An annotated checklist of the planthoppers of Iran (Hemiptera, Auchenorrhyncha, Fulgoromorpha) with distribution data

A list of Hemiptera Fulgoromorpha (planthoppers) of Iran is provided, based primarily on literature records from 1902 to the present. In total 15 families and 235 species are recorded, with taxonomic details. Distribution data in Iran are given. Iranissus ephedrinus Dlabola, 1980 is transferred from Issidae to Nogodinidae. To resolve nomenclatural difficulty the following new combinations in Issidae are given: Iranodus dumetorus (Dlabola, 1981), Iranodus khatunus (Dlabola, 1981) and Iranodus repandus (Dlabola, 1981). Due to published generic synonomy the following are new combinations: Duilius seticulosus (Lethierry, 1874), Duilius tamaricis (Puton & Lethierry, 1887), Duilius tamaricicola (Dubovsky, 1966) and Duilius v-atrum (Dlabola, 1985).     

A Molecular Phylogeny of Hemiptera Inferred from Mitochondrial Genome Sequences

Classically, Hemiptera is comprised of two suborders: Homoptera and Heteroptera. Homoptera includes Cicadomorpha, Fulgoromorpha and Sternorrhyncha. However, according to previous molecular phylogenetic studies based on 18S rDNA, Fulgoromorpha has a closer relationship to Heteroptera than to other hemipterans, leaving Homoptera as paraphyletic. Therefore, the position of Fulgoromorpha is important for studying phylogenetic structure of Hemiptera. We inferred the evolutionary affiliations of twenty-five superfamilies of Hemiptera using mitochondrial protein-coding genes and rRNAs. We sequenced three mitogenomes, from Pyrops candelaria, Lycorma delicatula and Ricania marginalis, representing two additional families in Fulgoromorpha. Pyrops and Lycorma are representatives of an additional major family Fulgoridae in Fulgoromorpha, whereas Ricania is a second representative of the highly derived clade Ricaniidae. The organization and size of these mitogenomes are similar to those of the sequenced fulgoroid species. Our consensus phylogeny of Hemiptera largely supported the relationships (((Fulgoromorpha,Sternorrhyncha),Cicadomorpha),Heteroptera), and thus supported the classic phylogeny of Hemiptera. Selection of optimal evolutionary models (exclusion and inclusion of two rRNA genes or of third codon positions of protein-coding genes) demonstrated that rapidly evolving and saturated sites should be removed from the analyses.     

“Issidisation” of Fulgoroid planthoppers (Homoptera, Fulgoroidea) as a case of parallel adaptive radiation

The term “issidisation” is proposed to describe the formation of the specific box-like body shape (including elytra) in several families of planthoppers (Acanaloniidae, Caliscelidae, Issidae, Nogodinidae, and Tropiduchidae). Parallel radiation resulting from adaptation to semi-arid and arid conditions is considered the cause of issidisation. The possibly rapid rate of evolution of higher Fulgoroidea and especially Issidae is discussed. Pyrogenic renewal of Mediterranean-type communities is hypothesized to be the cause of sympatric speciation in the Mediterranean insects.     

Neotapirissus gen. nov. of the tribe Issini (Hemiptera: Fulgoromorpha: Issidae) from Hainan Island

Neotapirissus gen. nov. , in the tribe Issini (Hemiptera: Fulgoromorpha: Issidae), is described from China, and type species Neotapirissus reticularis sp. nov. (China: Hainan) is described and illustrated. The female internal reproductive organs of the new species are also described and illustrated. The new genus is compared to the superficially similar genus Tapirissus Gnezdilov. The morphological diversity of the issids distributed in Hainan Island is discussed briefly.     

蜡蝉次目(昆虫纲:半翅目)昆虫化石研究进展

近200年来,全世界共报道半翅目蜡蝉次目化石昆虫28科、216属、400余种.本文回顾了蜡蝉次目化石昆虫的研究历史,统计了中国已报道的该类群化石种类名录,总结了蜡蝉次目化石昆虫的地质历史、地理分布和年代、以及属种组成特征,提出了该类群化石昆虫研究中有待解决的问题以及未来的研究前景.     

