树与其传粉小蜂的共生关系中起决定性作用。应用Y型嗅觉仪检测两种来源的薜荔榕小蜂对8种单一信息化合物和芳樟醇、苯甲醇和松油醇组成的混合信息化合物的趋向性反应。实验结果表明:3%和5%芳樟醇、5%的苯乙烯、7%苯甲醇、5%和7%松油醇以及7%金合欢醇对两种来源的薜荔榕小蜂均具有显著的吸引作用,7%苯甲酸甲酯对两种来源的薜荔榕小蜂有显著的趋避作用,香兰醛对两种来源的薜荔榕小蜂的行为没有显著的影响,从而表明信息化合物的种类和浓度均显著影响薜荔榕小蜂的选择行为。5%芳樟醇、3%松油醇和3%苯甲醇的混合信息化合物组合对两种来源的薜荔榕小蜂具极显著的吸引作用,信息化合物之间的交互作用极为显著。两种不同来源的薜荔榕小蜂对信息化合物反应的异同,表明两种不同来源的薜荔榕小蜂与各自宿主之间的化学通讯信号传导上存在差异,同时也表明两种不同来源的薜荔榕小蜂之间存在密切的亲缘关系。在榕树接受期花序的挥发物成分中广泛存在并占较大比例的信息化合物(例如芳樟醇),可能与榕小蜂对寄主的远程定位有关,而近距离的专性定位却是依赖于各种信息化合物严格按一定浓度、比例组成的化学图谱。实验结果为榕-蜂共生体系的化学生态学理论研究和爱玉子高产栽培提供科学依据。     

Rampant host switching and multiple female body colour transitions in Philotrypesis (Hymenoptera: Chalcidoidea: Agaonidae)

Figs (Ficus, Moraceae) and their associated fig waSPS (Hymenoptera, Chalcidoidea and Agaonidae) have attracted much attention and have been used as a model system for many studies. Fig waSPS belonging to the genus Philotrypesis are very common in most figs but their taxonomy, ecology and biology are currently poorly explored. A previous study on African Philotrypesis showed that their host association is phylogenetically conserved at subsection level. We reconstructed a molecular phylogeny with extended sampling from seven sections of figs. Our study suggested that the diversification of Philotrypesis is less constrained by host figs. Host switching is rampant between figs at species level and even at section level. We also investigated the evolution of the body colour forms in female Philotrypesis. Our study first suggested that female body colour is not evolutionarily stable and that there have been multiple transitions. Possible mechanisms for multiple colour transitions are expected to be determined in the near future.     

延腹小蜂属榕小蜂对榕果果内空腔大小差异的行为和形态适应

大多数榕小蜂的雄蜂终生都在密闭的榕果中生活,榕果间果内空腔的大小存在很大的差异,这种差异可能限制对小蜂的活动。已有研究表明榕果的大小会影响果内榕小蜂雄蜂的打斗行为,那么这种差异是否会影响雄蜂的果内爬行行为? 雄蜂是否会产生适应性的形态特征？针对这些问题,在2008年6月到2009年10月期间,我们在野外采集榕果,并带回实验室中, 对榕果内生活的延腹小蜂属Philotrypesis 5种雄蜂的果内爬行行为和足的形态特征进行了研究。结果表明,根据Philotrypesis spp.雄蜂后足的跗节形态特征的不同,其后足可分成2种形态型--Ⅰ型：后足跗节基部两节的突出部分排列紧密, 后足中跗节不发达; Ⅱ型：后足跗节基部两节的突出部分排列疏松,二者之间有较宽的分隔,后足中跗节发达。GLM分析结果表明各种榕树上生活的Philotrypesis雄蜂前足腿节长宽比存在极显著差异（F_{8, 81}=94.86, P<0.001）。在大果中生活的Philotrypesis雄蜂都具有较细长的前足腿节,在小果中生活的Philotrypesis雄蜂除了M7#c-ben外,都具有较粗壮的前足腿节。当雄蜂在榕果内寻偶时,主要依靠粗壮的前足或灵活的后足在果内爬行。结果提示, 榕果空腔大小可能限制了Philotrypesis雄蜂果内爬行行为, 进而影响了雄蜂足形态的适应性进化。     

The life history and larval morphology of Aperilampus (Hymenoptera: Chalcidoidea: Philomidinae), with a discussion of the phylogenetic affinities of the Philomidinae

Abstract. The host association and larval morphology of Aperilampus (Hymenoptera: Chalcidoidea: Philomidinae) are documented for the first time based on specimens collected during the excavation of ground-nesting bee nests in South Africa. Aperilampus varians Strand is a primary parasite of the pupa of Halictus (Seladonia) africanus Friese (Hymenoptera: Halictidae). The first and final-instar larvae are described and illustrated, and notes are provided on the behaviour of the first-instar larva. The first-instar larva is a planidium and synapomorphies are documented between the first-instar larva of Aperilampus and the planidia of Chrysolampinae, Perilampidae and Eucharitidae.     

