Tribal Level Research Articles

Review of the Oriental Leafhopper GenusLampridius(Hemiptera: Cicadellidae: Deltocephalinae), with Description of a Related New Genus

The leafhopper genus Lampridius Distant, 1918 (type species: L. spectabilis Distant, 1918 ) is redescribed, and a second species, L. cuspidatus sp. nov. (Thailand: Loei), is described. A related new genus Paralampridius gen. nov. and four new species are described: P. mimicus sp. nov. (China: Guangdong, Hainan), P. rotundatus sp. nov. (Thailand: Loei), P. sinuatus sp. nov. (Thailand: Loei, Suphanburi, Petchaburi), and P. truncatus sp. nov. (Thailand: Chaiyaphum, Ubon Ratchathani). Both genera are tentatively included in the tribe Opsiini, although only Lampridius has paired aedeagal shafts with separate gonopores. All six species are illustrated, and a key is provided for their identification. Problems with the current tribal classification are discussed in light of the present discovery of closely related species that show variation in characters previously assumed to be stable at the tribal level in the subfamily Deltocephalinae.

An updated phylogeny and classification of Lamiaceae subfamily Lamioideae

AbstractLamioideae comprise the second‐largest subfamily in Lamiaceae. Although considerable progress has recently been made in Lamioideae phylogenetics, the subfamily remains one of the most poorly investigated subfamilies in Lamiaceae. Here we present a taxonomic update of the subfamily based on earlier published data as well as 71 new DNA extracts from relevant in‐ and outgroup taxa, and DNA sequence data from four chloroplast regions (matK, rps16, trnL intron and trnL‐F spacer). The phylogenetic positions of 10 out of 13 previously unplaced small or monotypic Asian lamioid genera and 37 additional lamioid species have been identified, and the classification is updated accordingly. Results from parsimony and Bayesian phylogenetic methods corroborate earlier results, but phylogenetic resolution as well as overall branch support are improved. All newly added genera are assigned to earlier established tribes or the new tribe Paraphlomideae Bendiksby, which includes Ajugoides, Matsumurella and Paraphlomis. Acanthoprasium is resurrected as a genus. Transfer of species is proposed to ac ‐ commodate the monophyly of two genera (Lamium, Otostegia), whereas ten genera remain non‐monophyletic (Ballota s. str., Lagopsis, Leonotis, Leonurus, Leucas, Microtoena, Phlomoides, Sideritis, Stachys, Thuspeinanta). Eriophyton and Stachyopsis have been included in Lamieae, Hypogomphia in Stachydeae, and Loxocalyx in Leonureae. Betonica, Colquhounia, Galeopsis, and Roylea remain unclassified at the tribal level. Lamium chinense and three East Asian Galeobdolon species are transferred to Matsumurella. Sulaimania and four Otostegia species are transferred to Moluccella . Alajja and three Lamium species are transferred to Eriophyton. In total, 14 new combinations are made, one at the rank of subgenus and 13 at the rank of species.

On the evolution of raptorial legs - an insect example (Hemiptera: Reduviidae: Phymatinae).

The presence of chelate and subchelate fore legs in Phymatinae (Hemiptera: Reduviidae), or ambush bugs, provides a unique opportunity to study the evolution of different types of raptorial legs in a closely related group of arthropods. Themonocorini have simple, possibly raptorial legs, Phymatini and Macrocephalini distinct subchelate fore legs, and the charismatic Carcinocorini are the only insects with a chelate fore leg apart from female dryinid Chysidoidea (Hymenoptera). Relationships between the four phymatine tribes are here analyzed in a cladistic framework thus permitting testable hypotheses on the evolution of raptorial legs. The presented analysis of phymatine tribal level relationships is based on a dataset comprising 11 species of Phymatinae and 54 non-phymatine Reduviidae and Heteroptera. The molecular data set consists of ∼3500 MAFFT aligned bases of 16S, 28S D2-D3, and 18S ribosomal genes. Parsimony and maximum likelihood analyses resulted in identical topologies for the ingroup with the relationships Themonocorini + (Phymatini + (Carcinocorini + Macrocephalini)) receiving high support values. Eleven morphological characters, eight of them derived from fore leg morphology, were optimized on the parsimony analysis. These optimizations indicate that the ancestral ambush bug had a simple raptorial leg; that size reduction of the tarsus, enlargement of the femur, curvature of the fore tibia, armature of tibia and femur with rows of tiny tubercles that allow for gripping of a prey insect, and the large process on the ventral surface of the femur arose in the common ancestor of Carcinocorini + Macrocephalini + Phymatini. The chelate leg in Carcinocorini is likely derived from a subchelate precursor similar to the one seen in recent Macrocephalini and may have evolved through elongation of the ventral, proximal portion of the fore femur and modification of the median process to form part of the digitus fixus. © The Willi Hennig Society 2010.

