Molecular phylogeny of the beetle tribe Oxypodini (Coleoptera: Staphylinidae: Aleocharinae)

This is the first study to comprehensively address the phylogeny of the tribe Oxypodini Thomson and its phylogenetic relationships to other tribes within the staphylinid subfamily Aleocharinae. Using the hitherto largest molecular dataset of Aleocharinae comprising of 4599 bp for representatives of 22 tribes, the Oxypodini are recovered as non‐monophyletic. Members of the tribe belong to three distantly related lineages within the Aleocharinae: (i) the Amarochara group as sister clade to the tribe Aleocharini, (ii) the subtribe Tachyusina within a clade that also includes the tribes Athetini and Hygronomini, (iii) all other Oxypodini in a clade that also includes the tribes Placusini, Hoplandriini and Liparocephalini. Based on the inferred phylogeny, five subtribes of the Oxypodini are recognized: Dinardina Mulsant & Rey, Meoticina Seevers, Microglottina Fenyes, Oxypodina Thomson and Phloeoporina Thomson. The following changes in the classification of the Aleocharinae are proposed: (i) Amarochara Thomson is removed from the Oxypodini and placed in the tribe Aleocharini; (ii) the subtribe Taxicerina Lohse of the Athetini is reinstated as tribe Taxicerini to include Discerota Mulsant & Rey, Halobrecta Thomson (both removed from the Oxypodini) and Taxicera Mulsant & Rey; (iii) the subtribe Tachyusina Thomson is excluded from the Oxypodini and provisionally treated as tribe Tachyusini; (iv) the oxypodine subtribe name Blepharhymenina Klimaszewski & Peck is placed in synonymy with the subtribe name Dinardina Mulsant & Rey.     

Independent origins of coastal colonization in the tribe Athetini (Coleoptera,Staphylinidae)

Less than 1% of Staphylinidae are known to be confined to coastal habitats. To explore the origins of coastal colonization within the tribe Athetini Casey, we present a revised molecular phylogeny. The dataset comprised partial mitochondrial COI, COII, 16S rDNA, NADH1, partial nuclear 18S rDNA and 28S rDNA. We chose a total of 95 species in 51 genera, including 14 coastal species in eight genera and 21 outgroup species from other aleocharine tribes. The concatenated dataset was analysed simultaneously by both parsimony‐ and model‐based (Bayesian and maximum likelihood) methods. The tribe Athetini was not supported as a monophyletic group, but together with the tribes Tachyusini, Ecitocharini and Hygronomini did form a monophylum. The ecological association of species with a coastal habitat was mapped onto a phylogeny to assess the evolution of habitat specialization in the Athetini lineage. The results reveal that five independent origins of coastal colonization have occurred throughout the tribe Athetini: (a) Osakatheta + Adota minuta + coastal Atheta (Badura) (clade A); (b) Adota (clade B); (c) Pontomalota + Tarphiota + Thinusa (clade C); (d) Iotarphia (clade D); and (e) Psammostiba (clade E). The low species number of the coastal Athetini compared with the entire Athetini lineage indicates that coastal habitats are harsh environments and so only a few species were able to colonize this habitat. The following changes in classification are proposed: (a) Ad. minuta Lee and Ahn is removed from the genus Adota and tentatively included in Atheta (Badura); (b) The genus Saphocallus Sharp is transferred from Athetini to Geostibini.     

Exploring myrmecophily based on the phylogenetic interrelationships of Falagonia Sharp, 1883 (Coleoptera: Staphylinidae: Aleocharinae) and allied genera

