Gonoid thelytoky in soft scale insects (Coccidae: Homoptera)

Cytological analysis of the thelytokous soft-scale insects Coccus hesperidum L. (2n=14) and Saissetia coffeae (Walker) (2n=16) revealed that while in both species the chromosomes did not pair during prophase I, meiosis consisted of two divisions, the chromosome number was reduced, and diploidy was restored by the fusion of the female pronucleus with the polar nucleus II. The difficulty of trying to classify this type of thelytoky as either automictic or apomictic led to the proposal that a new criterion and new terms be used to classify thelytoky (and parthenogenesis). The new criterion is whether the number of chromosome elements present in the first (or only) metaphase of oogenesis is the same as that present in the oogonia (gonoid thelytoky) or different from it (agonoid thelytoky). The new criterion is superior to the existing criteria because it is unambiguous, and because it groups together forms with a similar tendency towards heterozygosity (or homozygosity). The possible evolution of the forms analyzed as well as the two other thelytokous forms of each species described by Thomsen (1927) are discussed. Another soft-scale insect, Physokermes hemicryphus Dalam, consisted of a diploid (2n=18) and a triploid (3n=27) form, in both of which the chromosomes also did not pair. Each of the three species contained a strain in which only a single nucleolus was present per cell. In C. hesperidum some strains with two nucleoli differed in the size of the nucleoli.     

Preliminary study of wing interference patterns (WIPs) in some species of soft scale (Hemiptera,Sternorrhyncha, Coccoidea,Coccidae)

The fore wings of scale insect males possess reduced venation compared with other insects and the homologies of remaining veins are controversial. The hind wings are reduced to hamulohalterae. When adult males are prepared using the standard methods adopted to females and nymphs, i.e. using KOH to clear the specimens, the wings become damaged or deformed, an so these structures are not usually described or illustrated in publications. The present study used dry males belonging to seven species of the family Coccidae to check the presence of stable, structural colour patterns of the wings. The visibility of the wing interference patterns (WIP), discovered in Hymenoptera and Diptera species, is affected by the way the insects display their wings against various backgrounds with different light properties. This frequently occurring taxonomically specific pattern is caused by uneven membrane thickness and hair placement, and also is stabilized and reinforced by microstructures of the wing, such as membrane corrugations and the shape of cells. The semitransparent scale insect’s fore wings possess WIPs and they are taxonomically specific. It is very possible that WIPs will be an additional and helpful trait for the identification of species, which in case of males specimens is quite difficult, because recent coccidology is based almost entirely on the morphology of adult females.     

Local maladaptation in the soft scale insect Saissetia coffeae (Hemiptera: Coccidae)

Local adaptation has often been documented in herbivorous insects. The potential for local maladaptation in phytophagous insects, however, has not been widely considered. I performed a two-generation reciprocal cross-transplant experiment with the generalist soft scale insect Saissetia coffeae (Hemiptera: Coccidae) on two common species of host plants in rain forest habitat in Costa Rica. In this system, S. coffeae showed significant local maladaptation at the level of the host species. Lineages originally collected from Witheringia enjoyed a strong advantage over those collected from Lomariopsis when both sets of lineages were placed on Lomariopsis; however, when both sets of lineages were raised on Witheringia, their fitnesses were statistically indistinguishable. While some aspects of the biology of S. coffeae may impair its ability to adapt to local selection pressures, scale insects are often locally adapted on fine spatial scales, and local maladaptation is therefore especially surprising. Other documented cases of local maladaptation in parasites appear to be due to evolution on the part of the host. The possibility that hosts or natural enemies may place local genotypes at a disadvantage, producing a pattern of local maladaptation, is one that deserves more consideration in the context of plant-insect interactions.     