The occurrence and phylogenetic implications of wing interference patterns in Cynipoidea (Insecta:Hymenoptera)

Wing interference patterns (WIPs) are a potentially rich source of taxonomic data. We surveyed a broad range of Cynipoidea (Hymenoptera) for the presence of these patterns. Further, we used phylogeny to investigate the evolutionary patterns of WIPs throughout the superfamily. We also documented that WIPs occur across nearly all species sampled, in both the hindwing and forewing; in many instances, the forewing WIP is carried over to the hindwing. Only Austrocynips mirabilis Riek lacked WIPs. Among species that possess WIPs, we were able to recognise four distinct categories: radiform (radiating), striatiform (longitudinally striate), campiform (continuous field) and galactiform (amorphous mixture of colours). Wing interference patterns are not found in infuscate wings or areas of infuscation on otherwise hyaline wings, and are reduced on large wings. Within Figitidae, WIPs are more phylogenetically conservative, especially at the tribal level: within Cynipidae the phylogenetic conservatism of WIPs is somewhat less clear, though all Synergini sampled had a very consistent WIPs. For certain genera, such as Ganaspidium and Andricus, WIPs may prove useful for species-level discrimination.

Phylogeny and Recircumscription of Artocarpeae (Moraceae) with a Focus on <I>Artocarpus</I>

Abstract Moraceae is a large (∼1,050 species) primarily tropical family with several economically and ecologically important species. While its monophyly has been well supported in recent studies, relationships within the family at the tribal level and below remain unresolved. Delimitation of the tribe Artocarpeae has been particularly difficult. Classifications based on morphology differ from those based on phylogenetic studies, and all treatments include highly heterogeneous assemblages of genera that seem to represent a cross section of the family. We evaluated chloroplast and nuclear DNA sequence data for 60 Moraceae taxa representing all genera that have been included in past treatments of Artocarpeae and also included species from several other Moraceae tribes and closely related families as outgroups. The data were analyzed using maximum parsimony and maximum likelihood methods and indicate that none of the past treatments of Artocarpeae represent a monophyletic lineage. We present the most complet...

An enigmatic new leafhopper genus, Protochiasmus (Hemiptera, Cicadellidae, Deltocephalinae), from Brazil

A new brachypterous leafhopper genus and species of Deltocephalinae, Protochiasmus mysticus gen. n., sp. n. from Brazil, are described. The genus is not easily placed to tribe partly due to the lack of known females which may provide characters that are diagnostic to the tribal level and also due to the unique combination of morphological characters in the male. Based on these morphological characters and on recent molecular and morphological phylogenetic analyses of Deltocephalinae, Protochiasmus is tentatively placed in Chiasmini. (© 2010 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)

Scanning electron microscopy of fruits in the West African Polygonaceae

The fruit morphology of 18 taxa representing seven genera of the family Polygonaceae in West Africa was investigated using scanning electron microscopy. The achenes are trigonous, lenticular, globose, subglobose, heart shaped, ovoid, or cone like. Sizes range from 0.12 x 0.10 cm(2) in Polygonum plebeium to 7.87 x 0.58 cm(2) in Afrobrunnichia erecta. Colors are brown to black. The cells are isodiametric in P plebeium, irregular in A. erecta, Antigonon leptopus, and Harpagocarpus snowdenii, and polygonal in other species. The walls are straight, curved, or undulate and are either raised or depressed. Afrobrunnichia erecta is characterized by deeply sinuate lateral walls. The cell surface may be smooth or tuberculate or fibrillate in the family, usually covered with wax deposits. The combination of these characters is mainly taxonomically useful at the tribal level and rarely at the specific or infraspecific level for the delimitation of the taxa.

