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.     

Ground pearls: a generic revision of the Margarodidae sensu stricto (Hemiptera: Sternorrhyncha: Coccoidea)

The ground pearls comprise the scale insect family Margarodidae s. str., and are subterranean plant sucking parasites on roots of a wide variety of plants. Some ground pearls are serious destructive pests of grape vines, sugar cane, oil palms, cotton or lawn grass around the world. Both female and male development is characterised by an apodous, feeding 2^nd-instar nymph, called a cyst. The cyst is primarily circular and often colourful, shiny, metallic or pearl-like, from which the common name “ground pearls” is derived. Other unique features are the strongly developed prothoracic legs for digging, the construction of a protective test from their own liquid excreta in which the cyst is enclosed and a behavioral adaptation by modification of their life-cycle to survive in adverse environmental conditions.

The 10 genera, currently include 105 species, form a monophyletic group with a worldwide distribution. First-instar nymphs and cysts are feeding instars; however, adults male and females lack mouthparts and do not feed. Species of ground pearls reproduce either bisexually or parthenogenetically, parthenogenesis is facultative in Eurhizococcus brasiliensis. Females undergo three, four or five developmental stages, while male have five. During the cyst stage, one, two or three molts may occur. Most species have a single generation each year, although development in Margarodes vitis requires three years including three instars of cysts. This revision redescribes and illustrates the type species of each genus, including Margarodes vitis (Philippi) and provides a key to the genera based on the morphology of adult females. Lectotypes of Dimargarodes medíterraneus (Silvestri), Heteromargarodes americanus Jakubski, Promargarodes sinensis Silvestri and Termitococcus carrotai Silvestri are designated. A nomenclatural change is proposed, the genus Sphaeraspis Giard, 1894 is considered to be a synonym of Margarodes Guilding, 1829. This work also provides information on the history of the group of Archaeococcoids; on the biology, economic importance and distribution of the margarodids.     

The ovaries of aphids (Hemiptera,Sternorrhyncha, Aphidoidea): morphology and phylogenetic implications

The ovaries of aphids belonging to the families Eriosomatidae, Anoeciidae, Drepanosiphidae, Thelaxidae, Aphididae, and Lachnidae were examined at the ultrastructural level. The ovaries of these aphids are composed of several telotrophic ovarioles. The individual ovariole is differentiated into a terminal filament, tropharium, vitellarium, and pedicel (ovariolar stalk). Terminal filaments of all ovarioles join together into the suspensory ligament, which attaches the ovary to the lobe of the fat body. The tropharium houses individual trophocytes and early previtellogenic oocytes termed arrested oocytes. Trophocytes are connected with the central part of the tropharium, the trophic core, by means of broad cytoplasmic processes. One or more oocytes develop in the vitellarium. Oocytes are surrounded by a single layer of follicular cells, which do not diversify into distinct subpopulations. The general organization of the ovaries in oviparous females is similar to that of the ovaries in viviparous females, but there are significant differences in their functioning: (1) in viviparous females, all ovarioles develop, whereas in oviparous females, some of them degenerate; (2) the number of germ cells per ovariole is usually greater in females of the oviparous generation than in females of viviparous generations; (3) in oviparous females, oocytes in the vitellarium develop through three stages (previtellogenesis, vitellogenesis, and choriogenesis), whereas in viviparous females, the development of oocytes stops after previtellogenesis; and (4) in the oocyte cytoplasm of oviparous females, lipid droplets and yolk granules accumulate, whereas in viviparous females, oocytes accrue only lipid droplets. Our results indicate that a large number of germ cells per ovariole represent the ancestral state within aphids. This trait may be helpful in inferring the phylogeny of Aphidoidea.     