The taxonomy of Lomechusini Fleming has a complex history. Recent studies have shown that this group is polyphyletic; however, little is known about the evolutionary interrelationships among its constituent genera. The goals of the present study are to infer the phylogenetic relationships of Falagonia Sharp and closely related genera; to define the boundaries of those genera based on synapomorphic characters; and to explore the evolution of myrmecophily within the lineage. The phylogenetic analyses are based exclusively on morphological characters of adults. A total of 36 operational taxonomic units were used for the analysis. The best trees were selected based on maximum parsimony and Bayesian inference. During the parsimony reconstruction, different weighting strategies were used to recover the most robust phylogenetic hypothesis. Although minor differences were observed in the results of the different analyses, the topologies were consistent throughout. Several groups of genera proposed by Seevers (1965), such as the ‘Tetradonia’ and ‘Ecitopora’ groups, were not recovered. Thus, these may represent nonmonophyletic groups that were based on nonsynapomorphic diagnostic characters. Our analyses consistently recovered the genera Asheidium Santiago‐Jiménez, Delgadoidium Santiago‐Jiménez, Falagonia, Newtonidium Santiago‐Jiménez, Pseudofalagonia Santiago‐Jiménez, Sharpidium Santiago‐Jiménez, Tetradonia Wasmann and Thayeridium Santiago‐Jiménez, forming a monophyletic group that we have called the ‘Asheidium complex’. Falagonia mexicana Sharp shows seven autapomorphies, none of which were used to establish the genus. Based on the phylogenetic results, myrmecophily has evolved independently at least three times within the lineage. This study, based on morphological characters, is one of the first approaches towards gaining an understanding of the phylogenetic relationships within the polyphyletic tribe Lomechusini.     

Phylogenetic structure in the grass family (Poaceae) as inferred from chloroplast DNA restriction site variation

A cladistic analysis of chloroplast DNA restriction site variation among representatives of all subfamilies of the grass family (Poaceae), using Joinvillea (Joinvilleaceae) as the outgroup, placed most genera into two major clades. The first of these groups corresponds to a broadly circumscribed subfamily Pooideae that includes all sampled representatives of Ampelodesmeae, Aveneae, Brachypodieae, Bromeae, Diarrheneae, Meliceae, Poeae, Stipeae, and Triticeae. The second major clade includes all sampled representatives of four subfamilies (Panicoideae [tribes Andropogoneae and Paniceae], Arundinoideae [Arundineae], Chloridoideae [Eragrostideae], and Centothecoideae [Centotheceae]). Within this group (the “PACC” clade), the Panicoideae are resolved as monophyletic and as the sister group of the clade that comprises the other three subfamilies. Within the latter group, Danthonia (Arundinoideae) and Eragroslis (Chloridoideae) are resolved as a stable monophyletic group that excludes Phragmites (Arundinoideae); this structure is inconsistent with the Arundinoideae being monophyletic as currently circumscribed. The PACC clade is placed within a more inclusive though unstable clade that includes the woody Bambusoideae (Bambuseae) plus several disparate tribes of herbaceous grasses of uncertain affinity that are often recognized as herbaceous Bambusoideae (Brachyelytreae, Nardeae, Olyreae, Oryzeae, and Phareae). Among eight most-parsimonious trees resolved by the analysis, four include a monophyletic Bambusoideae sensu lato (comprising Bambuseae and all five of these herbaceous tribes) as the sister group of the PACC clade; in the other four trees these bambusoid elements are not resolved as monophyletic, and the PACC clade is nested among these tribes. These results are consistent with those of previous analyses that resolve a basal or near-basal branch within the family between Pooideae and all other grasses. However, resolution by the present analysis of the PACC clade, which includes Centothecoideae, Chloridoideae, and Panicoideae, but excludes Bambusoideae, is inconsistent with the results of previous analyses that place Bambusoideae and Panicoideae in a monophyletic group that excludes Centothecoideae and Chloridoideae.     

Molecular phylogenetics of Braconidae (Hymenoptera: Ichneumonoidea), based on multiple nuclear genes,and implications for classification

