The phylogeny and revised classification of Machaerotidae,the tube‐making spittlebugs (Hemiptera: Auchenorrhyncha: Cercopoidea)

Machaerotidae (Hemiptera: Auchenorrhyncha: Cercopoidea) is a taxonomically small but morphologically diverse family of spittlebugs with approximately 115 described species in 31 genera and an exclusively Palaeotropical distribution. Results are presented of the first molecular phylogenetic investigation of Machaerotidae, examining relationships among the currently recognized subfamilies and tribes, as well as determining the phylogenetic placement of the genera Enderleinia Schmidt, Neuromachaerota Schmidt, Labramachaerota Bell & Cryan, and Kyphomachaerota Bell & Cryan. DNA nucleotide sequence data from eight loci (12s rDNA, 16s rDNA, 18S rDNA, 28S rDNA, histone 2A, histone 3, wingless and NADH Dehydrogenase subunit 4) were analysed to reconstruct the phylogeny. The evidence generated in this study supports the following systematic conclusions: (i) Machaerotidae is a monophyletic family; (ii) Machaerotini, Hindoloidini (with the new inclusion of Kyphomachaerota), and Enderleiniini (excluding Kyphomachaerota and Apomachaerota Schmidt) are monophyletic tribes; (iii) the genus Apomachaerota was recovered as the most anciently diverged lineage of extant Machaerotidae, and a new subfamily (Apomachaerotinae subfam.n. ), is proposed on the basis of its phylogenetic placement as sister lineage to all other extant Machaerotidae.     

Molecular phylogeny of Cicadomorpha (Insecta: Hemiptera: Cicadoidea, Cercopoidea and Membracoidea): adding evidence to the controversy

Abstract. The hemipteran infraorder Cicadomorpha comprises the superfamilies Cicadoidea (cicadas), Cercopoidea (spittlebugs or froghoppers) and Membracoidea (leafhoppers and treehoppers). Earlier attempts to determine relationships among these three monophyletic lineages using either morphological or molecular data suffered from insufficient sampling (taxonomic and data) and problematic tree rooting, leading to discordant results. Presented here are phylogenetic reconstructions within Cicadomorpha based on DNA nucleotide sequence data from multiple genetic markers (18S rDNA, 28S rDNA, and histone 3) sequenced from representative taxa of Cicadidae, Tettigarctidae, Cercopidae, Aphrophoridae, Clastopteridae, Machaerotidae, Epipygidae, Cicadellidae, Membracidae, Myerslopiidae and Aetalionidae. To test the robustness of the phylogenetic signal, these sequence data were analysed separately and in combination under various alignment parameters using both manual alignment (of both attenuated and full sequences) and alignment via clustal x . The results demonstrate clearly that, despite the alignment method used, basing a phylogeny on a single gene region is often misleading. Analyses of the combination of datasets support the major relationships within Cicadomorpha as (Membracoidea (Cicadoidea, Cercopoidea)). Internal relationships recovered within each superfamily shows evidence for: (1) the placement of Myerslopiidae as the sister group of the remaining Membracoidea; (2) the paraphyly of Cicadellidae; (3) the sister-group relationship between Machaerotidae and Clastopteridae; (4) the monophyly of Cercopidae; (5) the diversification of Epipygidae from within the possibly paraphyletic Aphrophoridae.     

Review of the Australian aphrophorid spittlebugs (Hemiptera: Aphrophoridae)

Abstract A review of the Aphrophoridae (Hemiptera: Auchenorrhyncha: Cercopoidea) occurring in Australia has revealed closer links between the Australian fauna and those of the Pacific region than were previously acknowledged. A key to the Australian genera is provided along with a diagnosis for each genus and a list of described Australian species with notes on each. Taxonomic changes proposed are: Neoaphrophora China synonymised with Anyllis Kirkaldy; Eurycercopis Kirkaldy synonymised with Interocrea Walker; Aphrophora rufiventris Walker and Philagra vittata Metcalf and Horton synonymised with Philagra parva (Donovan). New combinations proposed are: Amarusa australis (Jacobi) (from Eoptyelus ); Anyllis tiegsi (China) (from Neoaphrophora ); Interocrea nigrofasciata (Kirkaldy) (from Eurycercopis ); Interocrea regalis (Lallemand) (from Clovia ); Liorhina loxosema (Hacker) (from Clovia ). To provide a single reference point for the species and, for species described by Hacker (1926) and Jacobi (1928) , to validate apparent holotype intentions by those authors, lectotypes are designated for: Ptyelus homochrous Hacker; Eurycercopis nigrofasciata Kirkaldy; Clovia regalis Lallemand; Clovia loxosema Hacker; Philagra concolor Hacker; Philagra fulvida Hacker; Philagra recurva Jacobi. The probable whereabouts of the type specimen of Philagra parva (Donovan) are discussed. The origin of the holotype of Novaphrophara tasmaniae Lallemand is discussed; the Australian record is almost certainly incorrect.     