Towards a more robust molecular phylogeny of Chinese Apiaceae subfamily Apioideae: Additional evidence from nrDNA ITS and cpDNA intron ( rpl16 and rps16) sequences

China contains some of the world’s greatest diversity of Apiaceae (Umbelliferae), with ten endemic genera. Our previous investigation into the phylogenetic relationships of Chinese Apiaceae subfamily Apioideae, based exclusively on nrDNA ITS sequences, revealed several major clades heretofore unrecognized in the subfamily and confirmed the phylogenetic placements of five endemic genera. To further elucidate relationships among Chinese Apioideae, ascertain the phylogenetic placements of the remaining endemic genera, and test hypotheses of relationships proposed in our earlier study, additional taxa were examined for ITS and, for smaller sets of accessions, cpDNA rpl16 and rps16 intron sequence variation. These ITS and cpDNA data matrices comprised 158 and 131 accessions, respectively, with 110 accessions included in the analysis of combined ITS and cpDNA data. Maximum parsimony and Bayesian analyses of partitioned ITS and chloroplast data sets resulted in highly consistent phylogenies, whereas analyses of combined molecular data resulted in trees of greatest resolution and overall branch support. Two major clades identified in our previous study are recognized at the tribal level: Komarovieae J. Zhou & S. R. Downie trib. nov. and Chamaesieae J. Zhou & F. D. Pu trib. nov. The monotypic tribe Chamaesieae represents one of the earliest diverging lineages of subfamily Apioideae in Asia. The Acronema and East Asia clades of previous circumscription are each expanded. Of the five Chinese endemic genera not examined previously, Chaerophyllopsis finds affinity within tribe Scandiceae subtribe Scandicinae, Chuanminshen allies with Changium and Cyclorhiza in tribe Komarovieae, Harrysmithia falls within the Acronema Clade, Melanosciadium embeds in Angelica in tribe Selineae, and Dickinsia is confirmed as a member of Apiaceae subfamily Azorelloideae. The cpDNA-based phylogenies are not sufficiently resolved in their distal portions to elucidate the tribal placement of the endemic genus Nothosmyrnium, whereas the ITS trees strongly indicate an affinity with tribe Pimpinelleae. The affinities of several Chinese endemic species are also addressed. Peucedanum delavayi, for example, is phylogenetically distant from Sinodielsia and Meeboldia, genera with which it has been allied previously, and shows close affinity to three Chinese species of Ligusticum in tribe Selineae. Aside from providing a framework for taxonomic revisions, the phylogenetic structure recovered in this study for subfamily Apioideae will lay the foundation for future investigations of evolutionary patterns of morphological characters and biogeography.

Nomenclature of tribes within the Urticaceae

As part of systematic studies of the genus Elatostema (Urticaceae) and its sister genera, it was apparent that different names had been applied at the tribal level for these genera. This paper proposes that the tribal name Elatostemeae be used for the genera currently delineated as belonging to tribe Lecantheae (Friis 1993). It is also proposed that the authorship of the currently recognised tribes in the Urticaceae should be attributed to Charles Gaudichaud-Beaupres as described in H. L. C. de Freycinet’s Voyage autour du monde…execute sur les corvettes de S. M. l’Uranie et la Physiciene, published in 1830.

A new genus and species of the tribe Trigonogeniini Cobos, 1956, from Belize (Coleoptera: Buprestidae)

The name Trigonogenium Harold, 1869 was introduced as a replacement name for Trigonophorus Solier, 1849 (preoccupied by Serville,1834: Hymenoptera). The genus was revised by Cobos (1956) who discussed the systematic position of Trigonogenium and suggested for the genus a separate tribe Trigonogenini (sic !). The next revision was published also by Cobos (1986) and the placement of the genus was repeatedly discussed by Hołyński (1988, 1993), Bílý (2000), Volkovitsh (2001), Bellamy (2003, 2007) and the tribal level of the group was finaly confirmed by Bílý & Volkovitsh (2007) after a detailed study of the larval morphology. Up to now the tribe has contained only two genera: Trigonogenium (three species and one subspecies) and Hovorigenium Bellamy, 2007 (one species).

