Assessment of monophyly of species-groups within Afrotropical Empidini (Diptera: Empididae: Empidinae), with a cladistic analysis of the Empis setitarsus-group

Abstract.  The Afrotropical Empidini are reviewed from a phylogenetic point of view. Seven monophyletic groups are recognized within the tribe, namely Empis ( Coptophlebia ) bivittata , E. ( C ) chrysocera , E. ( C ) hyalea , E. ( C ) namwamba , E. ( C ) poecilosoma , E. ( Disneyempis ) and E. setitarsus . Their monophyly is assessed by comparison with the remaining Empidini. The E. setitarsus -group is cladistically analysed to study the evolution of the length of vein M1 by optimization on the cladogram, and the status of subgenus Coptophlebia Bezzi is discussed. A key to the seven Afrotropical species-groups of Empidini is provided, genus Rhamphomyia is no longer recognized for the Afrotropical Region, and four new species belonging to the E. setitarsus -group, namely E. cinerarius sp.n., E. pseudosetitarsus sp.n., E. spinosa sp.n and E. spungaberaensis sp.n., are described and included in a key to all known species of this group.     

Review of the genus Stephanospathius Belokobylskij, 1992 (Hymenoptera, Braconidae), with discussion of their tribal position

A review of the Afrotropical genus Stephanospathius Belokobylskij is provided. A new species Stephanospathius benoitisp. n. from the Republic of Congo, and the male and, for the first time, the female of Stephanospathius ornatipes (Kieffer) are described. A discussion of the status and composition of the tribe Stephaniscini is given and a new name for this tribe, Leptospathiini, nom. n., is proposed. A key to the included genera and a key to species of Stephanospathius are provided.     

A revision of the tribe Plectrothripini of fungus-feeding Thysanoptera (Phlaeothripidae: Phlaeothripinae)

Abstract The tribe Plectrothripini is redefined with examination of its systematic relationships. Keys are provided to the ten genera and forty-seven species in this tribe; one genus and eighteen species are newly described; one genus and two species are newly synonymized; two genera, Chiridothrips and Eurytrichothrips , are here removed from this tribe. The species of the tribe occur throughout the tropical and subtropical regions of the world; seventeen species are recorded from the Oriental Region, seventeen species from the Neotropical Region, ten species from the Afrotropical Region, two from Australia and one from the Pacific. They seem to be fungus-feeders, being found mainly under the bark of trees or on dead branches.     

Conochironomus Freeman: an Afro-Australian Chironomini genus revised (Diptera: Chironomidae)

The chironomid genus Conochironomus is revised, with diagnoses provided for both sexes of adults and, for the first time, the immature stages. The Afrotropical genotype C.acutistilus Freeman and the newly described Australian species C.australiensis (for misidentified acutistilus in Australia), C.cygnus and C.kakadu are described in all stages. C.avicula Freeman is redescribed from the male, and C.deemingi is described as new from the male alone, both from the Afrotropical Region. The Australian species C.cervus is described as new, based on the pupa alone. Comments are made on the ecology and distribution. A previously little-recognized 'tropical' Gondwanan biogeographic explanation for Conochironomus distribution is postulated.     

Molecular phylogenetics of Acacia (Fabaceae: Mimosoideae) based on the chloroplast MATK coding sequence and flanking TRNK intron spacer regions

The tribe Acacieae (Fabaceae: Mimosoideae) contains two genera, the monotypic African Faidherbia and the pantropical Acacia, which comprise about 1200 species with over 950 confined to Australia. As currently recognized, the genus Acacia is subdivided into three subgenera: subg. Acacia, subg. Aculeiferum, and the predominantly Australian subg. Phyllodineae. Morphological studies have suggested the tribe Acacieae and genus Acacia are artificial and have a close affinity to the tribe Ingeae. Based on available data there is no consensus on whether Acacia should be subdivided. Sequence analysis of the chloroplast trnK intron, including the matK coding region and flanking noncoding regions, indicate that neither the tribe Acacieae nor the genus Acacia are monophyletic. Two subgenera are monophyletic; section Filicinae of subgenus Aculeiferum does not group with taxa of the subgenus. Section Filicinae, eight Ingeae genera, and Faidherbia form a weakly supported paraphyletic grade with respect to subg. Phyllodineae. Acacia subg. Aculeiferum (s. s.) is sister to the grade. These data suggest that characters currently used to differentiate taxa at the tribal, generic, and subgeneric levels are polymorphic and homoplasious in cladistic analyses.     

