Phylogeny of the lacebugs (Insecta: Heteroptera: Tingidae) using morphological and molecular data

A phylogeny of the Tingidae is provided on the basis of parsimony analysis with pre‐alignment as well as with direct optimization of 2018 bp from five loci (COI, Leu‐tRNA, COII, 16S and 28S) and a set of 30 morphological characters. The results obtained with either direct optimization or pre‐alignment methods of parsimony clearly show that Cantacaderinae are sister group to (Phatnominae + Tinginae). Several apomorphies, retrieved in different studies, characterize Cantacaderinae, Phatnominae and Tinginae, but only one characterizes the grouping of Phatnominae + Tinginae. In addition, no clear tribal division as currently used was highlighted within the Tinginae. The results are discussed in the light of earlier morphologically based hypotheses.     

Correlated evolution and Bayesian divergence time estimates of the Cimicoidea (Heteroptera: Cimicomorpha) reveal the evolutionary history

The diverse habitat types and discrete morphological characters of cimicoid species provide a unique opportunity to study correlated evolution. Phylogenetic relationships within Cimicoidea were determined using Bayesian analyses of molecular data, allowing the generation of testable hypotheses of correlated evolution. An investigation of the correlation between habitat selection and morphological characters revealed that a dead plant habitat was correlated with the filiform antennal type. Furthermore, molecular dating analysis was used to examine divergence times within the Cimicoidea. Transitions to live plants from dead plants for most cimicoid clades started right after the mid‐Cretaceous, coinciding with the radiation of the angiosperms. Using contingency analyses, we determined that evolutionary changes in morphological characters were dependent on habitat selection. Based on these results, we propose evolutionary historical hypotheses for the Cimicoidea.     

Phylogeny and character evolution in the bee-assassins (Insecta: Heteroptera: Reduviidae)

Apiomerus, the charismatic bee-assassins (>108 spp.), belong to the New World resin bugs in the harpactorine tribe Apiomerini (12 extant genera) that is characterized by a novel predation strategy, resin trap predation. Apiomerini also exhibit striking genitalic diversity that has shaped subgeneric classifications within the genus Apiomerus and females of some species of Apiomerus are known to engage in unique maternal care behaviors. The lack of a phylogenetic framework currently hinders evolutionary interpretations of genitalic morphology and maternal care. We here present a molecular phylogeny based on 4, 477 bp of six ribosomal and protein coding genes and 95 terminal taxa using parsimony and maximum likelihood approaches as a way of addressing these shortcomings. Apiomerini are monophyletic, with Heniartes being the sistergroup to all remaining taxa that form the monophyletic Manicocoris (Calliclopius, Manicocoris, Micrauchenus, and Ponerobia) and Apiomerus (Agriocoris, Apiomerus, and Sphodrolestes) clades. Previously proposed subgeneric groups are polyphyletic, but several proposed species groups are recovered as monophyletic. Ancestral state reconstruction of the metatibial comb indicates that this structure evolved in the ancestor of all Apiomerini where it was present in males and in females; it became strongly sexually dimorphic (better developed in females than in males) in the Apiomerus clade (Apiomerus + Agriocoris + Sphodrolestes). Genitalic features reveal a pattern of homoplasy, but frequently are nonetheless useful to diagnose supraspecific groups within Apiomerus. The complex genitalia found within Apiomerus are derived for that clade. We conclude that, using the metatibial comb as a proxy, maternal care is relatively common in the tribe Apiomerini and propose that it likely evolved at the base of the Apiomerus clade if not at the base of Apiomerini.     

Phylogeny, biogeography, and the evolution of life-history traits in Leucadendron (Proteaceae)

Leucadendron is a moderately large genus of Proteaceae almost entirely restricted to the Cape Floristic Region of southern Africa. The genus is unusual in being dioecious and sexually dimorphic. ITS sequence data were obtained from 62 of the 96 currently recognized taxa (85 species and 11 subspecies). Phylogenetic analyses were conducted under Maximum Likelihood and parsimony and resolved nine groups of species with varying degrees of bootstrap support, but relationships between these groups are largely unsupported. The phylogeny conflicts with the current taxonomic arrangement, which is based mainly on fruit morphology. The two sections of the genus, Alatosperma and Leucadendron, and several subsections within these sections, are resolved as non-monophyletic. This means that taxonomically important characters (such as fruit shape) have evolved multiple times, as the species with nut-like fruit (resolved into two of the nine groups) appear to have evolved independently from ancestors with winged fruit. Based on the topology obtained, the life history traits of anemophily, myrmechochory, and re-sprouting have also originated multiple times. Dispersal-Vicariance (DIVA) analysis suggests that the genus had an ancestral area in the Karoo Mountain and Southeastern phytogeographic centres of endemism in the southwestern Cape.     