Maternal effects in the soft scale insect Saissetia coffeae (Hemiptera: Coccidae)

Effects of maternal environment on offspring performance have been documented frequently in herbivorous insects. Despite this, very few cases exist in which exposure of parent insects to a resource causes the phenotype of their offspring to be adjusted in a manner that is adaptive for that resource, a phenomenon called adaptive transgenerational phenotypic plasticity. I performed a two-generation reciprocal cross-transplant experiment in the field with the soft scale insect Saissetia coffeae (Hemiptera: Coccidae) on two disparate host plant species in order to separate genetic effects from possible transgenerational plasticity. Despite striking differences in quality between host species, maternal host had no effect on overall offspring performance, and I detected no "acclimatization" to the maternal host species. However, there was a significant negative association between maternal and offspring development times, with potentially adaptive implications. Furthermore, offspring of mothers reared in an environment where scale densities were higher and scales were more frequently killed by fungi were significantly less likely to suffer from fungal attack than were offspring of mothers reared in an environment where densities were low and fungal attack was rare. Although S. coffeae does not appear to alter offspring phenotype to increase offspring fitness on these two distinct plant species, it does appear that offspring phenotype may be responding to some subtler aspects of maternal environment. In particular, the possibility of induced transgenerational prophylaxis in S. coffeae deserves further investigation.     

中国蜡蚧科一新记录属（半翅目：蚧总科）

记述采自中国云南的1新记录属:类白蜡蚧属Ericeroides Danzig,1990及1新记录种:越南类白蜡蚧E.zaitzevi Danzig,1990。该新记录属近似于白蜡蚧属Ericerus Guérin-Méneville,二属均有成群气门刺及爪齿;主要区别是:类白蜡蚧属Ericeroides无背面管状腺及肛前孔,而白蜡蚧属Ericerus则有。文中对雌成虫形态特征进行了详细描述和绘图。     

The hinged back plate mechanism in glassy wax tests of New Zealand male soft scale insects (Hemiptera: Coccoidea: Coccidae)

Male scale insects (Hemiptera: Coccoidea) undergo a metamorphosis of the neometabola type, from scale-like nymph through prepupa and pupa to winged adult. The nymphal instar before prepupa secretes a waxy protective covering that remains in place throughout metamorphosis and these covers are characteristic of each family of scale insects. Most scale insect families (e.g. mealybugs, eriococcids, diaspidids) have rather loosely woven male covers, but male nymphs in the family Coccidae (soft scales) construct more rigid, glassy wax tests, which need a special mechanism for adult emergence. In the New Zealand male soft scales, a suture across the posterior quarter of the test enables the back plate to flex at a pair of hinges, to be raised up off the substrate, and so allow egress. The waxy back plate hinges are secreted by groups of tubular ducts on the abdominal dorsum of 2nd-instar males, during construction of the test. Scanning electron micrographs (SEMs) show the detail and diversity of hinge types. The wax tests of most New Zealand Coccidae, both female and male, are apparently unique in that they are constructed in rows of hexagonal plates, separated by sutures, however in the male test, the sutures are all fused except for the back plate suture. The two species in the endemic New Zealand genus Pounamococcus have male tests more like those of species in the Australian genus Austrolecanium.     

A preliminary phylogeny of the scale insects (Hemiptera: Sternorrhyncha: Coccoidea) based on nuclear small-subunit ribosomal DNA

Scale insects (Hemiptera: Sternorrhyncha: Coccoidea) are a speciose and morphologically specialized group of plant-feeding bugs in which evolutionary relationships and thus higher classification are controversial. Sequences derived from nuclear small-subunit ribosomal DNA were used to generate a preliminary molecular phylogeny for the Coccoidea based on 39 species representing 14 putative families. Monophyly of the archaeococcoids (comprising Ortheziidae, Margarodidae sensu lato, and Phenacoleachia) was equivocal, whereas monophyly of the neococcoids was supported. Putoidae, represented by Puto yuccae, was found to be outside the remainder of the neococcoid clade. These data are consistent with a single origin (in the ancestor of the neococcoid clade) of a chromosome system involving paternal genome elimination in males. Pseudococcidae (mealybugs) appear to be sister to the rest of the neococcoids and there are indications that Coccidae (soft scales) and Kerriidae (lac scales) are sister taxa. The Eriococcidae (felt scales) was not recovered as a monophyletic group and the eriococcid genus Eriococcus sensu lato was polyphyletic.     