科昆虫地理分布研究初探

本文就科昆虫的起源,各亚科、族及属的分布特点进行了分析和总结;并根据各属的世界分布情况,将其分为８种类型。文中还就中国科昆虫的地理分布及特点作了初步的探讨,结果表明：在我国已知的２亚科４族２０个属中,东洋区分布的有１０个属,其中有３个属仅分布于我国;古北区分布的有１个属;东洋－古北区分布的有３个属;东洋－新北区分布的有１个属;东洋－古北－新北区分布的有４个属;东洋－古北－新北－非洲区分布的有１个属。此外,科昆虫在中国的一个主要分布特点就是多数种类集中分布于华中区、华南区和西南区;而华中区则很可能是科种类的分化中心,并以此为中心向其他区扩散。     

Bobgunnia, a new African genus of tribe Swartzieae (Fabaceae, Faboideae)

A new genus, Bobgunnia, is established in tribe Swartzieae for the African species of Swartzia. Two new combinations are made: B. fistuloides and B. madagascariensis. The new genus has seeds, unlike the remainder of tribe Swartzieae but like the other tribes of Faboideae. The systematic position of Swartzieae is reevaluated.     

Revised systematics and biogeography of ‘Quediina’ of sub‐Saharan Africa: new phylogenetic insights into the rove beetle tribe Staphylinini (Coleoptera: Staphylinidae)

Abstract Quediina, a mega‐diverse conventional subtribe of the rove beetle tribe Staphylinini, is remarkably species rich in the north and south temperate regions of the world. Tropical faunas of this group, and the fauna of the entire Afrotropical biogeographical region (= Ethiopian region, = sub‐Saharan Africa), in contrast, are remarkably poor. The taxonomic study of the quediine genera of Staphylinini from the Afrotropical region reveals misidentifications for many of them. Their phylogenetic study demonstrates polyphyly of Quediina and reveals a new evolutionary pattern for the entire tribe Staphylinini. In particular, the formerly quediine genera Euristus Fauvel, 1899 , Ioma Blackwelder, 1952, Natalignathus Solodovnikov, 2005 , all endemic in the Afrotropical region, belong to the non‐related ‘Staphylinina’, ‘Philonthina propria’ and ‘Tanygnathinina sensu novo’ lineages of Staphylinini, respectively. Contrary to earlier records, the genus Quedius Stephens, 1929 does not occur in Africa south of Sahara: Quedius angularis Cameron, 1948 and Quedius cinctipennis Cameron, 1951 are moved to the genus Philonthus Stephens, 1829. The same is established for the Asian genus Algon Sharp, 1874, formerly for a long time associated with Quediina: African species Algon robustus Wendeler, 1928 is moved to the genus Moeocerus Fauvel, 1899 (here in the ‘Philonthina propria’ lineage); and the misidentification of Algon africanus Bernhauer, 1915, a species that probably belongs to a new genus, is discussed. The phylogenetic affiliation of Afroquedius Solodovnikov, 2006 , a South African endemic, is still ambiguous. Overall, the formerly seen bipolar distribution pattern for the ‘Quediina’ is demonstrated to be an artefact, not a reality to explain. Historical biogeographical explanations are proposed for some of the Afrotropical endemics, partly as an attempt to apply biogeography as an external criterion for the evaluation of the new phylogenetic pattern revealed for Staphylinini. The monotypic genera Euristus and Ioma, as well as Heterothops megalops Cameron, 1959 , the only representative of this widespread genus in the Afrotropical region, are redescribed. Limits and synapomorphies of the genus Heterothops are discussed. The following new combinations and new names are proposed: Philonthus cinctipennis ( Cameron, 1951 ) comb.n. (preoccupied by Philonthus cinctipennis Fauvel, 1875), here replaced by Philonthus pseudoquedius Solodovnikov nom.n. ; Philonthus angularis ( Cameron, 1948 ) comb.n. ; Moeocerus robustus ( Wendeler, 1928 ) comb.n. [preoccupied by Moeocerus robustus (Gestro, 1881)], here replaced by Moeocerus wendeleri Solodovnikov nom.n. A lectotype is designated for Heterothops megalops Cameron, 1959 .     