This study examined subfamilial relationships within Braconidae, using 4 kb of sequence data for 139 taxa. Genetic sampling included previously used markers for phylogenetic studies of Braconidae (28S and 18S rDNA) as well as new nuclear protein‐coding genes (CAD and ACC). Maximum likelihood and Bayesian inference of the concatenated dataset recovered a robust phylogeny, particularly for early divergences within the family. This study focused primarily on non‐cyclostome subfamilies, but the monophyly of the cyclostome complex was strongly supported. There was evidence supporting an independent clade, termed the aphidioid complex, as sister to the cyclostome complex of subfamilies. Maxfischeria was removed from Helconinae and placed within its own subfamily within the aphidioid complex. Most relationships within the cyclostome complex were poorly supported, probably because of lower taxonomic sampling within this group. Similar to other studies, there was strong support for the alysioid subcomplex containing Gnamptodontinae, Alysiinae, Opiinae and Exothecinae. Cenocoeliinae was recovered as sister to all other subfamilies within the euphoroid complex. Planitorus and Mannokeraia, previously placed in Betylobraconinae and Masoninae, respectively, were moved to the Euphorinae, and may share a close affiliation with Neoneurinae. Neoneurinae and Ecnomiinae were placed as tribes within Euphorinae. A sister relationship between the microgastroid and sigalphoid complexes was also recovered. The helconoid complex included a well‐supported lineage that is parasitic on lepidopteran larvae (macrocentroid subcomplex). Helconini was raised to subfamily status, and was recovered as sister to the macrocentroid subcomplex. Blacinae was demoted to tribal status and placed within the newly circumscribed subfamily Brachistinae, which also contains the tribes Diospilini, Brulleiini and Brachistini, all formerly in Helconinae.     

Phylogeny of Aphantochilinae and Strophiinae sensu Simon (Araneae; Thomisidae)

This study tests the monophyly of ant‐mimicking Thomisidae (Aphantochilinae and Strophiinae sensu Simon), redefines the composition of these taxa, proposes tribes and discusses aspects of their myrmecomorphy and biogeography. The analysis is based on a matrix composed of 113 morphological characters and 37 terminal taxa (11 Aphantochilinae, 16 Strophiinae and 10 belonging to the out‐group). The 12 most parsimonious trees with 232 steps, obtained with equally weighted characters, support the monophyly of Aphantochilinae sensu Simon. Strophiinae emerges as a paraphyletic group divided into two clades: a basal clade that groups Strophius and Strigoplus (Strophiini new status) and another clade that includes Ceraarachne, Simorcus and Ulocymus (Ceraarachnini new status) as the sister group of Aphantochilus + Bucranium (Aphantochilini new status). Diagnoses are presented for the tribes and genera in this analysis. The synonymy between Bucranium and Aphantochilus is rejected. Majellula and Acracanthostoma are considered junior synonyms of Bucranium, and Synstrophius of Ceraarachne. The monophyly of Synstrophius is not recovered, S. blanci is transferred to Ceraarachne and S. muricatus is transferred to Ulocymus. Ant‐preying behaviour appears to be basal and has been documented for Strophiini and Aphantochilini species. Myrmecomorphy, which was documented for Aphantochilus, is presumably derived. The biogeographical analysis of Aphantochilinae and Strophiinae suggests an ancient relation between Neotropical, Afrotropical and Oriental species, with probable origin after the breakup of Gondwana, that is, in early Paleogene.     

Phylogeny,taxonomy, and evolution of Neotropical cichlids (Teleostei: Cichlidae: Cichlinae)

Despite recent progress on the higher‐level relationships of Cichlidae and its Indian, Malagasy, and Greater Antillean components, conflict and uncertainty remain within the species‐rich African, South American, and Middle American assemblages. Herein, we combine morphological and nucleotide characters from the mitochondrial large ribosomal subunit, cytochrome c oxidase subunit I, NADH dehydrogenase four, and cytochrome b genes and from the nuclear histone H3, recombination activating gene two, Tmo‐4C4, Tmo‐M27, and ribosomal S7 loci to analyse relationships within the Neotropical cichlid subfamily Cichlinae. The simultaneous analysis of 6309 characters for 90 terminals, including representatives of all major cichlid lineages and all Neotropical genera, resulted in the first well‐supported and resolved generic‐level phylogeny for Neotropical cichlids. The Neotropical subfamily Cichlinae was recovered as monophyletic and partitioned into seven tribes: Astronotini, Chaetobranchini, Cichlasomatini, Cichlini, Geophagini, Heroini, and Retroculini. Chaetobranchini + Geophagini (including the “crenicichlines”) was resolved as the sister group of Heroini + Cichlasomatini (including Acaronia). The monogeneric Astronotini was recovered as the sister group of these four tribes. Finally, a clade composed of Cichlini + Retroculini was resolved as the sister group to all other cichlines. The analysis included the recently described ?Proterocara argentina, the oldest known cichlid fossil (Eocene), which was placed in an apical position within Geophagini, further supporting a Gondwanan origin for Cichlidae. These phylogenetic results were used as the basis for generating a monophyletic cichline taxonomy. © The Willi Hennig Society 2008.     