Phylogenetic relationships among superfamilies of Cicadomorpha (Hemiptera: Auchenorrhyncha) inferred from the wing base structure

The infraorder Cicadomorpha is a monophyletic group of the order Hemiptera, suborder Auchenorrhyncha, and is composed of three superfamilies: Cercopoidea (spittle bugs), Cicadoidea (cicadas) and Membracoidea (leafhoppers and treehoppers). Phylogenetic relationships among the superfamilies have been highly controversial morphologically and molecularly, but recent molecular phylogenetic analyses provided support for Cercopoidea + Cicadoidea. In this study, we examined morphology of the wing base structure in Cicadomorpha and tested the previous phylogenetic hypotheses using the characters selected from the wing base. As a result, a sister‐group relationship between Cicadoidea and Cercopoidea was supported by three synapomorphies (presence of a projection posterior to the anterior notal wing process, presence of a novel notal process anterior to the posterior notal wing process, presence of a novel sclerite between the distal median plate and the base of anal vein). The present study provides the first unambiguous and prominent morphological support for Cicadoidea + Cercopoidea.     

半翅目沫蝉总科化石研究进展

回顾半翅目沫蝉总科化石研究简史,列出已发表的化石名录、地理分布及地质年代,共6科、117属、252种;概述了沫蝉总科的地质历史,总结了世界沫蝉总科化石的组成,提出该类群昆虫化石研究中存在的问题,并对今后的研究前景进行展望.     

Morphology-based phylogeny of the treehopper family Membracidae (Hemiptera: Cicadomorpha: Membracoidea)

A parsimony‐based phylogenetic analysis of eighty‐three morphological characters of adults and immatures of seventy representatives of the tribes and subfamilies of Membracidae and two outgroup taxa was conducted to evaluate the status and relationships of these taxa. Centrotinae apparently gave rise to Nessorhinini and Oxyrhachini (both formerly treated as subfamilies, now syn.n. and syn.reinst., respectively, of Centrotinae). In contrast to previous analyses, a clade comprising Nicomiinae, Centronodinae, Centrodontinae, and the unplaced genera Holdgatiella Evans, Euwalkeria Goding and Antillotolania Ramos was recovered, but relationships within this clade were not well resolved. Nodonica bispinigera, gen.n. and sp.n., is described and placed in Centrodontini based on its sister‐group relationship to a clade comprising previously described genera of this tribe. Membracinae and Heteronotinae were consistently monophyletic. Neither Darninae nor Smiliinae, as previously defined, was monophyletic on the maximally parsimonious cladograms, but constraining both as monophyletic groups required only one additional step. The monophyly of Stegaspidinae, including Deiroderes Ramos (unplaced in Membracidae), was supported on some but not all equally parsimonious cladograms. More detailed analyses of individual subfamilies, as well as morphological data on the undescribed immatures of several membracid tribes and genera, will be needed to elucidate relationships among tribes and genera. A key to the subfamilies and tribes is provided.     

New Jurassic Cercopoidea from China and their evolutionary significance (Insecta: Hemiptera)

Abstract: A new family, Sinoalidae Wang and Szwedo fam. nov., is described from the Middle Jurassic Daohugou Biota (Inner Mongolia, China). Two new genera with three new species (Sinoala parallelivena Wang and Szwedo gen. et sp. nov., Jiania crebra Wang and Szwedo gen. et sp. nov., and Jiania gracila Wang and Szwedo sp. nov.) are erected based on well‐preserved whole‐bodied specimens. The morphological characters of Procercopidae and the new family are given, and a key to the species of Sinoalidae is presented. The genera Luanpingia Hong, 1983 and Huabeicercopis Hong, 1983 are revised and attributed to Sinoalidae. The genera Mesocercopis Hong, 1983 and Sinotettegarcta Hong, 1986 are considered junior synonyms of Anthoscytina Hong, 1983. Taxa assigned to Sinoalidae fam. nov. cannot be attributed to any previously known family, but share some general (plesiomorphic) characters with ancient Clypeata, for example, postclypeus distinctly swollen, transversely wrinkled, antenna with a flagellum of a few elongate segments, frons with median ocellus and lateral ocelli on crown. Moreover, the new family is closely related to Procercopidae (Cercopoidea) based on a pronotum with a median incision at the hind margin; tegmen slender, partly punctuate, basal portion of Sc not exceeding apex of basal cell, and hind legs with lateral spines. It has a mixture of ancestral characters shared with Hylicelloidea and Jurassic Procercopidae and some derived characters. Therefore, it is tentatively placed in Cercopoidea. A preliminary phylogram of Clypeata is presented based on the combination of fossil and morphological data.     