Phylogeny of Parabuthus (Scorpiones, Buthidae)

A cladistic analysis of the 20 southern African species of Parabuthus Pocock, 1 890 (Scorpiones, Buthidae) and five of the eight north-eastern African and Arabian species is presented, based on 53 characters, mostly of the adult morphology. The resultant topology is largely congruent with Lamoral's (1978) unpublished topology for 14 Namibian species. Monophyly of the genus Parabuthus is supported, but monophyly of the disjunct southern African vs. north-eastern African and Arabian species is unsupported. The implications of the cladogram for understanding ecological specialization in Parabuthus , Afrotropical arid biogeography and Parabuthus envenomation are discussed.     

Molecular evidence for the polyphyly of the genus Hemitragus (Mammalia, Bovidae)

The genus Hemitragus includes three species of tahr distributed in distant geographical areas: the Himalayan tahr, H. jemlahicus, occupies the southern flanks of the Himalaya Mountains; the Nilgiri tahr, H. hylocrius, is endemic to southern India; and the Arabian tahr, H. jayakari, is unique to the mountains of south-east Arabia. All previous investigations were based on morphology, and these three species together have never been included in a molecular phylogenetic study. In this study, we constructed a molecular phylogeny of the tribe Caprini sensu lato to determine the taxonomic status of the three species of tahr. Phylogenetic analyses were carried out on a matrix including most extant species currently described in the tribe Caprini sensu lato, and 3165 nucleotide characters, coming from four different markers, i.e., an intron of the nuclear gene coding for the protein kinase C iota, and three mitochondrial genes (subunit II of the cytochrome c oxidase, cytochrome b, and 12S rRNA). The results show that the genus Hemitragus is polyphyletic, as H. jemlahicus is associated with Capra (goats), H. hylocrius is the sister-group of Ovis (sheep), and H. jayakari is allied with Ammotragus lervia (aoudad). In the light of these unexpected results, we revaluate the validity of the morphological characters originally used for defining the genus Hemitragus. At least, we propose a new taxonomy, where the three species of tahr are ranged into three monospecific genera: the genus Hemitragus is restricted to the Himalayan tahr, and two new genera are created: Arabitragus for the Arabian tahr and Nilgiritragus for the Nilgiri tahr.     

Phylogeny and life history evolution of the genus Chrysoritis within the Aphnaeini (Lepidoptera: Lycaenidae), inferred from mitochondrial cytochrome oxidase I sequences

Phylogenetic relationships among 26 South African species in the tribe Aphnaeini (Lepidoptera: Lycaenidae) were inferred from DNA characters of the mitochondrial gene cytochrome oxidase I (COI), using maximum-parsimony methods. The resulting phylogenetic estimate supports the systematic hypothesis made by Heath (1997, Metamorphosis, supplement 2), based on morphological characters, that at least three preexisting genera (Chrysoritis, Poecilmitis, and Oxychaeta) should be collapsed into the single monophyletic genus Chrysoritis. Two of the species groups described by Heath within Chrysoritis are also monophyletic, while one is paraphyletic and thus unsupported by the molecular data. Strong node support and skewed transition/transversion ratios suggest that two Chrysoritis clades contain synonymous species. Aphytophagy appears as a derived feeding strategy. Evolutionary patterns of ant association indicate lability at the level of ant genus, while association with different ant subfamilies may have played an ancestral and chemically mediated role in the diversification of South African aphnaeines.     

A phylogeny of Delphinieae (Ranunculaceae) shows that Aconitum is nested within Delphinium and that Late Miocene transitions to long life cycles in the Himalayas and Southwest China coincide with bursts in diversification

The tribe Delphinieae (Ranunculaceae) comprises two species-rich genera, Aconitum and Delphinium, the latter including Consolida and Aconitella. The 650-700 species are distributed in Eurasia and North America; three species occur on tropical African mountains. Maximum likelihood analyses of 2088 aligned nucleotides of plastid and nuclear sequences obtained from up to 185 species of Delphinieae from throughout the geographic range (plus relevant outgroups) show that three short-lived (facultative annual or biennial) Mediterranean species belonging to Delphinium subgenus Staphisagria are the sister clade to all other Delphinieae, implying that Staphisagria needs to be raised to genus status if Delphinium and Aconitum are to become mutually monophyletic. Molecular clock dating suggests an origin of the sampled Delphinieae in the Early Oligocene (c. 32.3 Ma) and expansion to North America of Aconitum and Delphinium around 3.3 and 2.9 Ma ago, respectively; the East African Mts. were reached by long-distance dispersal some 2.4 Ma ago, coincident with the major uplift of the East African Rift system. The ancestral growth form of the Delphinieae could not be reconstructed, but Late Miocene bursts in diversification rates in the Himalayan and southwestern Chinese clades of Aconitum and Delphinium appear to be associated with transitions from short-lived to long-lived life histories.     