The function and phylogenetic implications of the tentorium in adult Neuroptera (Insecta)

Despite several recent analyses on the phylogeny of Neuroptera some questions still remain to be answered. In the present analysis we address these questions by exploring a hitherto unexplored character complex: the tentorium, the internal cuticular support structure of the insect head. We described in detail the tentoria of representatives of all extant neuropteran families and the muscles originating on the tentorium using 3D microCT images and analyzed differences in combination with a large published matrix based on larval characters. We find that the tentorium and associated musculature are a source of phylogenetically informative characters. The addition of the tentorial characters to the larval matrix causes a basad shift of the Sisyridae and clearly supports a clade of all Neuroptera except Sisyridae and Nevrorthidae. A sister group relationship of Coniopterygidae and the dilarid clade is further corroborated. A general trend toward a reduction of the dorsal tentorial arms and the development of laminatentoria is observed. In addition to the phylogenetic analysis, a correlation among the feeding habits, the development of the maxillary muscles, and the laminatentoria is demonstrated.     

Phylogeny and classification of the Stenophlebioptera (Odonata: Epiproctophora)

The Juraheterophlebiidae, new family of the “heterophlebioid” lineage, the Henrotayiidae, new family of the “anisopteroid” lineage, the Prostenophlebiidae and the Liassostenophlebiidae, new families of the Stenophlebioptera, and three new genera and species of the Stenophlebiidae are described from the Mesozoic of Germany, Spain, England, Kazakhstan, and Mongolia. The phylogenetic positions of the families Erichschmidiidae and Gondvanogomphidae are discussed. A tentative phylogenetic analysis of the Anisopteromorpha is proposed. This significantly extends our knowledge on the palaeogeographical distribution of the Stenophlebioptera and the Epiproctophora (“dragondamselflies”).     

A Preliminary Molecular Phylogeny of Planthoppers (Hemiptera: Fulgoroidea) Based on Nuclear and Mitochondrial DNA Sequences

The planthopper superfamily Fulgoroidea (Insecta: Hemiptera) is one of the most dominant groups of phytophagous insects. It comprises about 20 families, containing a total of 9000 species worldwide. Despite several recent studies, the phylogeny of Fulgoroidea is not yet satisfactorily resolved and the phylogenetic positions of several key families, especially Cixiidae, Delphacidae, Tettigometridae, Nogodinidae, Acanaloniidae and Issidae, are contentious. Here, we expand upon recent phylogenetic work using additional nuclear (18S and 28S) and novel mitochondrial (16S and cytb) markers. Maximum likelihood and Bayesian analyses yielded robust phylogenetic trees. In these topologies, a group containing Cixiidae and Delphacidae is recovered as the sister group to the remaining taxa. Tettigometridae is placed in a more nested position and is grouped with Caliscelidae. Sister relationships are found between Flatidae and Ricaniidae, and between Dictyopharidae and Fulgoridae. Nogodinidae and Issidae are confirmed to be non-monophyletic families. For major nodes of interest, divergence date estimates are generally older than those from the fossil record.     

Development of the mouthparts in embryos of Haplothrips verbasci (Osborn) (Insecta,Thysanoptera, Phlaeothripidae)

The asymmetric “punch and suck” mouthparts of larval Haplothrips verbasci develop from paired appendages in the late, post-anatrepsis embryo similar to those of other insects. Later, the labrum flexes ventrally over the stomodaeum, the right mandibular appendage degenerates, the maxillary appendages divide into inner (lacinial) and outer (stipital) lobes, and the hypopharynx arises from the venters of the mandibular and maxillary segments. All cephalic segments consolidate anteriorly prior to katatrepsis, their appendages flex ventrally, and the labial appendages fuse medially to form the labium and the primordia of the salivary glands and valve. The left mandible and the lacinial lobes of the maxillae invaginate into the head during and after katatrepsis to form the mandibular and maxillary stylet-secreting organs and these later deposit the cuticle of their respective stylets. Cuticle of the mandibular lever is deposited by labral cells at the apex of the mandibular sheath during and after hatching. That of each maxillary lever is secreted simultaneously into the lumen of a ventrally-directed diverticulum developing from stipital cells at the apex of each maxillary sheath. Shortly after katatrepsis, the maxillary and labial palpi originate respectively from cells in the outer wall of each stipital lobe and at the apex of the labium. Muscles of the mouthparts arise after katatrepsis from cephalic mesoderm and are fully-differentiated before cuticle of the mandibular and maxillary levers has been deposited. Gnathal morphogenesis in embryos of H. verbasci resembles that occurring in bug embryos and provides additional evidence that Thysanoptera and Hemiptera evolved from a common psocopteroid stem species having small, paired, biting and chewing mandibles and well developed lacinial stylets.     