Evolutionary relationships in the Asteraceae tribe Inuleae (incl. Plucheeae) evidenced by DNA sequences of ndhF; with notes on the systematic positions of some aberrant genera

The phylogenetic relationships between the tribes Inuleae sensu stricto and Plucheeae are investigated by analysis of sequence data from the cpDNA gene ndhF. The delimitation between the two tribes is elucidated, and the systematic positions of a number of genera associated with these groups, i.e. genera with either aberrant morphological characters or a debated systematic position, are clarified. Together, the Inuleae and Plucheeae form a monophyletic group in which the majority of genera of Inuleae s.str. form one clade, and all the taxa from the Plucheeae together with the genera Antiphiona, Calostephane, Geigeria, Ondetia, Pechuel-loeschea, Pegolettia, and Iphionopsis from Inuleae s.str. form another. Members of the Plucheeae are nested with genera of the Inuleae s.str., and support for the Plucheeae clade is weak. Consequently, the latter cannot be maintained and the two groups are treated as one tribe, Inuleae, with the two subtribes Inulinae and Plucheinae. The genera Asteriscus, Chrysophthalmum, Inula, Laggera, Pentanema, Pluchea, and Pulicaria are demonstrated to be non-monophyletic. Cratystylis and Iphionopsis are found to belong to the same clade as the taxa of the former Plucheeae. Caesulia is shown to be a close relative of Duhaldea and Blumea of the Inuleae-Inulinae. The genera Callilepis and Zoutpansbergia belong to the major clade of the family that includes the tribes Heliantheae sensu lato and Inuleae (incl. Plucheeae), but their exact position remains unresolved. The genus Gymnarrhena is not part of the Inuleae, but is either part of the unresolved basal complex of the paraphyletic Cichorioideae, or sister to the entire Asteroideae.     

Phylogenetic relationships of Combretoideae (Combretaceae) inferred from plastid,nuclear gene and spacer sequences

Phylogenetic relationships of the subfamily Combretoideae (Combretaceae) were studied based on DNA sequences of nuclear ribosomal internal transcribed spacer (ITS) regions, the plastid rbcL gene and the intergenic spacer between the psaA and ycf3 genes (PY-IGS), including 16 species of eight genera within two traditional tribes of Combretoideae, and two species of the subfamily Strephonematoideae of Combretaceae as outgroups. Phylogenetic trees based on the three data sets (ITS, rbcL, and PY-IGS) were generated by using maximum parsimony (MP) and maximum likelihood (ML) analyses. Partition-homogeneity tests indicated that the three data sets and the combined data set are homogeneous. In the combined phylogenetic trees, all ingroup taxa are divided into two main clades, which correspond to the two tribes Laguncularieae and Combreteae. In the Laguncularieae clade, two mangrove genera, Lumnitzera and Laguncularia, are shown to be sister taxa. In the tribe Combreteae, two major clades can be classified: one includes three genera Quisqualis, Combretum and Calycopteris, within which the monophyly of the tribe Combreteae sensu Engler and Diels including Quisqualis and Combretum is strongly supported, and this monophyly is then sister to the monotypic genus Calycopteris; another major clade includes three genera Anogeissus, Terminalia and Conocarpus. There is no support for the monophyly of Terminalia as it forms a polytomy with Anogeissus. This clade is sister to Conocarpus. Electronic Publication     

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.     

