Superfamily Membracoidea (Homoptera: Auchenorrhyncha). II. Cladistic analysis and conclusions

Abstract. Homologies among traditional morphological characters in the Membracoidea ( sensu lato ) are reassessed and the phylogenetic relationships among higher membracoid taxa are explored, incorporating new morphological evidence from nymphs and adults. Weighted and unweighted parsimony analyses of a matrix of sixty–three characters and thirty-nine OTUs representing the families Aetalionidae, Cicadellidae, Melizoderidae and Membracidae, and an outgroup (superfamily Cercopoidea) yielded various topologies that were largely congruent but presented alternative hypotheses of relationships among the Membracidae. These analyses indicate that the superfamily consists of the following clades: Cicadellidae + (Melizoderidae + (Aetalionidae + Membracidae)). The family Membracidae, traditionally characterized by the presence of a posterior pronotal process, apparently gave rise to Nicomia Stål and other genera that lack this process.     

Revision of the genera Hovadelium Ardoin and Mimolaena Ardoin (Coleoptera,Tenebrionidae, Laenini) from Madagascar,with remarks on tribal assignment

The genera Hovadelium Ardoin, 1961 and Mimolaena Ardoin, 1961, endemic in Madagascar, are revised and assigned to the tribe Laenini Seidlitz, 1896 (subfamily Lagriinae Latreille, 1825). New species: Hovadelium ardoini sp. n., Hovadelium bremeri sp. n. and Mimolaena janaki sp. n. An identification key is compiled for all taxa. Distribution of Hovadelium (5 species) and Mimolaena (3 species) is mapped. The congeners might be indicator species for the highly endangered mature forests in Madagascar.     

Phylogeny and classification of the leafhopper subfamily Eurymelinae (Hemiptera: Cicadellidae) inferred from molecules and morphology

Phylogenetic analysis of the globally distributed arboreal leafhopper subfamily Eurymelinae was conducted based on DNA sequence data from three nuclear and two mitochondrial genes in addition to 86 discrete morphological characters. The analysis included 89 species representing 61 genera from all major biogeographic regions including six species from outgroups, Megophthalminae and Ulopinae. Trees resulting from partitioned Bayesian and maximum likelihood analyses of the combined data were well resolved and largely congruent, differing mainly in the relationships among the earliest diverging lineages. The results are consistent with an expanded concept of Eurymelinae, including tribes Austroagalloidini and Macropsini. Six monophyletic groups are recognized as new tribes, Balocerini, Chiasmodolini, Chileanoscopini, Idioceroidini, Kopamerrini and Nesocerini, tribe n. , and the previously recognized tribes Eurymelini, Idiocerini and Megipocerini are redefined. A new synonym, Busonini Zhang & Li, 2015 syn.n. is proposed here for Megipocerini Isaev, 1988. Molecular divergence time estimates were calibrated using two fossil taxa and suggested that the earliest divergences occurred in the Lower Cretaceous and that most major lineages of this group arose during the Cretaceous. Reconstruction of ancestral areas revealed considerable continental-scale biogeographical structure. The place of origin of Eurymelinae is equivocal but major lineages arose in the Neotropical, Australian and Afrotropical regions. A key to tribes and a checklist of genera showing current tribal placements are provided.     

Molecular phylogeny of Sterrhinae moths (Lepidoptera: Geometridae): towards a global classification