An annotated checklist of the scale insects of Iran (Hemiptera,Sternorrhyncha, Coccoidea) with new records and distribution data

A list of scale insects (Hemiptera: Sternorrhyncha: Coccoidea) of Iran is present based mainly on the literature records since 1902. In total, 13 families and 275 species have been recorded and these are listed along with their locality data and host plants. The families are as follows: Asterolecaniidae, Cerococcidae, Coccidae, Diaspididae, Eriococcidae, Kermesidae, Margarodidae, Monophlebidae, Ortheziidae, Phoenicococcidae, Pseudococcidae, Putoidae and Rhizoecidae. The following ten species are recorded for the first time from Iran: Diaspidiotus lenticularis (Lindinger), Diaspidiotus wuenni (Lindinger), Fiorinia proboscidaria Green, Koroneaspis lonicerae Borchsenius, Eriococcus cingulatus Kiritchenko, Eriococcus pamiricus (Bazarov), Eriococcus reynei Schmutterer, Eriococcus sanguinairensis Goux and Eriococcus saxidesertus (Borchsenius) and Porphyrophora victoriae Jashenko.     

Biology and behavior of Metaphycus angustifrons Compere (Hymenoptera: Encyrtidae), a newly discovered parasitoid of soft scale insects (Hemiptera: Coccidae) in California

Metaphycus angustifrons Compere has recently been found to be the most abundant parasitoid of brown soft scale, Coccus hesperidum L., in southern California. In laboratory experiments we examined several biological parameters of this species. M. angustifrons both oviposits and host feeds in brown soft scale and is a facultatively gregarious endoparasitoid of this soft scale insect. In contrast with other Metaphycus spp., M. angustifrons is a koinobiont parasitoid, allowing its host to grow up to 40% beyond its size at parasitism. Despite its high abundance on brown soft scale in the field, in the laboratory, high rates of parasitoid egg encapsulation are observed; about half of parasitized hosts failed to issue parasitoids. Furthermore, host scales that encapsulated parasitoids eggs showed significant reduction in development. Increased scale size at oviposition influences the size of emerging females but not the size of males. Female M. angustifrons are synovigenic. They emerge from their hosts without mature eggs and begin maturing eggs after they are provided a carbohydrate source. Carbohydrates prolong the life span of both female and male M. angustifrons. The size of female wasps influences egg load but not longevity. Finally, based on laboratory observations, M. angustifrons uses citricola scale almost exclusively for host feeding and not for oviposition. These results suggest that the role of this species in citricola scale’s decline in southern California in the 1950s–1960s was negligible.     

我国发现一新入侵蜡蚧:三列鬃软蚧(半翅目,蚧总科,蜡蚧科)

本文记述采自中国惠州的1新记录属鬃软蚧属Trijuba De Lotto,该属目前只有1种,区别于蜡蚧科其他属的特征是雌成虫背面具有3列长刚毛。新记录种三列鬃软蚧Trijuba oculata(Brain)寄生于猴耳环Pithecellobium clypearia的枝干和叶柄上。     

中国克里蜡蚧属二新记录种（半翅目：蚧总科：蜡蚧科）

记述采自海南的克里蜡蚧属Kilifia 2新记录种:Kilifia deltoides De Lotto, 1965和K. americana Ben-Dov, 1979。对雌成虫形态特征进行了详细描述和绘图,提供了克里蜡蚧属中国已知种的检索表。研究标本保存在西北农林科技大学昆虫博物馆。     

Characterization of Burkholderia bacteria clade compositions in soil and Riptortus pedestris (Hemiptera: Alydidae) in South Korea