Evolution of the planthoppers (Insecta: Hemiptera: Fulgoroidea)

The planthopper superfamily Fulgoroidea (Insecta: Hemiptera) comprises approximately 20 described insect families, depending on which classification is followed. Multiple competing hypotheses of fulgoroid phylogeny have been published, based on either morphological character coding or DNA sequence data; however, those hypotheses disagree in several key aspects regarding the evolution of planthoppers. The current paper seeks to test these hypotheses, including the Asche (Asche, M. 1987. Preliminary thoughts on the phylogeny of Fulgoromorpha (Homoptera Auchenorrhyncha). In: Proceedings of the 6th Auchenorrhyncha Meeting, Turin, Italy, 7-11 September, 1987, pp. 47-53.) hypothesis of a trend in ovipositor structure, which may be correlated with planthopper feeding ecology. Presented here are phylogenetic reconstructions of Fulgoroidea based on analysis of DNA nucleotide sequence data from four loci (18S rDNA, 28S rDNA, Histone 3, and Wingless) sequenced from 83 exemplar taxa representing 18 planthopper families and outgroups. Data sets were analyzed separately and in various combinations under the maximum parsimony criterion, and the total combined dataset was analyzed via both maximum parsimony and partitioned Bayesian criteria; results of the combined analyses were concordant across reconstruction paradigms. Relationships recovered suggest several major planthopper lineages, including: (1) Delphacidae+Cixiidae; (2) Kinnaridae+Meenoplidae; (3) Fulgoridae+Dictyopharidae; (4) Lophopidae+Eurybrachidae (possibly+Flatidae); (5) Ricaniidae+Caliscelidae (possibly+Tropiduchidae). Results also suggest the placement of Achilixiidae outside of Cixiidae and of Tettigometridae as one of the more recently diversified lineages within Fulgoroidea. The resulting phylogeny supports Asche's (1987) hypothesis of a functional trend in ovipositor structure across families.     

Revision of the tribe Colpopterini Gnezdilov, 2003 (Homoptera, Fulgoroidea: Nogodinidae)

The tribe Colpopterini Gnezdilov, 2003 is revised, upgraded to the subfamily Colpopterinae, stat. n., and transferred from the family Issidae Spinola to Nogodinidae Melichar. The tribe Colpopterini is recorded from the Afrotropical Region for the first time—Bumerangum deckerti gen. et sp. n. is described from Southern Africa. The tribe Colpopterini s. str. comprises 6 genera: Bumerangum gen. n., Caudibeccus Gnezdilov et O’Brien, 2008, Colpoptera Burmeister, 1835, Jamaha Gnezdilov et O’Brien, 2008, Neocolpoptera Dozier, 1931, and Ugoa Fennah, 1945. The genera Cheiloceps Uhler, 1895, Tempsa Stål, 1866, Eupilis Walker, 1857, and Gabaloeca Walker, 1870 are transferred to the tribe Issini Spinola of the family Issidae. Issus longulus Lethierry, 1890 is transferred to the genus Colpoptera Burmeister. A key to the genera and a list of the species of the tribe Colpopterini are given. Morphological data confirming independent evolution of similar ovipositor types in the families Issidae and Nogodinidae are provided. The term “styletization” is suggested for describing the transformation of the ovipositor from a rounded to an elongate type.     

Phylogenetic relationships among monogenean gill parasites (Dactylogyridea, Ancyrocephalidae) infesting tilapiine hosts (Cichlidae): systematic and evolutionary implications

We studied the systematics of 14 species of monogenean (Ancyrocephalidae) gill parasites from West African tilapiine hosts (Cichlidae) using both morphological and genetic data. With these tools, we were able to: (i) confirm the validity of the previously described morphological parasite species and of the genus Scutogyrus; (ii) propose that some stenoxenous species (i.e., parasite species with more than one host) may be composed of sister species (e.g., Cichlidogyrus tilapiae); (iii) state that the use of the morphology of the haptoral sclerites is more suitable to infer phylogenetic relationships than the morphology of the genitalia (which seems to be more useful to resolve species-level identifications, presumably because of its faster rate of change). These results imply that: (i) the specificity of these monogenean parasites is greater than initially supposed (what were thought to be stenoxenous species may be assemblages of oïoxenous sister species); (ii) related species groups (i.e., “tilapiae,” “halli,” and “tiberianus”) have to be, as genus Scutogyrus, validated within the 54 ancyrocephalid species described from 18 species of tilapiine hosts in West Africa, (iii) the group “tilapiae,” due to its morphology and host range, have to be considered as being the most primitive; (iv) the occurrence of lateral transfers and parallel speciation processes are necessary to describe the repartition of the newly described parasite groups on the three host genera studied (Tilapia, Oreochromis, and Sarotherodon).     