Phylogeny of the plant bug subfamily Mirinae (Hemiptera: Heteroptera: Cimicomorpha: Miridae) based on total evidence analysis

The first comprehensive phylogenetic analyses of the most diverse subfamily of plant bugs, Mirinae, is presented in this study, for 110 representative taxa based on total evidence analysis. A total of 85 morphological characters and 3898 bp of mitochondrial (16S, COI) and nuclear (18S, 28S) sequences were analysed for each partitioned and combined dataset based on parsimony, maximum likelihood and Bayesian inference. Major results obtained in this study include monophyly of the tribe Mecistoscelini. The largest tribe, Mirini, was recovered as polyphyletic, and Stenodemini was recovered as paraphyletic. The clade of Stenodemini + Mecistoscelini is the sister group of the remaining Mirinae. The monophyly of two complexes composed of superficially similar genera were tested; the Lygus complex was recovered as nonmonophyletic, and the Adelphocoris–Creontiades–Megacoelum complex was confirmed to be monophyletic. The generic relationships of the main clades within each tribe based on the phylogeny, as well as their supported morphological characters, are discussed.     

Cytogenetics of the true bug infraorder Cimicomorpha (Hemiptera, Heteroptera): a review

The Cimicomorpha is one of the largest and highly diversified infraorders of the Heteroptera. This group is also highly diversified cytogenetically and demonstrates a number of unusual cytogenetic characters such as holokinetic chromosomes; m-chromosomes; multiple sex chromosome systems; post-reduction of sex chromosomes in meiosis; variation in the presence/absence of chiasmata in spermatogenesis; different types of achiasmate meiosis. We present here a review of essential cytogenetic characters of the Cimicomorpha and outline the chief objectives and goals of future investigations in the field.     

Hairy attachment structures in Reduviidae (Cimicomorpha, Heteroptera), with observations on the fossula spongiosa in some other Cimicomorpha

Reduviidae and some other groups of cimicomorphan Heteroptera possess a hairy attachment structure on the apex of the tibia called “fossula spongiosa”. The fossula spongiosa was never studied comparatively across Reduviidae, its fine structure and mode of function is not well documented, and attachment structures in immature stages are virtually unknown. Here, a sample of 171 species of Reduviidae representing 22 subfamilies is examined for presence-absence of the fossula spongiosa on the three pairs of legs. Representatives of 11 of the 22 subfamilies are shown to possess a fossula spongiosa. The fine structure of the fossula spongiosa is examined for a more limited sample of Reduviidae and several Pachynomidae and Nabidae. In addition, scanning micrographs for the fossula spongiosa in other Cimicomorpha are given, among them Anthocoridae, Cimicidae, Microphysidae (first record of a fossula spongiosa), and Thaumastocoridae. The fossula spongiosa in Reduviidae consists of tenent hairs (acanthae) with spatulate or tapering apices interspersed with sensory setae, both of which are embedded in a thick and flexible cuticle, underlain by a hemolymph cavity separated almost entirely from the interior of the remaining tibia by a cuticular invagination. Judging from comparison with non-reduviid Cimicomorpha, this separation of the fossula spongiosa cavity from the tibial interior may be unique to Reduviidae. A simple experiment using live specimens of Platymeris biguttata (Reduviinae) revealed a liquid on the tip of the tenent hairs that might be involved in the attachment of the fossula spongiosa by adhesion mechanisms. The nymphs of Reduviidae whose adults have a fossula spongiosa are here documented for the first time to possess pads of ventrally barbed setae instead of tenent hairs and their tibia lacks the internal cuticular invagination. The nymphal attachment structures seem to rely on increase of friction rather than the adhesion mechanism proposed to be present in the adult. Combined with the tenent setae on the third tarsomere known in some Emesinae and here documented for Saicinae, three types of hairy attachment structures occur on the legs of Reduviidae: tenent hairs (acanthae), which form the fossula spongiosa in many Reduviidae, barbed setae that substitute the fossula in the immatures, and tenent setae on the tarsus which are restricted to only a few taxa.     

New perspectives on the evolution of exaggerated traits

The scaling of body parts is central to the evolution of morphology and shape. Most traits scale proportionally with each other and body size such that larger adults are essentially magnified versions of smaller ones. This pattern is so ubiquitous that departures from it – disproportionate scaling between trait and body size – pique interest because it can generate dramatically exaggerated traits. These extreme morphologies are frequently hypothesized to result from sexual selection and their study has a long history, with several hypotheses seeking to explain their evolution. Despite this effort, surprisingly little progress has been made in demonstrating the forms of selection that produce different scaling patterns or in identifying the mechanisms that underlie the expression and evolution of scaling relationships. Here we review recent insights regarding the proximate mechanisms that regulate and integrate trait growth and that offer a new framework for studying the evolution of morphological scaling.     