Phylogeny of ensign scale insects (Hemiptera: Coccoidea: Ortheziidae) based on the morphology of Recent and fossil females

The Ortheziidae (ensign scale insects) is a morphologically well‐defined family. The morphology and occurrence in the fossil record suggests a probable early origin of the family in scale insect evolution. The present phylogenetic analysis – based on 69 morphological characters of female ortheziids, using 39 exemplar Recent species – provides the first analytical assessment of relationships among living genera of the family, as well as the relationships of eight fossil species, based on complete, well‐preserved specimens in amber. Monophyly of the subfamilies Newsteadiinae, Ortheziinae and Ortheziolinae is supported, but Nipponortheziinae is found to be paraphyletic by inclusion of the Ortheziolinae. Thus, the subfamily Ortheziolinae is reduced in rank to tribe Ortheziolini stat.n. , which now includes Matileortheziola Kozár & Foldi, Ortheziolacoccus Kozár, Ortheziolamameti Kozár and Ortheziola?ulc. Consequently, the tribes Matileortheziolini, Ortheziolacoccini and Ortheziolamametini are synonymized ( syn.n. ) here under Ortheziolini. Five new species and one new genus of fossil ensign scales are described from three amber deposits: Burmorthezia gen.n. with type species Burmorthezia kotejai sp.n. and also B. insolita sp.n ., both in mid‐Cretaceous Burmese amber (98 Ma) and Arctorthezia baltica sp.n. in Eocene Baltic amber (c. 43 Ma) based on second‐instar nymphs; Mixorthezia kozari sp.n . and M. dominicana sp.n . in Miocene Dominican amber (c. 17 Ma) based on adult females. Fossil placements are unambiguous, with Burmorthezia forming a stem to crown‐group (Recent and Tertiary) Ortheziidae. A summary of described fossil ortheziids is provided.     

Two new species of Megacanthaspis Takagi (Hemiptera, Sternorrhyncha, Coccoidea, Diaspididae) from China

Two new species of armored scale, Megacanthaspis hangzhouensis Wei & Feng, sp. n. and Megacanthaspis hainanensis Wei & Feng, sp. n. are described and illustrated from specimens collected from China. A key to adult female of Megacanthaspis species is provided.     

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.     

A comprehensive phylogeny of extinct and extant Rhizomyinae (Rodentia): evidence for multiple intercontinental dispersals

The subfamily Rhizomyinae is known from the Late Oligocene up to the present. Today this group comprises six species, which live in southern Asia and eastern Africa. Despite the current moderate diversity of the rhizomyines, they had a greater diversification and wider distribution in the past: from Asia, their land of origin, to Africa, which they entered during the Early Miocene. So far 33 fossil species can be referred to this group. A cladistic analysis involving fossil and living species has been carried out. Prokanisamys spp. turned out to be the most basal taxa of the ingroup. This analysis calls into question the monophyly of several genera, and allows the proposal of a phylogenetic definition of the tribes Tachyoryctini and Rhizomyini. It also provides information about the origin of the African rhizomyines and allows inferring multiple dispersal phenomena from Asia to Africa in Early and Late Miocene times.     

Treehopper trees: phylogeny of Membracidae (Hemiptera: Cicadomorpha: Membracoidea) based on molecules and morphology

Abstract. Recent independent phylogenetic analyses of membracid relationships based on molecular and morphological data have identified monophyletic lineages within the family. However, the results of these studies have not fully resolved treehopper phylogeny, and relationships among some higher membracid lineages remain in doubt. Portions of three datasets (958 aligned nucleotides from elongation factor‐1α, 2363 aligned nucleotides from 28S ribosomal DNA, and eighty‐three morphological features of adults and nymphs) introduced in recent studies were reanalysed separately and in combination with two new molecular datasets (321 aligned nucleotides from wingless and 1829 aligned nucleotides from 18S ribosomal DNA). The results of the combined data analyses, contrary to previous analyses of morphological data alone, grouped membracids into two well‐supported lineages, one comprising Stegaspidinae and Centrotinae, the other comprising Membracinae, Darninae and Smiliinae. The analyses recovered Centrotinae, Membracinae and Darninae as monophyletic groups, but Stegaspidinae was paraphyletic with respect to Centrotinae, and Smiliinae was polyphyletic with Micrutalini placed as a sister group to the clade comprising Membracinae, Darninae and Smiliinae. These results are consistent with the following hypotheses, proposed previously based on an analysis of morphological data: (1) the posterior pronotal process was derived and lost multiple times during the evolution of Membracidae; (2) Membracidae originated in the New World and reached the Old World subsequently via dispersal; (3) maternal care evolved independently multiple times and may or may not have been preceded by the acquisition of ant mutualism.     