Lateral line system and its innervation in Tetraodontiformes with outgroup comparisons: Descriptions and phylogenetic implications

The lateral line system and its innervation in ten tetraodontiform families and five outgroup taxa were examined. Although some homology issues remained unresolved, tetraodontiforms were characterized by having two types (at least) of superficial neuromasts (defined by the presence or absence of supporting structures) and accessory lateral lines and neuromasts (except Molidae in which “accessory” elements were absent). The preopercular line in Tetraodontiformes was not homologous with that of typical teleosts, because the line was innervated by the opercular ramule that was newly derived from the mandibular ramus, the condition being identical to that in Lophiidae. Within Tetraodontiformes, the number of neuromasts varied between 70 and 277 in the main lines and between 0 and 52 in accessory elements. Variations were also recognized in the presence or absence of the supraorbital commissure, mandibular line, otic line, postotic line, ventral trunk line, and some lateral line nerve rami, most notably the dorsal branch of the opercular ramule, being absent in Aracanidae, Ostraciidae, Tetraodontidae, Diodontidae, and Molidae. Morphological characteristics derived from the lateral line system and its innervation provided some support for a sister relationship of tetraodontiforms with lophiiforms. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Systematic significance of pollen arrangement in MICROSPORANGIA OF POACEAE AND Cyperaceaei REVIEW AND OBSERVATIONS ON REPRESENTATIVE TAX A

An anatomical survey of anthers of the Poaceae and Cyperaceae and two related families, the Restionaceae and Flagellariaceae, was conducted to determine the taxonomic distribution and possible phylogenetic significance of pollen arrangement. An unusual pollen arrangement was known from a small number of taxa in the Poaceae and Cyperaceae, in which a single, uniseriate cylinder of pollen grains is arranged in the anther locule such that each grain is in contact with the tapetum (termed here “peripheral” pollen). This contrasts with the prevailing arrangement in other angiosperms in which the locule contains a relatively large number of pollen grains in no special configuration, with many interior grains that never touch the tapetum (termed here “central” pollen). A total of 48 species in these four families was examined in this study, and observations on pollen arrangement for numerous additional species in these families were gleaned from the literature. We confirm that the peripheral arrangement is predominant in Poaceae and Cyperaceae, although both families also include species with central pollen, whereas only central pollen is found in Restionaceae and Flagellariaceae. In the peripheral arrangement, the pore of the pollen grain or pseudomonad, when observed, is in contact with the tapetum, but it has not been definitively demonstrated that this is always the case. Peripheral pollen in the Poaceae and Cyperaceae is nonhomologous because of the presence of pseudomonads in the latter family. It remains unexplained why peripherally arranged pollen or pseudomonads, with the attendant reduction in the number of pollen grains, should be associated with anemophily in these two families.     

A generalized framework to expand incomplete phylogenies using non-molecular phylogenetic information

Aim

The increasing availability of molecular information has lifted our understanding of species evolutionary relationships to unprecedent levels. However, current estimates of the world's biodiversity suggest that about a fifth of all extant species are yet to be described, and we still lack molecular information for many of the known species. Hence, evolutionary biologists will have to tackle phylogenetic uncertainty for a long time to come. This prospect has urged the development of software to expand phylogenies based on non-molecular phylogenetic information, and while the available tools provide some valuable features, major drawbacks persist and some of the proposed solutions are hardly generalizable to any group of organisms.

Innovation

Here, we present a completely generalized and flexible framework to expand incomplete phylogenies. The framework is implemented in the R package “randtip”, a toolkit of functions that was designed to randomly bind phylogenetically uncertain taxa in backbone phylogenies through a fully customizable and automatic procedure that uses taxonomic ranks as a major source of phylogenetic information. Although randtip can generate fully operative phylogenies for any group of organisms using just a list of species and a backbone tree, we stress that the “blind” expansion of phylogenies using “quick-and-dirty” approaches often leads to suboptimal solutions. Thus, we discuss a variety of circumstances that may require customizing simulation parameters beyond default settings to optimally expand the trees, including a detailed step-by-step tutorial that was designed to provide guidelines to non-specialist users.

Main Conclusions

Phylogenetic uncertainty should be tackled with caution, assessing potential pitfalls and opportunities to optimize parameter space prior to launch any simulation. Used judiciously, our framework will help evolutionary biologists to efficiently expand incomplete phylogenies and thereby account for phylogenetic uncertainty in quantitative analyses.     