Cladistic analysis of Reduviidae (Heteroptera: Cimicomorpha) based on morphological characters

Abstract.  With more than 6600 species worldwide, Reduviidae (Insecta: Heteroptera), or assassin bugs, form the second largest and one of the most diverse groups of true bugs. The poor condition of the higher-level classification of Reduviidae is reflected by the facts that different authors recognize between 21 and 32 subfamily-level names and that Reduviidae were never subjected to a rigorous cladistic analysis using an exemplar approach. In the present study, a cladistic analysis of higher-level taxa of Reduviidae based on 162 morphological characters and 75 ingroup and outgroup species is presented. Twenty-one subfamily-level taxa of Reduviidae were examined, accounting for 28 tribes. In addition to characters previously used for diagnosis in Reduviidae, information on recently published character complexes is used in the present analysis, supplemented with new character information gathered specifically for this project. Reduviidae are supported as a monophyletic group with Pachynomidae as their sister taxon. The major results of this study are the support of a sistergroup relationship of Hammacerinae with the remaining Reduviidae, the monophyly of the Phymatine Complex, the relatively basal position of Harpactorinae within Reduviidae as well as a novel hypothesis on the relationships within this group, and the sistergroup relationship of Ectrichodiinae + Tribelocephalinae and their placement in a clade that also contains Emesinae, Saicinae, and Visayanocorinae. The analysis further supports a clade formed by paraphyletic Salyavatinae + Sphaeridopinae, renders Vesciinae non-monophyletic, and demonstrates the polyphyly of Reduviinae. Pseudocetherinae are shown to group with some Reduviinae. Triatominae are supported as a monophyletic group and are nested among additional Reduviinae and Stenopodainae.     

A new family of anthocoroid hemipterans (Hemiptera: Heteroptera,Cimicomorpha) from Cretaceous Taimyr amber

A new unusual anthocoroid monotypic family, Taimyrocoridae fam. nov., is described from Late Cretaceous resin of the Taimyr Peninsula (Yantardakh, Coniacian–Santonian). The available data on the other records of hemipterans from the Late Cretaceous localities Yantardakh (Coniacian–Santonian), Baikura (Santonian), and Nizhnyaya Agapa (Cenomanian) are presented.     

Azalea growth in response to azalea lace bug (Heteroptera: Tingidae) feeding

The effects of azalea lace bug, Stephanitis pyrioides (Scott), feeding injury on azalea growth and development were investigated using 'Girard's Rose' azaleas during a 2-yr field study in Georgia Low, medium, and high injury treatments, which corresponded to 6, 8, and 14% maximum canopy area injury, were compared with control azaleas that received no lace bug infestation. Flower number, whole-shrub leaf and stem dry mass, and dry mass and size of new growth tissues were unaffected by treatments. In contrast, growth index measurements, a general measure of variability frequently used for horticultural differentiation, showed significant reductions for all treatments in comparison to control azaleas after 20 wk. Though not directly quantified, this apparent discrepancy may be explained as an artifact of lace bug feeding-induced leaf abscission. Growth index measurements had considerable variability and may not be the most reliable measurement of size. In July 1998, plant canopy densities among azaleas maintained in the high injury treatments were approximately 15% less full than the canopies of control shrubs. Predaceous insects had a significant negative association with azalea lace bug number during the 2-yr study. Flower and new tissue production, measured destructively during two growing seasons, revealed azalea tolerance to 14% of maximum canopy area lace bug feeding-injury levels.     

Phylogeny of the water strider genus Rheumatobates (Heteroptera: Gerridae)

The genus Rheumatobates comprises thirty‐seven species and subspecies of New World water striders belonging to subfamily Rhagadotarsinae. Among species, males vary dramatically in the degree and nature of modifications of the antennae, three pairs of legs and abdominal and genital segments. Characters describing this modification have traditionally been used to differentiate and group species. The general assumption has been that modified species belong to one group and unmodified species to another. These two ‘species groups’ are subdivided into ‘subgroups’, but little effort has been made to resolve relationships among them. We conduct the first numerical cladistic analysis of Rheumatobates using a data set comprised of 102 characters, primarily describing modification of male external morphology. To address concerns about the inclusion of characters to be optimized on the phylogeny, characters describing modification of antennae and hind legs were included and then excluded in separate analyses. A preferred phylogeny was chosen from the four equally parsimonious cladograms found after successive reweighting of characters. There was good resolution at all levels of the phylogeny. Most of the major clades and terminal relationships were moderately to strongly supported, whereas the basal relationships were less well supported. The general assumption that unmodified and modified species form two monophyletic groups was not supported. However, traditionally recognized ‘subgroups’ within the modified species group were largely upheld. The analysis also suggested several major clades and relationships among these clades that were not previously recognized. The exclusion of characters describing modification of antennae and hind legs did not change the resolved major clades of the reconstructed phylogeny.     