A phylogeny of extant penguins (Aves: Sphenisciformes) combining morphology and mitochondrial sequences

The phylogenetic relationships among the penguins have received little attention, despite their well‐known anatomy and the conspicuous nature of the group. Previous attempts have included datasets limited to few, mostly osteological characters, and one study was based on integumentary and breeding characters. We developed a morphological matrix comprising 159 morphological characters of osteology (70 characters), myology (15), digestive tract (1), integument (66), and breeding (7 characters), scored in 18 extant forms (all currently recognized species plus one distinct subspecies). A gaviiform was placed at the root, and 11 species of representative procellariiform groups completed the outgroup. A heuristic parsimony analysis under equal weights was performed. We also compiled DNA sequences available in GenBank for the mitochondrial genes 12S rDNA and cytochrome b. We included the two data partitions in a combined analysis under direct optimization. Both analyses recovered the monophyly of Sphenisciformes and all the traditional polytypic genera. Morphological characters performed optimally at the ordinal and generic nodes, also providing resolution and varying degrees of support at supra‐ and intrageneric nodes. The comparison of molecular and morphological results indicated that the most significant problem in the phylogeny of extant penguins is rooting the ingroup. The mutual interaction of molecular and morphological data decreases the ambiguity regarding the placement of the root, and provides a resolved, relatively well‐supported phylogeny of extant penguins. Biogeographical patterns based on breeding ranges and derived from the combined analysis show that the major intercontinental vicariance events detected are consistent with cold marine current patterns of the Southern Hemisphere. © The Willi Hennig Society 2005.     

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.     

The biology of extinct and extant sawfish (Batoidea: Sclerorhynchidae and Pristidae)

Sclerorhynchids (extinct sawfishes, Batoidea), pristids (extant sawfish, Batoidea) and pristiophorids (sawsharks, Squalomorphi) are the three elasmobranch families that possess an elongated rostrum with lateral teeth. Sclerorhynchids are the extinct sawfishes of the Cretaceous period, which reached maximum total lengths of 100 cm. The morphology of their rostral teeth is highly variable. Pristid sawfish occur circumtropically and can reach maximum total lengths of around 700 cm. All pristid species are globally endangered due to their restricted habitat inshore. Pristiophorid sawsharks are small sharks of maximum total lengths below 150 cm, which occur in depths of 70–900 m. Close examination of the morphology of pectoral fin basals and the internal structure of the rostrum reveals that sclerorhynchids and pristids evolved independently from rhinobatids, whereas pristiophorids are squalomorph sharks. The elongation of the rostrum may be an adaptation for feeding, as all marine vertebrate taxa that possess this structure are said to use it in the context of feeding.     

Extensive chromosomal variation associated with taxon divergence and host specificity in the gall-inducing scale insect Apiomorpha munita (Schrader) (Hemiptera: Sternorrhyncha: Coccoidea: Eriococcidae)

Apiomorpha Rubsaamen (Hemiptera: Coccoidea: Eriococcidae) is one of the most chromosomally diverse of all animal genera. There is extensive karyotypic variation within many of the morphologically defined species, including A. munita (Schrader) which is here reported to have diploid chromosome counts ranging from 6 to more than 100. Each of the three morphologically defined subspecies of A. munita also displays considerable chromosomal variation: A. m. tereticornuta Gullan (2 n = 6, 8, 20, 22 or 24), A. m. malleensis Gullan (2 n = 6, 20, 22, 24 or 26), and A. m. munita (Schrader) (2 n = 54 or >100). Apiomorpha munita appears to occur only on eucalypts of the informal subgenus Symphyomyrtus , with each of the subspecies of A. munita restricted to discrete symphyomyrt sections. Several different karyotypic forms within each subspecies of A. munita appear to be restricted to only one or a few eucalypt species or series. The association between apparent host specificity and chromosomal rearrangements in A. munita suggests that both may be playing an active role in taxon divergence in Apiomorpha.     