Molecular Phylogeny of Rhizophoraceae Based on rbcL Gene Sequences

rbcL sequences to clarify the inter- and intrarelationships of Rhizophoraceae which have been variously discussed. The analyses included 12 of the 15 genera of Rhizophoraceae (4/7 of Macarisieae, 4/4 of Gynotrocheae, and 4/4 of Rhizophoreae) and a few putatively related taxa, including two of the four genera of Anisophylleaceae. The most parsimonious trees supported the monophyly of Rhizophoraceae as well as each of the three traditionally recognized tribes Macarisieae, Gynotrocheae, and Rhizophoreae. The family Rhizophoraceae is a sister taxon to Erythroxylum (Erythroxylaceae) and is further closely related to Byrsonima (Malpighiaceae), Passiflora (Passifloraceae), Turnera (Turneraceae), Ochna (Ochnaceae), Drypetes (Euphorbiaceae), and Humiria (Humiriaceae). Anisophylleaceae, which have often been included in Rhizophoraceae as a tribe or subfamily, are placed in a common clade with Begonia (Begoniaceae), Cucurbita (Cucurbitaceae), Coriaria (Coriariaceae), Corynocarpus (Corynocarpaceae), Datisca (Datiscaceae), Tetrameles (Datiscaceae), and Octomeles (Datiscaceae). Within Rhizophoraceae the mangrove tribe Rhizophoreae is sister to the inland tribe Gynotrocheae, with inland tribe Macarisieae positioned as a sister taxon to these two tribes. This pattern of relationships within the family basically agrees with those suggested by cladistic analyses based on morphological characters, except that Gynotrocheae are monophyletic with Crossostylis as a derived taxon within the tribe in the present study. Based on this cladogram for Rhizophoraceae, we discuss evolutionary trends of a few ecological and morphological characters, including the formation of aerial roots and the ovary position. Received 12 August 1999/ Accepted in revised form 11 October 1999     

First phylogeny of predatory flower flies (Diptera, Syrphidae, Syrphinae) using mitochondrial COI and nuclear 28S rRNA genes: conflict and congruence with the current tribal classification

The family Syrphidae (Diptera) is traditionally divided into three subfamilies. The aim of this study was to address the monophyly of the tribes within the subfamily Syrphinae (virtually all with predaceous habits), as well as the phylogenetic placement of particular genera using molecular characters. Sequence data from the mitochondrial protein-coding gene cytochrome c oxidase subunit I ( COI ) and the nuclear 28S ribosomal RNA gene of 98 Syrphinae taxa were analyzed using optimization alignment to explore phylogenetic relationships among included taxa. Volucella pellucens was used as outgroup, and representatives of the tribe Pipizini (Eristalinae), with similar larval feeding mode, were also included. Congruence of our results with current tribal classification of Syrphinae is discussed. Our results include the tribe Toxomerini resolved as monophyletic but placed in a clade with genera Ocyptamus and Eosalpingogaster . Some genera traditionally placed into Syrphini were resolved outside of this tribe, as the sister groups to other tribes or genera. The tribe Bacchini was resolved into several different clades. We recovered Paragini as a monophyletic group, and sister group of the genus Allobaccha . The present results highlight the need of a reclassification of Syrphinae.
© The Willi Hennig Society 2008.     

The systematic position of Meruidae (Coleoptera, Adephaga) and the phylogeny of the smaller aquatic adephagan beetle families