A multigene phylogenetic study was carried out to test current, mostly morphology-based hypotheses on Sterrhinae phylogeny with additional material included from further geographical areas and morphologically different lineages. A maximum likelihood analysis (11 molecular markers and 7665 bp) was conducted on 76 species and 41 genera using iq-tree software. The resulting phylogenetic hypothesis is well resolved and branches have high support values. Results generally agree with earlier hypotheses at tribal levels and support the hypothesis that Sterrhinae comprises two major lineages. Based on the molecular phylogeny and extensive morphological examination, nine tribes are considered valid and the following taxonomic changes are introduced to recognize monophyletic groups: Mecoceratini Guenée, 1858 (= Ametridini Prout, 1910) is transferred from Desmobathrinae to Sterrhinae, and it is considered valid at tribal level new classification ; Haemaleini Sihvonen & Brehm is described as a new tribe and deemed sister to Scopulini + Lissoblemmini; Lissoblemmini Sihvonen & Staude is described as a new tribe and sister to Scopulini; Lythriini Herbulot, 1962 is now a junior synonym of Rhodometrini Agenjo, 1952 syn.n. ; and Rhodostrophiini Prout, 1935 is now a junior synonym of Cyllopodini Kirby, 1892 syn.n. In addition, 48 taxa are transferred from other geometrid subfamilies to Sterrhinae, or within Sterrhinae from one tribe to another, or they are classified into a tribe for the first time, or a new genus classification is proposed. The results demonstrate the limited explanatory power of earlier classifications, particularly at the tribal level. This is probably a result of earlier classifications being based on superficial characters and biased towards the European and North American fauna. The species richness and distribution of Sterrhinae and its constituent tribes are reviewed, showing that the globally distributed Sterrhinae are most diverse in the Neotropics (31% of global fauna). They are species-rich in the Palaearctic (22%), Afrotropics (19%) and Indo-Malay (16%) regions, whereas they are almost absent in Oceania (1%). In terms of the described fauna, the most species-rich tribes are Scopulini (928 species), Sterrhini (876 species) and Cosymbiini (553 species), all of which have a cosmopolitan distribution. Mecoceratiini and Haemaleini are almost entirely Neotropical. Timandrini and Lissoblemmini, by contrast, are absent in the Neotropics. We present a revised classification of the global Sterrhinae fauna, which includes about 3000 putatively valid species, classified into nine tribes and 97 genera. Four genera are of uncertain position within Sterrhinae. Our results highlight the compelling need to include more genera from a global perspective in molecular phylogenetic studies, in order to create a stable global classification for this subfamily. This published work has been registered on ZooBank, http://zoobank.org/urn:lsid:zoobank.org :pub:A66F5DDD-06D6-4908-893E-E8B124BB99B1.     

The spider family Selenopidae (Arachnida, Araneae) in Australasia and the Oriental Region

We relimit and revise the family Selenopidae to include four new genera and 27 new species from Australia and the Oriental Region. The family is redefined, as are the genera Anyphops Benoit, Garcorops Corronca, Hovops Benoit, Selenops Latreille, and Siamspinops Dankittipakul & Corronca, to accommodate the new genera and to correct previous inconsistencies in the diagnoses and definitions of the aforementioned genera. The species of Selenops that occur throughout India and China are also reviewed. Three species occur in China: Selenops bursarius Karsch 1879, also known from Japan, Korea and Taiwan, Selenops ollarius Zhu, Sha, & Chen 1990, and Selenops radiatus Latreille 1819, the type of the genus and most widespread selenopid. Selenops cordatus Zhu, Sha & Chen syn. n. is recognized as a junior synonym of Selenops radiatus. Amamanganopsgen. n. is monotypic, with Amamanganops baginawasp. n. (♀; from the Philippines). Godumopsgen. n. is monotypic, with Godumops caritussp. n. (♂; from Papua New Guinea). Karaopsgen. n. occurs throughout Australia and includes 24 species. A new combination is proposed for Karaops australiensis (L. Koch 1875) comb. n. (ex. Selenops), and the new species: Karaops gangariesp. n. (♀, ♂), Karaops monteithisp. n. (♀), Karaops alanlongbottomisp. n. (♂), Karaops keithlongbottomisp. n. (♂), Karaops larryoosp. n. (♂), Karaops jarritsp. n. (♂,♀), Karaops marrayagongsp. n. (♀), Karaops ravenisp. n. (♂,♀), Karaops badgeraddasp. n. (♀), Karaops burbidgeisp. n. (♂,♀), Karaops karrawarlasp. n. (♂,♀), Karaops julianneaesp. n. (♀), Karaops martamartasp. n. (♀), Karaops manaaynsp. n. (♀, ♂), Karaops vadlaadambarasp. n. (♀, ♂), Karaops pilkingtonisp. n. (♀, ♂), Karaops deserticolasp. n. (♀), Karaops ngarutjaranyasp. n. (♂,♀), Karaops francesaesp. n. (♂,♀), Karaops toolbrunupsp. n. (♀, ♂), the type species Karaops ellenaesp. n. (♂,♀), Karaops jenniferaesp. n. (♀), and Karaops dawarasp. n. (♀).The genus Makdiopsgen. n. contains five species from India and Nepal. A new combination is proposed for Makdiops agumbensis (Tikader 1969), comb. n., Makdiops montigenus (Simon 1889), comb. n., Makdiops nilgirensis (Reimoser 1934) comb. n.,(ex. Selenops). Also, there are two new species the type of the genus Makdiops mahishasurasp. n. (♀; from India), and Makdiops shivasp. n. (♀). The genus Pakawopsgen. n. is monotypic. A new combination is proposed for Pakawops formosanus (Kayashima 1943) comb. n. (ex. Selenops), known only from Taiwan. A new combination is proposed for Siamspinops aculeatus (Simon)comb. n. (ex. Selenops). The distribution and diversity of the studied selenopid fauna is discussed. Finally, keys are provided to all of the selenopid genera and to the species of Karaopsgen. n.and Makdiopsgen. n.     