Digestive system of larval black flies (Diptera: Simuliidae): structure and phylogeny

Abstract. Larval gut characters of 45 species of black flies, representing 17 subgenera and 15 genera, were studied for their utility in taxon diagnosis and phylogenetic inference. Phase‐contrast and scanning electron microscopy revealed a wealth of new characters in the esophageal armature of the foregut. Larval gut morphology, particularly characters of the esophageal armature, varied significantly among taxa, permitting generic diagnoses. A cladogram inferred from 13 gut characters supported a sister‐group relationship between Parasimulium and the remaining simuliids, but did not show a strong phylogenetic signal for the current, well‐supported concept of tribal level monophyly.

Systematic position and cladistic analysis of Gyrasida Koch, a remarkable genus of Praocini (Coleoptera: Tenebrionidae) from Chile

Gyrasida Koch (Pimeliinae: Praocini), distributed in central Chile is elevated to generic status and transferred from Asidini to Praocini on the basis of constant tribal level characters. Prior to this study Gyrasida was considered a subgenus of the South African Afrasida Wilke. Character states of Gyrasida shared with the other genera of Praocini and not shared with genera of Asidini are discussed. A cladistic analysis of the six species of the genus is presented including: G. propensa (Wilke) (type species), G. lucianoi Flores & Vidal, G. fernandoi Flores & Vidal, G. tomasi Vidal, G. franciscae Vidal, and G. camilae Vidal. This article includes a redescription of the genus, habitus photographs, illustrations of external morphology and genitalic features, a cladogram, and a distribution map.

Phylogenetic relationships within the Cimicomorpha (Hemiptera: Heteroptera): a total‐evidence analysis

AbstractA phylogenetic analysis for the Cimicomorpha was conducted using 92 taxa, including eight outgroups and six species of Thaumastocoridae. Density of taxon sampling allows for tests of relationships at the family level for most taxa, whereas in the Miridae denser sampling allows for doing so on the tribal level. This level of sampling also corresponds with the availability of testable published hypotheses of relationships. Morphological data for 73 characters are coded for all taxa. Approximately 3500 base pairs of DNA were sequenced for the following gene regions for 83 taxa: 16S rDNA, 18S rDNA, 28S rDNA and COI. Results are presented for analysis of morphological data, individual molecular partitions, combined molecular data, combined morphological and molecular data for 83 taxa and combined morphological and molecular data for 92 taxa. Analyses of morphological data were performed using the parsimony programsnonaandpiwe: molecular and combined data were analysed using direct optimization with the programpoy. Major conclusions of the present study include recognition of the following monophyletic groups: The Geocorisae is a monophyletic group. The monophyly of the Cimicomorpha – including Thaumastocoridae – is not supported in most analyses. The Reduviidae is monophyletic, with the Phymatinae Complex being the sister‐group of the remaining subfamilies. The circumscription of the Cimiciformes is altered from the prior conception of Schuh and Štys to also include the Joppeicidae, Microphysidae and Velocipedidae, as well as the recently described family Curaliidae; the monophyly of the Cimiciformes is supported in most analyses; the Cimiciformes is treated as the sister‐group of the Miroidea in most analyses. The monophyly of the Cimicoidea, including Curaliidae, is supported in all analyses including molecular data, whereas Curaliidae is treated as a more basal cimiciform in all other analyses. The monophyly and placement of the Thaumastocoridae is ambiguous across the range of analyses, and the monophyly of the Miroidea sensu Schuh and Štys receives limited support in the combined analyses of morphology + molecular data. The Tingidae and Miridae are each monophyletic and together almost invariably form a monophyletic group. Within the Miridae, several inclusive monophyletic groups at the subfamily/tribal level are more or less consistently recognized when molecular data are included; however, the interrelationships of the subfamilies vary substantially across the range of analyses. Of the individual molecular partitions, only 18S rDNA shows significant congruence with combined analyses of morphological, combined molecular or combined morphological and molecular data. Scenarios are discussed for the evolution of the metathoracic scent‐efferent system and the origin of the fossula spongiosa.