Revised classification and phylogeny of an Afrotropical species group based on molecular and morphological data,with the description of a new genus (Coleoptera: Scarabaeidae: Onthophagini)

The worldwide distributed Onthophagus genus comprises at present more than 2000 species, that often show a complicated and uncertain systematic history. In particular, the many Afrotropical species included in this genus have never been entirely reviewed after the division into 32 species groups proposed by d’Orbigny in 1913, although subsequent researches focusing on some of these species suggested that Onthophagus constituted a nonmonophyletic taxon. In order to highlight their phylogenetic relationships, the various Afrotropical species groups of d’Orbigny must thus be examined, and it would be advisable to study them separately to avoid misunderstanding. In this framework, the taxonomic position of the three species currently included in the 21st d’Orbigny group was examined. Both morphological and biomolecular analyses contributed in confirming that these species (i.e., Onthophagus caffrarius d’Orbigny, 1902; Onthophagus quadraticeps Harold, 1867; and Onthophagus signatus Fåhraeus, 1857) constituted a well-defined monophyletic group that cannot be maintained within the genus Onthophagus. Therefore, the Kurtops gen.n. is here described to accommodate these Afrotropical species, that are nevertheless always included within the Onthophagini tribe. On the basis of the phylogenetic relationships here elucidated, it was also emphasized that the new genus is strictly related to Digitonthophagus and Phalops; thus, it was proposed to include the three genera into a single clade of suprageneric rank naming it as Phalops complex.     

Pugs of the tribe Eupitheciini (Lepidoptera,Geometridae): the history of study and distribution over biogeographic regions

The World fauna of the tribe Eupitheciini is the most species-rich in the family Geometridae. This tribe includes about 1900 species (almost 3000 species-group names) from 47 genera; about one third of the genera (15) are monotypic. The generic diversity of Eupitheciini is the highest in the Australian (38 genera, 11 of them endemic) and Oriental regions (32 genera, 4 endemic) and the lowest in the Neotropical Region (possibly one genus only). The faunas of different biogeographic regions can be arranged in following order by their species richness: the Palaearctic (487 species), Oriental (397), Neotropical (346), Australian (251), Afrotropical (198), and Nearctic Regions (166 species). Eupithecia is the most species-rich genus in the family Geometridae and the entire order Lepidoptera, and one of the largest genera in the whole World fauna of insects. The greatest number of species of this genus is recorded in the Palaearctic Region (466 species), where Eupithecia accounts for about 95% of the tribe Eupitheciini. The mainland of the Oriental Region (especially the Himalayas) is also very species-rich; however the proportion of the Eupithecia representatives decreases towards Malaysia, Sundaland, and the Australian Region (about 2% of the tribe). The Eupitheciini faunas have the greatest similarity at the generic level between the Oriental and Australian Regions (the Jaccard and Sørensen coefficient values being 0.62 and 0.77, respectively). The Palaearctic fauna is more similar to the Afrotropical and Oriental faunas at the genus-group level. On the whole, the fauna of the Nearctic Region is similar to that the West Palaearctic, with the exception of the fact that representatives of the genera Gymnoscelis and Chloroclystis are absent in North America, although two endemic genera Nasusina and Prorella are present. At the genus-group level, the Nearctic fauna of Eupitheciini is more similar to the Neotropical (the Jaccard and Sørensen coefficients 0.20 and 0.33, respectively) than to the Palaearctic fauna (0.17 and 0.29). The number of synonymies is very high in the tribe Eupitheciini because of the homogeneity of this group, whose species are difficult to identify without the use of elaborate anatomical techniques. Modern revisions, catalogues, surveys, and atlases on Eupitheciini are absent for many countries and large geographic regions. Revisions of pugs of the tribe Eupitheciini for some biogeographic regions are extremely difficult because of fragmentation of entomological collections including the type specimens of many species-group taxa. A large fraction of synonyms is characteristic of parts of the World with the best known faunas: Europe (64% of synonyms) and North America (39%). On the contrary, the lowest levels of synonymy are typical of the less known faunas of the regions situated at the equatorial latitudes, namely the Neotropical (9%) and Afrotropical (8%) ones.     

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.