Comprehensive phylogeny of the Cleroidea (Coleoptera: Cucujiformia)

The first thorough molecular phylogeny of the superfamily Cleroidea, represented by 377 taxa, and the first with an emphasis on Trogossitidae, was undertaken. Maximum likelihood and Bayesian analyses were performed on a four‐gene dataset (18S, 28S, cox1, cytb) of 395 taxa (along with 18 outgroups), including all 16 currently recognized families of Cleroidea and all current and formerly recognized tribes of Trogossitidae. The superfamily as a whole received strong support in Bayesian analyses. On the basis of phylogenetic results, 18 families in Cleroidea are recognized, including three taxa elevated to family for the first time and two reinstated families. The former tribe Rentoniini (Trogossitidae: Peltinae) was strongly supported as a monophyletic group apart from the remainder of Trogossitidae, and is herein elevated to family status, Rentoniidae stat.n. Protopeltis was also found to be an isolated lineage and becomes Protopeltidae stat.n. Peltini + Larinotini were recovered as a weakly supported sister grouping; Peltini (including only Peltis) becomes Peltidae stat.rest. The trogossitid subfamily Lophocaterinae, to the exclusion of Decamerini, formed a clade which is here designated Lophocateridae stat.rest. and sensu n. The Trogossitinae tribes Calityini, Egoliini (represented by Egolia) and Larinotini were recovered apart from core Trogossitidae but showed no strong affinities to other taxa or congruence between analyses; they are here conservatively retained in Trogossitidae as Calityinae stat.rest. , Egoliinae stat.rest. and Larinotinae stat.rest. The genus Thymalus of the peltine tribe Thymalini was indicated with moderate to strong support as the sister group of the Decamerini (Trogossitidae: Lophocaterinae); together these represent Thymalidae stat.n. and sensu n. with subfamilies Decamerinae stat.rest. ( new placement ) and Thymalinae stat.n. The remainder of Trogossitinae, the tribes Trogossitini and Gymnochilini, formed a well‐supported clade which comprises the Trogossitidae: Trogossitinae sensu n. The tribe Gymnochilini syn.n. is synonymized with Trogossitini. The monotypic family Phloiophilidae was recovered, contradicting a recent placement within Trogossitidae. The melyrid lineage was recovered with moderate (maximum likelihood) to strong (Bayesian analyses) support and includes the families Phycosecidae, Rhadalidae, Mauroniscidae, Prionoceridae and Melyridae (including Dasytidae and Malachiidae). The genus Dasyrhadus is tentatively transferred from Rhadalidae to Mauroniscidae. The genus Gietella, once proposed as a distinct family but recently placed within Dasytidae, was recovered as strongly sister to Rhadalidae sensu n. , and we transfer it to that family as Gietellinae new placement . Attalomiminae (formerly Attalomimidae) syn.n. is synonymized with Melyridae: Malachiinae: Lemphini sensu n. Melyridae sensu n. includes only Dasytinae, Malachiinae and Melyrinae. Metaxina is returned to the Chaetosomatidae sensu n. , of which Metaxinidae syn.n. becomes a junior synonym. Resolution within Cleridae was generally poor, but a broadly defined Korynetinae stat rest. + Epiclininae received high support (Bayesian analyses). Outside of Trogossitidae, the main focus of this study, major rearrangements of the classification of Cleroidea were not undertaken, despite evidence indicating such changes are needed.     