Cephalic morphology of Hymenopus coronatus (Insecta: Mantodea) and its phylogenetic implications

External and internal head structures of the mantodean Hymenopus coronatus are examined and described in detail. The results are elaborately compared with the literature. Strong crests on the anterior tentorial arms that articulate with the subantennal suture, a parietal suture and glossae and paraglossae with anteriorly bent tips are proposed as new potential apomorphies for Mantodea while a head capsule being wider than long, enlarged compound eyes, the presence of a frontal shield or scutellum, lateral lobes in the anterior tentorial arms, the presence of a transverse and an interantennal suture and the reduction of the mentum are confirmed as apomorphies, As potential apomorphies for Dictyoptera the reduction of Musculus tentoriobuccalis lateralis (M. 49) is newly presented and a “perforate” tentorium, lacinial incisivi that are located in a galeal pouch and the presence of a postmola are confirmed. The present study shows the value of cephalic morphology for phylogenetic analysis but also points out that further studies including evolutionary key taxa are essential for resolving the evolutionary adaptations among dictyopterans.     

Global pattern of plant utilization across different organisms: Does plant apparency or plant phylogeny matter?

The present study is the first to consider human and nonhuman consumers together to reveal several general patterns of plant utilization. We provide evidence that at a global scale, plant apparency and phylogenetic isolation can be important predictors of plant utilization and consumer diversity. Using the number of species or genera or the distribution area of each plant family as the island “area” and the minimum phylogenetic distance to common plant families as the island “distance”, we fitted presence–area relationships and presence–distance relationships with a binomial GLM (generalized linear model) with a logit link. The presence–absence of consumers among each plant family strongly depended on plant apparency (family size and distribution area); the diversity of consumers increased with plant apparency but decreased with phylogenetic isolation. When consumers extended their host breadth, unapparent plants became more likely to be used. Common uses occurred more often on common plants and their relatives, showing higher host phylogenetic clustering than uncommon uses. On the contrary, highly specialized uses might be related to the rarity of plant chemicals and were therefore very species‐specific. In summary, our results provide a global illustration of plant–consumer combinations and reveal several general patterns of plant utilization across humans, insects and microbes. First, plant apparency and plant phylogenetic isolation generally govern plant utilization value, with uncommon and isolated plants suffering fewer parasites. Second, extension of the breadth of utilized hosts helps explain the presence of consumers on unapparent plants. Finally, the phylogenetic clustering structure of host plants is different between common uses and uncommon uses. The strength of such consistent plant utilization patterns across a diverse set of usage types suggests that the persistence and accumulation of consumer diversity and use value for plant species are determined by similar ecological and evolutionary processes.     

Elevational filtering and the evolution of planthoppers (Hemiptera,Fulgoromorpha) in Papua New Guinea

Along elevational gradients, phylogenetic relatedness patterns constitute a considerable source of information and may shed light on ecological processes that structure communities. This study focuses on community phylogenetic structure of planthoppers, specifically the species-rich and abundant Fulgoromorpha families (Hemiptera, Auchenorrhyncha), Cixiidae and Derbidae + Achilidae, along an elevational gradient on Mount Wilhelm (Papua New Guinea). In order to assess the factors driving planthoppers community composition, we recorded abundance data for planthoppers species at each elevation and we generated a molecular phylogeny of the local species, using Bayesian inference. We analyzed 168 individuals representing 59 local morphospecies. Using a fully resolved and well-supported phylogeny, we then investigated the phylogenetic structure of the communities by performing a Spatial Analysis of Community Diversity. We show that Cixiidae are phylogenetically clustered along the elevational gradient, whereas Derbidae + Achilidae harbor a random structure, suggesting that local adaptation to elevation shapes community structure of Cixiidae, but not that of Derbidae + Achilidae. Our findings highlight the importance of phylogenies in the study of tropical elevational gradients.     

Fish germinal cells. I. Comparative spermatology of seven Cyprinid species

The spermatozoa of seven fishes belonging to Cyprinid family are examined. They have no acrosome, like all Teleost fishes, a spheroidal or slightly elliptic nucleus, always eccentrically placed on the tail, two variously oriented centrioles, and a postnuclear cytoplasmic region of various size that contains some mitochondria (2 to 10) and surrounds a periaxonemal postnuclear canal. The tail is of moderate length (from 36 to 60 μm) and contains a “9 + 2” axoneme: both dynein arms are present. Comparative examination of the spermatozoa in the seven species shows that significant differences occur among them, even when they belong to the same genus. These concern the tail length; the position of the centrioles, the proximal with respect to the central one and with respect to the nucleus; the number of mitochondria, which is in relationship to the depth of the postnuclear canal. In the uniform general pattern of the ultrastructure of the Cyprinid spermatozoa, each species is characterized by a particular organization of the sperm organelles; in this respect, the two species examined by us, Leuciscus cephalus and souffia, are more closely related, even if easily recognizable one from the other. From a phylogenetic point of view, the comparative spermatology of the Cyprinid fishes suggests that the mitochondrial number is a good character, which enables us to order them in a phylogenetic arrangement.