Some species of the hypogeal scale insect Porphyrophora Brandt (Hemiptera: Sternorrhyncha: Coccoidea: Margarodidae) from Europe,the Middle East and North Africa

Porphyrophora is one of several hypogeal genera in the scale insect family Margarodidae (Hemiptera: Sternorrhyncha: Coccoidea). Historically, it has been of great economic importance as a source of carmine dye but has been superseded by artificial aniline dyes. Some Porphyrophora species are currently important crop pests, particularly in western Asia. However, adult female Porphyrophora are all remarkably uniform in structure and are difficult to identify. The present paper redescribes the adult females of 24 species of Porphyrophora from Europe, the Middle East and North Africa and describes 3 new species (P. chelodonta Vahedi, P. jakubskii Vahedi and P. jashenkoi Vahedi). A key is provided to separate the species of Porphyrophora covered here. It is considered that P. cynodontis (Archangelskaia) is probably a synonym of P. hamelii Brandt although this synonymy is not being formally introduced here.     

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.     

Insect, not plant, determines gall morphology in the Apiomorpha pharetrata species-group (Hemiptera: Coccoidea)

Abstract  Scale insects of the genus Apiomorpha Rübsaamen (Hemiptera: Coccoidea: Eriococcidae) induce sexually dimorphic galls on Eucalyptus, but the Apiomorpha pharetrata species-group is unusual in that nymphal males aggregate on the surface of the maternal gall where they induce a compound structure. Although originally described as distinct species on the basis of differences in gall morphology, A. pharetrata (Schrader) and A. thorntoni (Froggatt) were subsequently synonymised, primarily because the adult females of the two are morphologically indistinguishable. The two gall types of A. pharetrata sensu lato are allopatric and found on different host eucalypt species. To test the hypothesis that gall morphology may be determined by the host species on which the female feeds, we reared crawlers of field-collected females from both gall types on eucalypts in a glasshouse. Both gall types were induced on the same eucalypt species, with the gall type matching that of the maternal gall from which the crawlers had emerged. This indicates that it is the insect, not the host, that determines the gall morphology in these taxa. In addition, insects from the two gall types were different chromosomally and could be distinguished by one fixed allozyme difference. Thus, Brachyscelis (=  Apiomorpha ) thorntoni (Froggatt) revised status is removed from synonymy with Brachyscelis (=  Apiomorpha ) pharetrata (Schrader), and recognised as a distinct biological species. However, the extent of chromosomal variation among the populations of both A. pharetrata and A. thorntoni suggests that there may be further cryptic species present.     

宁夏粉蚧科昆虫研究（半翅目：蚧总科）

记述宁夏同族自治区粉蚧科Pseudococcidae昆虫24属41种,其中包括2新种:宁夏草粉蚧Fonscolombia ningxiana,sp.nov.和冰草长粉蚧Longicoccus agropyri.sp.nov.;2中国新记录种:多管刺粉蚧spinococcus multitubulatus(Danzig,1980)和孤独条粉蚧Trionymus singularis Schmutterer,1952:1新组合:赖草长粉蚧Longicoccus leymicola(Tang,1992),comb.n.(移自少粉蚧属Mirococcus Borchsenius);Puto jarudensis Tang(1992)为Ceroputopilosellae Sulc.1898的新异名.模式标本保存在山西农业大学蚧虫研究中心.     

The gall-inducing habit has evolved multiple times among the eriococcid scale insects (Sternorrhyncha: Coccoidea: Eriococcidae)

The habit of inducing plant galls has evolved multiple times among insects but most species diversity occurs in only a few groups, such as gall midges and gall wasps. This phylogenetic clustering may reflect adaptive radiations in insect groups in which the trait has evolved. Alternatively, multiple independent origins of galling may suggest a selective advantage to the habit. We use DNA sequence data to examine the origins of galling among the most speciose group of gall-inducing scale insects, the eriococcids. We determine that the galling habit has evolved multiple times, including four times in Australian taxa, suggesting that there has been a selective advantage to galling in Australia. Additionally, although most gall-inducing eriococcid species occur on Myrtaceae, we found that lineages feeding on Myrtaceae are no more likely to have evolved the galling habit than those feeding on other plant groups. However, most gall-inducing species-richness is clustered in only two clades ( Apiomorpha and Lachnodius  +  Opisthoscelis ), all of which occur exclusively on Eucalyptus s.s . The Eriococcidae and the large genus Eriococcus were determined to be non-monophyletic and each will require revision.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 441–452.     

双孔皮珠蚧形态学研究（半翅目：蚧总科：珠蚧科）

重描了双孔皮珠蚧Kuwania bipora Borchsenius, 1960 的雌成虫,首次记述和图示了各龄若虫和雄成虫的形态特征,并提供了分龄检索表。