Subforaminal bridges in Hymenoptera (Insecta), with a focus on Chalcidoidea

Variation in structures of the posterior surface of the head in Hymenoptera is compared and interpreted according to theories of head capsule evolution, with focus on understanding previously baffling conditions in the superfamily Chalcidoidea. Features are investigated separately without first classifying subforaminal bridges into subcategories. In Proctotrupomorpha (including Chalcidoidea), Ceraphronoidea and some Ichneumonoidea, there are multiple posterior pits associated with the tentorium. In most examined Hymenoptera with a subforaminal bridge, there was a differentiated median area, typically with highly variable microtrichia. This area is elevated in Cephoidea and Pamphilioidea, but is not elevated in other Hymenoptera. Subforaminal bridges in Apocrita previously classified as hypostomal bridges are discussed in the context of A.P. Rasnitsyn's hypothesis that relative importance of adult feeding drives subforaminal bridge evolution.     

A new whitefly parasitoid (Hymenoptera: Pteromalidae: Eunotinae), with comments on its placement, and implications for classification of Chalcidoidea with particular reference to the Eriaporinae (Hymenoptera: Aphelinidae)

Idioporus affinis gen.n. , sp.n . (Hymenoptera: Pteromalidae: Eunotinae) is described from Central America, and the new tribe, Idioporini, is created for it. Reasons for its placement in the Eunotinae are discussed, as well as implications for higher classification within the Chalcidoidea, particularly with regard to the Eriaporinae (currently placed in the Aphelinidae). Idioporus affinis is a parasitoid of the giant whitefly Aleurodicus dugesii Cockerell (Homoptera: Aleyrodidae), and is a potential biological control agent of this whitefly which is a recently introduced pest in California, Louisiana and Texas.     

A new genus of Encyrtidae (Hymenoptera: Chalcidoidea) from India

A new genus, probably related to Anagyrus, is erected for a new species parasitic on mealy-bugs, the male and female of which are described.     

The pretarsus in Chalcidoidea (Hymenoptera Parasitica): functional morphology and possible phylogenetic implications

The structure of the pretarsus of chalcid wasps (Hymenoptera: Chalcidoidea) was examined with light and scanning electron microscopy. The pretarsus of these wasps is characterized by a distal elastic widening of the planta that spreads over the arcus, by a pair of folding plates at the dorsal side of the arolium (the dorsal plates), and by the absence of auxiliary sclerites. The surface of the fully spread arolium of chalcids has a spongiform structure. The arcus of chalcids is an apodeme of the planta. The peculiarities of the inverting/everting biomechanics of the pretarsus of chalcids involve: 1) interactions between the elastic part of the planta, the dorsal plates and the manubrium, and 2) the functioning of the elastic part of the planta and the arcus together as a single unit. A single apical seta situated distally from the campaniform sensillae and proximal row of setae on the manubrium are regarded as putative synapomorphies of Chalcidoidea. A manubrium with a distinct proximal row of three setae characterizes almost all Eulophidae, Aphelinidae and Signiphoridae (‘eulophid lineage’) and Tetracampidae, whereas a row of two setae characterizes Mymaridae, Rotoitidae and Trichogrammatidae. Other studied families (Pteromalidae, Eurytomidae, Torymidae, Ormyridae, Eupelmidae, Encyrtidae, Perilampidae), which represent a ‘pteromalid lineage’, are characterized mostly by five setae in a proximal row, which could represent a synapomorphy for these groups, or a symplesiomorphy in Chalcidoidea, depending on rooting. However, the characters may be correlated with differences in body size that characterize the different lineages rather than being phylogenetically important. Other characters that may be phylogenetically informative are: 1) shape of the manubrium (spindle‐like in Mymaridae, Rotoitidae, Trichogrammatidae and the ‘eulophid lineage’, but mostly bottle‐like in representatives of the ‘pteromalid lineage’), and 2) pubescence of the proximal part of the planta (sparse, thick setae in Rotoitidae, Trichogrammatidae and the ‘eulophid lineage’, but dense, slender setae in representatives of the ‘pteromalid lineage’).     