Wing base morphology of Aetalionidae (Hemiptera: Cicadomorpha) and its phylogenetic implications

The Aetalionidae is a small family belonging to the treehopper superfamily Membracoidea (Hemiptera: Cicadomorpha). Although the wing‐base morphology of Cicadomorpha was examined in detail recently, the wing base of this family has not been investigated to date. We examined morphology of the wing‐base structure of Aetalionidae. Using the characters selected from the wing base, we inferred the phylogenetic placement of this family and confirmed that it belongs to the superfamily Membracoidea and is likely a sister group of the Membracidae.     

Cicadomorpha community (Hemiptera: Auchenorrhyncha) in different agroecosystems in the north of Portugal

1. The infraorder Cicadomorpha is a diverse group comprising several species considered important pests of economic crops and species that may act as vectors of plant pathogens. In Europe, the gram-negative bacterium Xylella fastidiosa is one of the most important and severe insect-borne plant pathogens associated with the infraorder Cicadomorpha.
2. Therefore, the knowledge of the abundance and diversity of native Cicadomorpha insect vectors related to the different agroecosystems is essential to design and implementing specific measures to control insect-borne plant pathogens.
3. In this work, in two consecutive years (2018 and 2019), five almond orchards, five vineyards, five olive orchards, and five scrublands distributed in the North of Portugal were sampled in three different periods (early summer, summer and autumn) to investigate species composition, richness, and diversity of the Cicadomorpha community. Also, in 2019, five lemon orchards were sampled.
4. A total of 6056 individuals were collected (2322 in 2018 and 3734 in 2019), belonging to 71 species of three families. Observing several considered pests or vectors of vicious pathogens within these species was possible. The confirmed vectors of X. fastidiosa (Philaenus spumarius (Linnaeus, 1758) and Neophilaenus campestris (Fallen, 1805)) were captured in all agroecosystems and, in general, with more abundance in autumn. The highest abundance, richness, and diversity of Cicadomorpha were observed in vineyards. However, these parameters (abundance, richness and diversity) significantly differed between the agroecosystem and sampling period.
5. Further research on how the composition of the vegetation cover shapes the Cicadomorpha community is essential to implement strategies to reduce the spread of insect-borne pathogens if they are introduced into agroecosystems.

     

Sperm morphology in five species of cicadettine cicadas (Hemiptera: Cicadomorpha: Cicadidae)

Mature spermatozoa from five species of cicadas of the subfamily Cicadettinae (Quintilia wealei, Melampsalta leucoptera, Stagira simplex, Xosopsaltria thunbergi and Monomatapa matoposa) were examined by light and electron microscopy. In each species sperm are elongate, aggregated into organized bundles with their heads embedded in a homogenous matrix to form spermatodesmata, and exhibit polymegaly. The head of the sperm consist of an anteriorly positioned conical acrosome that has a tubular substructure and a deep, posterior invagination that forms the subacrosomal space (eccentrically positioned anteriorly). The acrosome is flattened anteriorly; posteriorly it extends along either side of the nucleus as two tubular processes that gradually decrease in diameter. The filiform nucleus tapers anteriorly and intrudes into the subscrosomal space. Posteriorly the nucleus has a lateral invagination that houses material of the so-called centriolar adjunct. Posterior to the centriolar adjuct and the nucleus are two crystalline mitochondrial derivatives and a centriole, respectively, the latter giving rise to the axoneme, which has a 9 + 9 + 2 arrangement of microtubules. In these respects the sperm are similar to those of platypleurine cicadas. However, some features seem unique to cicadettines, including the structural organization of an enlarged centriolar adjunct and the dimensions of the tails. The enlarged centriolar adjunct has a lamella-like substructure and can be considered a synapomorphic character in the Cicadettinae. It is, therefore, potentially useful in the separation of this subfamily from the Cicadinae. In addition, the great length of the sperm nucleus of long-headed sperm in M. matoposa could be a synapomorphy of this genus and related taphurine and cicadettine species.     