A phylogenetic analysis of Adephaga is presented. It is based on 148 morphological characters of adults and larvae and focussed on a placement of the recently described Meruidae, and the genus‐level phylogeny of the smaller aquatic families Gyrinidae, Haliplidae and Noteridae. We found a sister group relationship between Gyrinidae and the remaining adephagan families, as was found in previous studies using morphology. Haliplidae are either the sister group of Dytiscoidea or the sister group of a clade comprising Geadephaga and the dytiscoid families. Trachypachidae was placed as the sister group of the rhysodid‐carabid clade or of Dytiscoidea. The monophyly of Dytiscoidea including Meru is well supported. Autapomorphies are the extensive metathoracic intercoxal septum, the origin of the metafurca from this structure, the loss of Mm. furcacoxalis anterior and posterior, and possibly the presence of an elongated subcubital setal binding patch. Meruidae was placed as sister group of the Noteridae. Synapomorphies are the absence of the transverse ridge of the metaventrite, the fusion of abdominal segments III and IV, the shape of the strongly asymmetric parameres, and the enlargement of antennomeres 5, 7 and 9. The Meru‐noterid clade is the sister group of the remaining Dytiscoidea. The exact position of Aspidytes within this clade remains ambiguous: it is either the sister group of Amphizoidae or the sister group of a clade comprising this family and Hygrobiidae + Dytiscidae. The sister group relationship between Spanglerogyrinae and Gyrininae was strongly supported. The two included genera of Gyrinini form a clade, and Enhydrini are the sister group of a monophylum comprising the remaining Enhydrini and Orectochilini. A branching pattern (Peltodytes + (Brychius + Haliplus)) within Haliplidae was confirmed. Algophilus, Apteraliplus and the Haliplus‐subgenus Liaphlus form a clade. The generic status of the two former taxa is unjustified. The Phreatodytinae are the sister group of Noterinae, and Notomicrus (+ Speonoterus), Hydrocoptus, and Pronoterus branch off successively within this subfamily. The search for the larvae of Meru and a combined analysis of morphological and molecular data should have high priority. © The Willi Hennig Society 2006.     

Molecular phylogeny of the Byrrhoidea–Buprestoidea complex (Coleoptera,Elateriformia)

The superfamilies of Elateriformia have been in a state of flux since their establishment. The recent classifications recognize Dascilloidea, Buprestoidea, Byrrhoidea and Elateroidea. The most problematic part of the elateriform phylogeny is the monophyly of Byrrhoidea and the relationships of its families. To investigate these issues, we merged more than 500 newly produced sequences of 18S rRNA, 28S rRNA, rrnL mtDNA and cox1 mtDNA for 140 elateriform taxa with data from GenBank. We assembled an all‐taxa (488 terminals) and a pruned data set, which included taxa with full fragment representation (251 terminals); both were aligned in various programs and analysed using maximum‐likelihood criterion and Bayesian inference. Most analyses recovered monophyletic superfamilies and broadly similar relationships; however, we obtained limited statistical support for the backbone of trees. Dascilloidea were sister to the remaining Elateriformia, and Elateroidea were sister to the clade of byrrhoid lineages including Buprestoidea. This clade mostly consisted of four major lineages, that is (i) Byrrhidae, (ii) Dryopidae + Lutrochidae, (iii) Buprestoidea (Schizopodidae sister to Buprestidae) and (iv) a clade formed by the remaining byrrhoid families. Buprestoidea and byrrhoid lineages, with the exception of Byrrhidae and Dryopidae + Lutrochidae, were usually merged into a single clade. Most byrrhoid families were recovered as monophyletic. Callirhipidae and Eulichadidae formed independent terminal lineages within the Byrrhoidea–Buprestoidea clade. Paraphyletic Limnichidae were found in a clade with Heteroceridae and often also with Chelonariidae. Psephenidae, represented by Eubriinae and Eubrianacinae, never formed a monophylum. Ptilodactylidae were monophyletic only when Paralichas (Cladotominae) was excluded. Elmidae regularly formed a clade with a bulk of Ptilodactylidae; however, elmid subfamilies (Elminae and Larainae) were not recovered. Despite the densest sampling of Byrrhoidea diversity up to date, the results are not statistically supported and resolved only a limited number of relationships. Furthermore, questions arose which should be considered in the future studies on byrrhoid phylogeny.     

Taxonomic studies in Chiloscyphus Corda (Jungermanniales: Lophocoleaceae) based on nrITS sequences and morphology

Maximum parsimony and likelihood analyses of 40 Lophocoleaceae nrITS sequences and 6 Plagiochilaceae sequences (outgroup) lead to a robust phylogeny of Chiloscyphus. Four main lineages are assigned to as Chiloscyphus subgenera Chiloscyphus, Lophocolea, Connati and Notholophocolea. Chiloscyphus subgen. Connati is resolved sister to the remainder of this genus. Chiloscyphus subgenus Lophocolea is subdivided into sections Heterophylli (incl. sect. Semiteretes, syn. nov.), Lophocolea, Microlophocolea, and Novae-Zeelandiae. Five accessions of Chiloscyphus pallescens with a chromosome number of n = 18 form a robust monophyletic lineage that is placed sister to a well supported clade with 4 accessions of C. polyanthos [n = 9]. Chiloscyphus mandonii is placed in the synonymy of C. latifolius.     