A molecular phylogeny of the Trichogrammatidae (Hymenoptera: Chalcidoidea), with an evaluation of the utility of their male genitalia for higher level classification

Abstract A molecular phylogeny of the Trichogrammatidae (Hymenoptera: Chalcidoidea) is presented. This group of minute egg parasitoids is known from a broad range of host insects. The phylogeny produced, the first of any kind for the family, utilizes 121 taxa in fifty‐two of the eighty‐four recognized genera. Results were inferred from the ribosomal RNA regions 18S, 28S‐D2 and 28S‐D3, all aligned according to secondary structure models. Parsimony analysis was performed on both a complete and reduced dataset, in which ambiguous regions as defined by secondary structure were eliminated. The reduced dataset produced a much less resolved phylogenetic hypothesis. Only the complete dataset was utilized for Bayesian and maximum likelihood analyses. A robust‐choice Bayesian hypothesis stemmed from the concatenation of five distinct character set parameters. The results are compared with the current classification based primarily on male genitalia. Although our conclusions are partially congruent with the accepted hypothesis of trichogrammatid relationships, none of the currently adopted assemblages was recovered as monophyletic. Nevertheless, the structure of the male genitalia does correspond with relative taxon position in the molecular hypotheses. In general, the greatest genitalic simplification and fusion characterizes taxa treated as the most derived in the molecular hypotheses. Several groups are consistently recovered, but relationships between these groups and other genera vary with the analytical method. A new classification of the Trichogrammatidae is proposed. It includes one tribe, Trichogrammatini, within the subfamily Trichogrammatinae, and three tribes, Paracentrobiini, Chaetostrichini and Oligositini, within the Oligositinae. All tribes are more narrowly defined than previously and many genera are treated as incertae sedis within each subfamily. The results are interpreted in the light of morphological evidence, and the placement of genera not represented in molecular analysis is inferred by morphology alone. Host associations throughout the family are varied, although the molecular results suggest that Trichogrammatidae primitively parasitized Coleoptera, with more recent radiations onto other insect orders, such as Lepidoptera and Hemiptera.     

Laying the foundations for a bio-economy

Biological technologies are becoming an important part of the economy. Biotechnology already contributes at least 1% of US GDP, with revenues growing as much as 20% annually. The introduction of composable biological parts will enable an engineering discipline similar to the ones that resulted in modern aviation and information technology. As the sophistication of biological engineering increases, it will provide new goods and services at lower costs and higher efficiencies. Broad access to foundational engineering technologies is seen by some as a threat to physical and economic security. However, regulation of access will serve to suppress the innovation required to produce new vaccines and other countermeasures as well as limiting general economic growth.

Two new British species of Halticoptera (Hymenoptera: Chalcidoidea,Pteromalidae)

The female of a new species of Halticoptera is described from a single specimen from Britain, and the male and female of a second new species are described from material from Britain and Czechoslovakia. Halticop-terina triannulata is recorded from Britain for the first time.     

Phylogenetic implications of the mesofurca in Chalcidoidea (Hymenoptera), with emphasis on Aphelinidae

The skeletomusculature of the mesofurcal–mesopostnotal complex is surveyed within the Chalcidoidea. Four internal character systems are assessed for their phylogenetic significance: the mesofurcal bridge, the structure and position of the furcal–laterophragmal muscle, the structure of the lateral arms of the mesofurca, and the supporting structures for the interfurcal muscles. Among Hymenoptera, Chalcidoidea are unique in having the furcal–laterophragmal muscle attached along the entire length of the laterophragmal apodeme. Also the furcal–laterophragmal muscle originates medial to the junction of the mesofurcal bridge and lateral mesofurcal arm in most Chalcidoidea. Mymarommatidae do not share either of these apomorphic states with Chalcidoidea. Within Chalcidoidea, apomorphic character states were found in each of Aphelinidae, Encyrtidae, Eulophidae, Mymaridae, Rotoitidae, Signiphoridae, Tanaostigmatidae and Trichogrammatidae. For taxa classified as Aphelinidae, the plesiomorphic complement of structures and muscle attachments is retained in Eriaphytinae and Eriaporinae. The mesofurcal bridge is considered to have been lost at least twice in each of Aphelininae and Coccophaginae. Similar interfurcal processes, resulting from loss of the mesofurcal bridge, support the monophyly of Aphelininae (Aphelinini, Aphytini and Eutrichosomellini). Azotinae are placed as the sister group of Aphelininae because of a similar lateral origin of the laterophragmal muscle and the shape of the mesofurcal arms. Other than loss of the mesofurcal bridge, no character states were shared by Azotinae and Coccophaginae. Coccophaginae (Coccophagini and Pteroptricini) are regarded as monophyletic based on the loss of the mesofurcal bridge, the peculiar shape of the mesofurca, and a unique modification of the laterophragmal muscle. Euxanthellus is removed from synonomy with Coccophagus and may be best treated as a separate tribe of Coccophaginae based on the shape of the lateral mesofurcal arms and the presence of a mesofurcal bridge. The shape of the mesofurca suggests a monophyletic grouping of Cales, Eretmocerus and Trichogrammatidae that could render Aphelinidae paraphyletic.     

Rotoitidae, a curious new family of Chalcidoidea (Hymenoptera) from New Zealand

Abstract. A new chalcidoid family, Rotoitidae fam.n., is described from New Zealand. The probable systematic affinities of Rotoitidae are discussed and the type genus and a single included species, Rotoita basalis gen. et sp.n., are described and illustrated.