Morphology of the mouthparts of the spittlebug Philagra albinotata Uhler (Hemiptera: Cercopoidea: Aphrophoridae)

Mouthparts associated with feeding behavior and feeding habits are important sensory and feeding structures in insects. To obtain a better understanding of feeding in Cercopoidea, the morphology of mouthparts of the spittlebug, Philagra albinotata Uhler was examined using scanning electron microscopy. The mouthparts of P. albinotata are of the typical piercing–sucking type found in Hemiptera, comprising a cone-shaped labrum, a tube-like, three-segmented labium with a deep groove on the anterior side, and a stylet fascicle consisting of two mandibular and two maxillary stylets. The mandibles consist of a dorsal smooth region and a ventral serrate region near the apical half of the external convex region, and bear five nodules or teeth on the dorsal external convex region on the distal extremity; these are regarded as unique features that distinguish spittlebugs from other groups of Hemiptera. The externally smooth maxillary stylets, interlocked to form a larger food canal and a smaller salivary canal, are asymmetrical only in the internal position of longitudinal carinae and grooves. One dendritic canal is found in each maxilla and one in each mandible. Two types of sensilla trichodea, three types of sensilla basiconica and groups of multi-peg structures occur in different locations on the labium, specifically the labial tip with two lateral lobes divided into anterior sensory fields with ten small peg sensilla arranged in a 5 + 4 + 1 pattern and one big peg sensillum, and posterior sensory fields with four sensilla trichodea. Compared with those of previously studied Auchenorrhyncha, the mouthparts of P. albinotata may be distinguished by the shape of the mandibles, the multi-peg structures and a tooth between the salivary canal and the food canal on the extreme end of the stylets. The mouthpart morphology is illustrated using scanning electron micrographs, and the taxonomic and putative functional significance of the different structures is briefly discussed.     

陕西省蝉科二新种（半翅目: 蝉亚目）

描述了1936年6月采自陕西省的蝉科两新种： Neotanna shensiensis sp.n. 和 Terpnosia. shaanxiensis sp.n.。正模标本保存在北卡罗来纳州立大学昆虫标本馆(NCSU); 副模保存在作者处。     

A remarkable new genus of Procercopidae (Hemiptera: Cercopoidea) from the Middle Jurassic of China

Jurocercopis grandis, a new genus and species belonging to Procercopidae (Hemiptera: Cercopoidea), is described, based on seven specimens from the Middle Jurassic of Daohugou, China. It differs distinctly from other genera as follows: body and wings obviously large; pronotum with posterior margin straight; tegmen with costal margin convex at about basal 0.4 wing length; hind basitarsomere with a row of apical small teeth.     

Sperm morphology in four species of African platypleurine cicadas (Hemiptera: Cicadomorpha: Cicadidae)

Mature spermatozoa from four species of platypleurine cicadas (Albanycada albigera, Azanicada zuluensis, Platypleura capensis and P. hirtipennis) were examined by light and electron microscopy. The filiform sperm have a similar ultrastructure in all species but notable variations were found in sperm dimensions. All species produce more than one discrete length of nucleated, motile sperm, a form of polymorphism termed polymegaly. Polymegaly is expressed in two ways: sperm have bi- or trimodal head and tail lengths. The anterior parts of sperm heads are embedded in an elongate homogenous matrix forming a spermatodesm. The conical acrosome is deeply invaginated posteriorly, and sits on top of the nucleus. The acrosomal contents are differentiated internally with an electron-lucent central medulla and a denser cortex. The homogenously electron-dense nucleus is pointed anteriorly and is generally cylindrical, although posteriorly there is a lateral invagination that extends part-way along the nucleus. This invagination houses fine granular material of the centriolar adjunct. Vesicle-like elements that are associated with both the posterior nucleus and the centriolar adjunct are also found within the invagination. Immediately posterior of and adjoining the centriolar adjunct is a pair of mitochondrial derivatives that are elongated and extend for almost the entire length of the tail. The absence of accessory bodies in cicada sperm suggests that within the Cicadomorpha, the families Cicadidae and Cercopidae are closely affiliated.     