Revision of the Plant Bug Genus Tytthus (Hemiptera,Heteroptera, Miridae,Phylinae)

The phyline plant bug genus Tytthus Fieber, previously containing 19 species, is revised. Isoproba Osborn and Drake, 1915, incorrectly placed in the subfamily Bryocorinae, tribe Dicyphini, is synonymized as a junior synonym of Tytthus Fieber, syn. n.; the only included species, Isoproba picea Osborn and Drake is transferred to Tytthus, comb. n., as the senior synonym of Tytthus hondurensis Carvalho, syn. n.; and Tytthus koreanus Josifov and Kerzhner, 1972 is synonymized with Tytthus chinensis (Stål 1860), syn. n.; and a lectotype for Tytthus parviceps is designated. The six new species Tytthus femoralis from Cuba, Ecuador, Guatemala, Jamaica, Mexico, and Peru,Tytthus fuscicornis from New Mexico (USA), Tytthus mexicanus from Mexico, Tytthus pallidus from Brazil and Panama, Tytthus uniformis from Arizona and New Mexico (USA), and Tytthus wheeleri from the eastern United States are described, bringing the total number of species for the genus to 24. A color adult habitus illustration of Tytthus wheeleri, color photographs for each species (except Tytthus juturnaiba Carvalho and Wallerstein), illustrations of male genitalia, scanning electron photomicrographs of selected structures of certain species, and an identification key are provided to facilitate species recognition. A phylogenetic analysis is offered to help infer relationships.     

The cladistics and biology of the Callajoppa genus-group (Hymenoptera: Ichneumonidae, Ichneumoninae)

A cladistic analysis is presented for the genera of the former ichneumonine tribe Trogini. The tribe Heresiarchini is paraphyletic with respect to the Trogini, and so maintaining Trogini as a separate tribe is unsatisfactory. Within Heresiarchini, the following changes are made: (a) the subtribes Apatetorina and Heresiarchina are referred to as the Apatetor and Heresiarches genus‐groups, (b) the genera of the paraphyletic subtribe Protichneumonina are treated as incerta sedis within Heresiarchini, and (c) the Trogini are referred to as the Callajoppa genus‐group, with the former subtribe Trogina referred to as the Trogus subgroup. Thirty‐five genera are recognized as valid within the Callajoppa genus‐group. Catadelphops, Catadelphus, Cobunus, and Facydes are transferred to this group; Holojoppa is removed and is incertae sedis within Heresiarchini. Three new synonyms are proposed: Araeoscelis and Cryptopyge are junior synonyms of Macrojoppa, and Neamblyjoppa is a junior synonym of Catadelphops. Trogus latipennis Cresson is transferred to Pedinopelte from Macrojoppa, and Trogus mactator Tosquinet and its related species (T. bicolor Radoszkowski, T. heinrichi Uchida, and T. tricephalus Uchida) are transferred to Holcojoppa. Tricyphus is redefined and a neotype is designated for Tricyphus cuspidiger Kriechbaumer, the type‐species of the genus. Thirteen new genera are described (authorship of all is Wahl & Sime): Charmedia (type‐species: Charmedia chavarriai Wahl & Sime, sp. n.) , Daggoo (type‐species: Daggoo philoctetes Wahl & Sime, sp. n.), Dothenia (type‐species: Dothenia hansoni Wahl & Sime, sp. n.), Humbert (type‐species: Humbert humberti Wahl & Sime, sp. n.), Laderrica (type‐species: Laderrica feenyi Wahl & Sime, sp. n.), Mokajoppa (type‐species: Tricyphus respinozai Ward & Gauld), Metallichneumon (type‐species: Metallichneumon neurospastarchus Wahl & Sime, sp. n.), Myocious (type‐species: Myocious orientalis Wahl & Sime, sp. n.) , Quandrus (type‐species: Trogus pepsoides Smith, transferred from Callajoppa), Queequeg (type‐species: Gathetus flavibasalis Uchida, transferred from Neofacydes), Saranaca (type‐species: Trogus elegans Cresson; includes Trogus apicalis Cresson, Tricyphyus ater Hopper, and Tricyphus floridanus Heinrich), Tashtego (type‐species: Tashtego janzeni Wahl & Sime, sp. n.), and Xanthosomnium (type‐species: Xanthosomnium froesei Wahl & Sime, sp. n.). A key to the genera of the Callajoppa genus‐group is provided. The evolution of biological traits within the Callajoppa genus‐group is discussed with reference to the elucidated phylogeny. The groundplan biology is parasitism of Sphingidae, with oviposition into a host pupa/prepupa. There have been two transitions to butterfly parasitism within the Trogus subgroup: one a transition to Papilionidae (followed by a switch to Nymphalidae at Psilomastax) and the other to Nymphalidae (followed by a switch to Papilionidae within Macrojoppa). ©2002 The Linnean Society of London. Zoological Journal of the Linnean Society, 134 , 1–56.