Proximate Factors Regulating Maternal Options in the Eggplant Lace Bug, Gargaphia solani (Heteroptera: Tingidae)

The eggplant lace bug, Gargaphia solani, was used to investigate the proximate factors regulating maternal care and a noncaring, condition dependent strategy called egg dumping. We hypothesize that the act of delaying oviposition while searching for a dumping opportunity suppresses oogenesis and triggers guarding behavior. We examined several predictions of this hypothesis by measuring: (1) whether females do delay oviposition in the absence of dumping stimuli, (2) whether females in transition between egg dumping and egg guarding are capable of expressing either reproductive option, (3) the effect of nymphal interactions and antennal ablation on the duration of maternal care, and (4) oogenesis in guarding and dumping females. We found that females without a dumping opportunity wait, on average, 30 h longer to oviposit than females exposed to a dump mass. Females that had initiated their own egg mass could resume eggdumping if they had laid less than half of their eggs but were unlikely to abandon their eggs when most had been laid. Maternal care in G. solani can be prolonged if interactions with nymphs are artificially prolonged. Females require antennae to maintain maternalcare. Presumably antennae transduce cues from eggs and nymphs. Dissections of dumping and guarding females 72 h after their first oviposition demonstrated that dumpers continue to produce primary oocytes after a dumping event but guarders terminate oogenesis whilecaring for their first brood. We interpret all of these results within the context of the hypothesis that juvenile hormone titers regulate the expression of both egg dumping and egg guarding.     

Patterns of chromosome banding in four nabid species (Heteroptera, Cimicomorpha, Nabidae) with high chromosome number karyotypes

Male Nabis (Aspilaspis) indicus (St?l), N. (A.) viridulus Spinola, Himacerus (Himacerus) mirmicoides (O. Costa) (2n=32+XY) and Prostemma guttula (Fabricius) (2n=26+XY) were studied using C-banding, silver nitrate staining and base-specific fluorochrome (DAPI and CMA(3)) staining. N. indicus differed from N. viridulus in distribution pattern of C-bands, which were telomeric in the former while interstitial in the latter. H. mirmicoides showed interstitial C-bands in the majority of autosomes. P. guttula had no conspicuous C-bands in other chromosomes, but only in the Y, which was totally heterochromatic. C-heterochromatin was labelled with DAPI, indicating that it was AT-rich. In every species, both X and Y chromosomes were NOR-bearing, and the NOR regions were GC-rich. In H.mirmicoides and P. guttula, NORs showed sub-median location in the X and distal in the Y, such a pattern being probably common in Nabidae. The present paper provides new information on the genome organization and new cytological markers useful for a better insight into karyotype evolution of nabid species.     

Relationships among male sexually selected traits in the bean bug,Riptortus pedestris (Heteroptera: Alydidae)

Current concepts of sexual selection suggest that male reproductive success is determined by multiple sexual traits. As expression and production of multiple sexual traits are frequently associated with each other, positive or negative correlations among multiple sexual traits ensue. These relationships among traits associated with male reproductive success may be crucial in the evolution of male reproductive strategies. Here, we investigate phenotypic relationships among sexually selected traits in the armed bean bug Riptortus pedestris. In this insect, males with a larger body and weapon are more likely to win male–male competitions, and males with a larger weapon or higher courtship rate are more attractive to females. There was a significant positive correlation between body size and weapon size, whereas the courtship rate had significant negative correlations with body size and weapon size. Our results suggested that there was a phenotypic trade‐off between courtship rate and male morphology. In this insect, smaller males may make more effort in courtship behavior as an alternative mating tactic.     

THREE NEW SPECIES OF GENUS Stephanitis FROM CHINA (Hemiptera: Heteroptera: Tingidae)

本文报道采自中国西部的冠网蝽属（ＳｔｅｐｈａｎｉｔｉｓＳｔａｌ）３新种：祁连冠网蝽ＳｔｅｐｈａｎｉｔｉｓｑｉｌｉａｎｅｎｓｉｓＢｕｓｐ．ｎｏｖ．,采自甘肃省天祝县乌鞘岭;萧氏冠网蝽ＳｔｅｐｈａｎｉｔｉｓｈｓｉａｏｉＢｕｓｐ．ｎｏｖ．,采自四川省小金县和雅安县,曾被错误鉴定为“Ｓｔｅｐｈａｎｉｔｉｓｆａｓｃｉ－ｉｃａ－ｒｉｎａＴａｋｅｙａ”（萧采瑜等,１９８１）;狭冠网蝽ＳｔｅｐｈｎｉｔｉｓａｎａｇｕｓｔａｔａＢｕｓｐ．ｎｏｖ．采自甘肃省武都县和武威县。模式标本均保存在南开大学生物系昆虫标本室。