The genus Xenaclopus Arrow (Coleoptera: Scarabaeidae): redescription and removal from the Aclopinae, with systematic notes

The monotypic genus Xenaclopus Arrow is redescribed and illustrated, based in its lectotype and paralectotypes. Characters examined indicate that this genus should be removed from the Aclopinae and placed into the subfamily Melolonthinae as insertae sedis at the tribal level.

Wood anatomy of Rauvolfioideae (Apocynaceae): a search for meaningful non‐DNA characters at the tribal level

Wood anatomical studies in the economically important Apocynaceae or dogbane family are fragmentary. This study represents a first attempt to unravel the phylogenetic significance and major evolutionary trends in the wood of the family, using existing and new microscopic wood observations within the large subfamily Rauvolfioideae. On the basis of LM and SEM observations of 91 species representing all 10 currently recognized tribes, we found that most of the tribes are characterized by a unique combination of wood characters, such as vessel grouping, vessel element length, fiber type, frequency of uniseriate rays, and fused multiseriate rays. Climbing rauvolfioid taxa can generally be distinguished from erect species by their wider vessels, tendency to form paratracheal axial parenchyma, presence of tracheids, and occurrence of laticifers in rays. With respect to the entire family, there is a general phylogenetic trend toward shorter vessel elements, a higher proportion of vessels in multiples and more vessels per multiple, higher tracheid abundance, more paratracheal parenchyma, and fewer cells per axial parenchyma strand in the more derived Apocynaceae. Most of these evolutionary trends are likely to be triggered by drier environmental conditions and/or shifts from an erect to a climbing habit.

Palynological evolutionary trends within the tribe Mentheae with special emphasis on subtribe Menthinae (Nepetoideae: Lamiaceae)

The pollen morphology of subtribe Menthinae sensu Harley et al. [In: The families and genera of vascular plants VII. Flowering plants·dicotyledons: Lamiales (except Acanthaceae including Avicenniaceae). Springer, Berlin, pp 167–275, 2004] and two genera of uncertain subtribal affinities (Heterolamium and Melissa) are documented in order to complete our palynological overview of the tribe Mentheae. Menthinae pollen is small to medium in size (13–43 μm), oblate to prolate in shape and mostly hexacolpate (sometimes pentacolpate). Perforate, microreticulate or bireticulate exine ornamentation types were observed. The exine ornamentation of Menthinae is systematically highly informative particularly at generic level. The exine stratification in all taxa studied is characterized by unbranched columellae. Orbicules are consistently absent in Menthinae. Our palynological data are interpreted in a phylogenetic context at tribal level in order to assess the systematic value of pollen characters and to evaluate the existing molecular phylogenies for this group. Pollen morphology suggests Heterolamium as a close relative of subtribe Nepetinae and supports the molecular affinity of Melissa to subtribe Salviinae.

Phylogeny and biogeography of African Murinae based on mitochondrial and nuclear gene sequences, with a new tribal classification of the subfamily

BackgroundWithin the subfamily Murinae, African murines represent 25% of species biodiversity, making this group ideal for detailed studies of the patterns and timing of diversification of the African endemic fauna and its relationships with Asia. Here we report the results of phylogenetic analyses of the endemic African murines through a broad sampling of murine diversity from all their distribution area, based on the mitochondrial cytochrome b gene and the two nuclear gene fragments (IRBP exon 1 and GHR).ResultsA combined analysis of one mitochondrial and two nuclear gene sequences consistently identified and robustly supported ten primary lineages within Murinae. We propose to formalize a new tribal arrangement within the Murinae that reflects this phylogeny. The diverse African murine assemblage includes members of five of the ten tribes and clearly derives from multiple faunal exchanges between Africa and Eurasia. Molecular dating analyses using a relaxed Bayesian molecular clock put the first colonization of Africa around 11 Mya, which is consistent with the fossil record. The main period of African murine diversification occurred later following disruption of the migration route between Africa and Asia about 7–9 Mya. A second period of interchange, dating to around 5–6.5 Mya, saw the arrival in Africa of Mus (leading to the speciose endemic Nannomys), and explains the appearance of several distinctive African lineages in the late Miocene and Pliocene fossil record of Eurasia.ConclusionOur molecular survey of Murinae, which includes the most complete sampling so far of African taxa, indicates that there were at least four separate radiations within the African region, as well as several phases of dispersal between Asia and Africa during the last 12 My. We also reconstruct the phylogenetic structure of the Murinae, and propose a new classification at tribal level for this traditionally problematic group.