Riptortus pedestris (Hemiptera: Alydidae) is known to acquire the genus Burkholderia, symbiotic bacteria, from soil. Therefore, symbiont acquisition of R. pedestris would be directly affected by bacterial diversity in soil. Soil typically harbors diverse microbes including different Burkholderia clades such as SBE (stinkbug-associated beneficial and environmental), PBE (plant-associated beneficial and environmental), and BCC (Burkholderia cepacia and complex). Nevertheless, little is known about Burkholderia acquisition patterns of R. pedestris in nature, especially in the context of bacteria clade compositions in soil. Therefore, based on diagnostic PCR analysis, we investigated Burkholderia clade compositions in field-collected soil itself and R. pedestris when the insects were provided with the soil. Also, wild R. pedestris were surveyed to characterize their Burkholderia compositions. First, 88.44% of soil samples were detected with the genus Burkholderia, and triple clade (SBE + PBE + BCC) was most frequently detected. Second, R. pedestris nymphs readily acquired Burkholderia bacteria from field-collected soil where 91.25% of the reared insects harbored the bacteria in their midguts. In contrast to soil, the detection of single BCC clade was the most dominant among the three identified Burkholderia clades. Third, from wild R. pedestris, 80.62% of the insects were found to harbor the genus Burkholderia, and single BCC clade was most frequently detected. Finally, 29.13% and 47.06% of the reared and wild R. pedestris were detected with unidentified Burkholderia clade, which does not belong to any of the three identified clades. Our findings provide baseline information to better understand ecological associations between R. pedestris and Burkholderia bacteria in different clades.     

An out-of-body experience: the extracellular dimension for the transmission of mutualistic bacteria in insects

Across animals and plants, numerous metabolic and defensive adaptations are a direct consequence of symbiotic associations with beneficial microbes. Explaining how these partnerships are maintained through evolutionary time remains one of the central challenges within the field of symbiosis research. While genome erosion and co-cladogenesis with the host are well-established features of symbionts exhibiting intracellular localization and transmission, the ecological and evolutionary consequences of an extracellular lifestyle have received little attention, despite a demonstrated prevalence and functional importance across many host taxa. Using insect–bacteria symbioses as a model, we highlight the diverse routes of extracellular symbiont transfer. Extracellular transmission routes are unified by the common ability of the bacterial partners to survive outside their hosts, thereby imposing different genomic, metabolic and morphological constraints than would be expected from a strictly intracellular lifestyle. We emphasize that the evolutionary implications of symbiont transmission routes (intracellular versus extracellular) do not necessarily correspond to those of the transmission mode (vertical versus horizontal), a distinction of vital significance when addressing the genomic and physiological consequences for both host and symbiont.     

Higher‐level molecular phylogeny of jumping plant lice (Hemiptera: Sternorrhyncha: Psylloidea)

Jumping plant lice (Hemiptera: Psylloidea) are known for a few deleterious pest species worldwide, yet the phylogeny of the group has been poorly understood until very recently. Here, we reconstruct the higher‐level phylogeny for the superfamily Psylloidea based on multilocus DNA sequences, three mitochondrial (COI‐tRNA^leu‐COII, 12S, 16S) and five nuclear (18S, 28S D2, 28S D3, 28S D6–7a, 28S D9–10) gene fragments, using maximum likelihood and Bayesian inference phylogenetic frameworks. Our results are largely congruent with the recent phylogenomic study and partly support prior classification of Psylloidea based mainly on morphology, with the following major exceptions: the family Calophyidae is revealed as polyphyletic, Aphalaridae as paraphyletic with respect to most other taxa of Psylloidea, and Liviidae as paraphyletic with respect to Calophyinae, Psyllidae and Triozidae. Our phylogenetic hypothesis identifies Phacopteronidae and the genus Cecidopsylla Kieffer as the very basal taxa within extant Psylloidea. Sister‐group relationships of Rhinocolinae with Togepsyllinae and of Pachypsyllinae with Homotomidae are also suggested. We present specific discussions for each group of interest recovered in our phylogenetic analysis. One nomenclatorial change is proposed: Spanioneura longicauda (Konovalova) comb.n. , from Psylla Geoffroy.     

Natural biological control of green scale (Hemiptera: Coccidae): a field life-table study

Understanding how the biotic and abiotic factors influence pest-population dynamics is important to implement sound pest management strategies in biological control and integrated pest management (IPM) programmes. Coccus viridis (Green) is an important indirect pest of coffee plants, but very little has been done to understand the factors that contribute most for its biological control in the field. In the present study, we examined the critical life stage and the key factors associated with the mortality of C. viridis in coffee plantations in Brazil by conducting field-based life table studies. Predators, parasitoids, fungi, infested leaf abscission and rainfall were collectively responsible for a total C. viridis mortality of 96.08%. Predation by coccinellids was the key factor governing the mortality of C. viridis. The parasitism of early instars by parasitoids was the second most important factor contributing to C. viridis mortality. Unlike the parasitoids, the fungus Lecanicillium lecanii caused mortality of scales in more advanced life stages. The abscission of infested leaves from the trees, and rainfall also contributed to the mortality of C. viridis. The nymph stage was considered the critical stage for mortality of C. viridis in the field. The results suggest that predators (Coccinellidae) are the most important factors controlling C. viridis, and thus should be the target of conservation measures in coffee plantations infested with this pest.     