Antifungal effect and chitinase activities of the froth of spittlebug Poophilus costalis (Walker) (Hemiptera: Cercopoidea: Aphrophoridae)

Nymphs of xylophagous spittlebug Poophilus costalis (Walker, 1851) produce froth that covers their entire body surface. However, no mold has ever been found on this froth under natural conditions. This study has shown that the froth of P. costalis can repress the growth of Fusarium oxysporum f. sp. pisi. Three chitinases, i.e., β-N-acetylglucosaminidase, chitobiosidase, and endochitinase, with specific activities of 1.759, 0.365, and 1.172 U/mg protein, respectively, and a protease with minimal activity have been detected in this froth. β-N-actylglucosaminidase activity was also detected in the source of the froth, the nymphal Malpighian tubules and the host plant Commelina diffusa Burm. f. These findings suggest that the antifungal effect of the froth from P. costalis may be attributed to the chitinase produced by its Malpighian tubules and/or its host plant.     

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.     

Anatomy and fine structure of the alimentary canal of the spittlebug Lepyronia coleopterata (L.) (Hemiptera: Cercopoidea)

The alimentary canal of the spittlebug Lepyronia coleopterata (L.) differentiates into esophagus, filter chamber, midgut (conical segment, tubular midgut), and hindgut (ileum, rectum). The filter chamber is composed of the anterior extremity of the midgut, posterior extremity of the midgut, proximal Malpighian tubules, and proximal ileum; it is externally enveloped by a thin cellular sheath and thick muscle layers. The sac-like anterior extremity of the midgut is coiled around by the posterior extremity of the midgut and proximal Malpighian tubules. The tubular midgut is subdivided into an anterior tubular midgut, mid-midgut, posterior tubular midgut, and distal tubular midgut. Four Malpighian tubules run alongside the ileum, and each terminates in a rod closely attached to the rectum. Ultrastructurally, the esophagus is lined with a cuticle and enveloped by circular muscles; its cytoplasm contains virus-like fine granules of high electron-density. The anterior extremity of the midgut consists of two cellular types: (1) thin epithelia with well-developed and regularly arranged microvilli, and (2) large cuboidal cells with short and sparse microvilli. Cells of the posterior extremity of the midgut have regularly arranged microvilli and shallow basal infoldings devoid of mitochondria. Cells of the proximal Malpighian tubule possess concentric granules of different electron-density. The internal proximal ileum lined with a cuticle facing the lumen and contains secretory vesicles in its cytoplasm. Dense and long microvilli at the apical border of the conical segment cells are coated with abundant electron-dense fine granules. Cells of the anterior tubular midgut contain spherical secretory granules, oval secretory vesicles of different size, and autophagic vacuoles. Ferritin-like granules exist in the mid-midgut cells. The posterior tubular midgut consists of two cellular types: 1) cells with shallow and bulb-shaped basal infoldings containing numerous mitochondria, homocentric secretory granules, and fine electron-dense granules, and 2) cells with well-developed basal infoldings and regularly-arranged apical microvilli containing vesicles filled with fine granular materials. Cells of the distal tubular midgut are similar to those of the conical segment, but lack electron-dense fine granules coating the microvilli apex. Filamentous materials coat the microvilli of the conical segment, anterior and posterior extremities of the midgut, which are possibly the perimicrovillar membrane closely related to the nutrient absorption. The lumen of the hindgut is lined with a cuticle, beneath which are cells with poorly-developed infoldings possessing numerous mitochondria. Single-membraned or double-membraned microorganisms exist in the anterior and posterior extremities of the midgut, proximal Malpighian tubule and ileum; these are probably symbiotic.     

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.     

Family‐level variation in early life‐cycle traits of kelp

Temperate kelp forests (Laminarians) are threatened by temperature stress due to ocean warming and photoinhibition due to increased light associated with canopy loss. However, the potential for evolutionary adaptation in kelp to rapid climate change is not well known. This study examined family‐level variation in physiological and photosynthetic traits in the early life‐cycle stages of the ecologically important Australasian kelp Ecklonia radiata and the response of E. radiata families to different temperature and light environments using a family × environment design. There was strong family‐level variation in traits relating to morphology (surface area measures, branch length, branch count) and photosynthetic performance (F_v/F_m) in both haploid (gametophyte) and diploid (sporophyte) stages of the life‐cycle. Additionally, the presence of family × environment interactions showed that offspring from different families respond differently to temperature and light in the branch length of male gametophytes and oogonia surface area of female gametophytes. Negative responses to high temperatures were stronger for females vs. males. Our findings suggest E. radiata may be able to respond adaptively to climate change but studies partitioning the narrow vs. broad sense components of heritable variation are needed to establish the evolutionary potential of E. radiata to adapt under climate change.