Molecular phylogeny of Pemphiginae (Hemiptera: Aphididae) inferred from nuclear gene EF-1α sequences

Three traditional tribes of Fordini, Pemphigini and Eriosomatini comprise Pemphiginae, and there are two subtribes in Fordini and Pemphigini, respectively. Most of the species in this subfamily live heteroecious holocyclic lives with distinct primary host specificity. The three tribes of Pemphigini (except Prociphilina), Eriosomatini and Fordini use three families of plants, Salicaceae (Populus), Ulmaceae (Ulums) and Anacardiaceae (Pistacia and Rhus), as primary hosts, respectively, and form galls on them. Therefore, the Pemphigids are well known as gall makers, and their galls can be divided into true galls and pseudo-galls in type. We performed the first molecular phylogenetic study of Pemphiginae based on molecular data (EF-1alpha sequences). Results show that Pemphiginae is probably not a monophylum, but the monophyly of Fordini is supported robustly. The monophyly of Pemphigini is not supported, and two subtribes in it, Pemphigina and Prociphilina, are suggested to be raised to tribal level, equal with Fordini and Eriosomatini. The molecular phylogenetic analysis does not show definite relationships among the four tribes of Pemphiginae, as in the previous phylogenetic study based on morphology. It seems that the four tribes radiated at nearly the same time and then evolved independently. Based on this, we can speculate that galls originated independently four times in the four tribes, and there is no evidence to support that true galls are preceded by pseudo-galls, as in the case of thrips and willow sawflies.

Phylogeny and evolutionary history of the blister beetles (Coleoptera, Meloidae)

Meloid beetles are well characterised by both morphological and biological features. Previous phylogenetic hypotheses based on morphological characters assumed the repeated parallel evolution of complex biological novelties. In this work relationships among several taxa of the four subfamilies and almost all tribes representing meloid diversity are examined by using mitochondrial (16S) and nuclear (ITS2) DNA sequences, in 25 genera (using Anthicidae as outgroup). Secondary structure of 16S and ITS2 rRNAs were modelled. ITS2 structure represents a synapomorphic condition for the family and informative characters at the tribal level. Phylogenetic hypotheses based on separate and combined analysis of the 16S and ITS2 rDNA sequences, and morpho-biological characters were tested, and compared with previous morphological classifications. Molecular dating allowed an outline of the main steps of the evolutionary history of Meloidae, which evolved during Early Cretaceous and then radiated considerably with the adoption of hypermetaboly and parasitic behaviour, and with repeated, parallel evolution of larval phoresy on its hosts.

A new record and an evaluation of the phylogenetic relationships of Abrothrix olivaceus markhami (Rodentia: Sigmodontinae)

Recent phylogenetic studies of the subfamily of cricetid rodents (Sigmodntinae) have validated the taxonomic classification at the tribal level of the Andean Clade. It is possible that some endemic species from Patagonian South America are part of this new tribe, but previous studies have not evaluated this hypothesis due to the difficulty of obtaining samples. In this study, we evaluate the phylogenetic relationships of Akodon markhami (Pine, 1973), a species endemic to the Chilean Patagonia, using individuals recently captured at the type locality of this taxon. Our results indicate that this form of Akodon corresponds to a subspecies of Abrothrix olivaceus, and should be incorporated into the Andean Clade as a geographic race of this widely distributed species on the South American continent. Based on a molecular clock calibration, we dated the origin of this geographic race during the last glacial cycles of the Quaternary, as the result of a vicariant process.