Discovery of cryptic species among North American pine‐feeding Chionaspis scale insects (Hemiptera: Diaspididae)

Cryptic species are present in many animal groups and they may be best detected through large sample sizes collected over broad geographic ranges. Fine‐scale local adaptation has been hypothesized to occur in armoured scale insects (Hemiptera: Diaspididae) and a consequence of this process may be multiple cryptic species. We estimate species diversity of pine‐feeding Chionaspis scale insects across North America by inferring species boundaries using genealogical concordance across allele genealogies of two nuclear loci and one mitochondrial locus. Our ingroup sample includes 366 individual insects from 320 localities and 51 host species within the Pinaceae. We also conducted a morphological survey of all insect specimens and assigned them as either one of the two currently recognized pine‐feeding species, Ch. heterophyllae, Ch. Pinifoliae, or with undescribed morphology. Using maximum likelihood allele genealogies in a majority‐rule consensus to assess congruence, we conservatively detect ten species in this group. Most of these species are robust to alternative methods of genealogical inference (Bayesian inference, maximum parsimony) and a few are robust to the use of strict consensus to assess congruence. Species show both narrow and more widespread ranges where almost half of the individuals sampled belong to a single very widespread polyphagous species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 47–62.     

Tests of male soft scale insects (Homoptera: Coccidae) from America north of Mexico, including a key to the species

Tests (pupal covers) of twenty-eight species of male Coccidae, representing fifteen genera from America north of Mexico, are described and illustrated. Tests of each of the species exhibit morphological differences which are useful in identification. Terminology is given for the sutures, corresponding plates, and related structures of the generalized test and a key is provided for separation of the described species.     

Cytoarchitecture of the ovarioles in scale insects (Hemiptera,Coccinea)

Telotrophic ovarioles of scale insects are subdivided into tropharia (=trophic chambers) and vitellaria that contain single developing oocytes. Tropharium encloses trophocytes (=nurse cells) and arrested oocytes. The central area of the tropharium, termed the trophic core, is devoid of cells. Both trophocytes and oocytes are connected to the trophic core: trophocytes by cytoplasmic processes, oocytes by means of nutritive cords. The trophic core, processes and nutritive cords are filled with bundles of microtubules. The trophocytes contain large lobated nuclei with giant nucleoli. Fluorescent labelling with DAPI has shown that trophocyte nuclei are characterized by high contents of DNA. In the cortical cytoplasm of trophocytes, numerous microfilaments are present. The developing oocyte is surrounded by a simple follicular epithelium. The cortical cytoplasm of follicular cells contains numerous microtubules and microfilaments.     

Cellular immune response of brown soft scale Coccus hesperidum L. (Hemiptera: Coccidae) to eggs of Metaphycus luteolus Timberlake (Hymenoptera: Encyrtidae)

The parasitic wasp, Metaphycus luteolus Timberlake, is an endoparasitoid of various soft scale insects including brown soft scale, Coccus hesperidum L. Development of this parasitoid in scale hosts is hindered by encapsulation. In the present study, using transmission and scanning electron microscopy, we show that hemocytes are responsible for encapsulation, which is mediated by the direct deposition of cells and melanin on the surface of M. luteolus eggs. By 12 h post-oviposition, scale hemocytes, presumably granulocytes, aggregate, spread and directly lyse on the surface of parasitoid eggs. This process continues for at least 1 day and results in the gradual formation of a capsule. Two to three days post-oviposition, a melanized capsule is well formed and signs of chemical deposition are evident by examination of the outer surface of the capsule. These results demonstrate that soft scale insects are fully capable of melanotic encapsulation of foreign material mediated by hemocytes.