A New Heterotrophic Cryptomonad: Hemiarma marina n. g., n. sp.

We report a new heterotrophic cryptomonad Hemiarma marina n. g., n. sp. that was collected from a seaweed sample from the Republic of Palau. In our molecular phylogenetic analyses using the small subunit ribosomal RNA gene, H. marina formed a clade with two marine environmental sequences, and the clade was placed as a sister lineage of the freshwater cryptomonad environmental clade CRY1. Alternatively, in the concatenated large and small subunit ribosomal RNA gene phylogeny, H. marina was placed as a sister lineage of Goniomonas. Light and electron microscopic observations showed that H. marina shares several ultrastructural features with cryptomonads, such as flattened mitochondrial cristae, a periplast cell covering, and ejectisomes that consist of two coiled ribbon structures. On the other hand, H. marina exhibited unique behaviors, such as attaching to substrates with its posterior flagellum and displaying a jumping motion. H. marina also had unique periplast arrangement and flagellar transitional region. On the basis of both molecular and morphological information, we concluded that H. marina should be treated as new genus and species of cryptomonads.     

An updated classification of Orchidaceae

Since the last classification of Orchidaceae in 2003, there has been major progress in the determination of relationships, and we present here a revised classification including a list of all 736 currently recognized genera. A number of generic changes have occurred in Orchideae (Orchidoideae), but the majority of changes have occurred in Epidendroideae. In the latter, almost all of the problematic placements recognized in the previous classification 11 years ago have now been resolved. In Epidendroideae, we have recognized three new tribes (relative to the last classification): Thaieae (monogeneric) for Thaia, which was previously considered to be the only taxon incertae sedis; Xerorchideae (monogeneric) for Xerorchis; and Wullschlaegelieae for achlorophyllous Wullschlaegelia, which had tentatively been placed in Calypsoeae. Another genus, Devogelia, takes the place of Thaia as incertae sedis in Epidendroideae. Gastrodieae are clearly placed among the tribes in the neottioid grade, with Neottieae sister to the remainder of Epidendroideae. Arethuseae are sister to the rest of the higher Epidendroideae, which is unsurprising given their mostly soft pollinia. Tribal relationships within Epidendroideae have been much clarified by analyses of multiple plastid DNA regions and the low‐copy nuclear gene Xdh. Four major clades within the remainder of Epidendroideae are recognized: Vandeae/Podochileae/Collabieae, Cymbidieae, Malaxideae and Epidendreae, the last now including Calypsoinae (previously recognized as a tribe on its own) and Agrostophyllinae s.s. Agrostophyllinae and Collabiinae were unplaced subtribes in the 2003 classification. The former are now split between two subtribes, Agrostophyllinae s.s. and Adrorhizinae, the first now included in Epidendreae and the second in Vandeae. Collabiinae, also probably related to Vandeae, are now elevated to a tribe along with Podochileae. Malaxis and relatives are placed in Malaxidinae and included with Dendrobiinae in Malaxideae. The increased resolution and content of larger clades, recognized here as tribes, do not support the ‘phylads’ in Epidendroideae proposed 22 years ago by Dressler. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 151–174.     

Morphology, molecules, and character congruence in the phylogeny of South American geophagine cichlids (Perciformes, Labroidei)

Phylogenetic relationships among the Neotropical cichlid subfamily Geophaginae were examined using 136 morphological characters and a molecular dataset consisting of six mitochondrial and nuclear genes. Topologies produced by morphological and combined data under parsimony were contrasted, congruence among different partitions was analysed, and potential effects of character incongruence and patterns of geophagine evolution on phylogenetic resolution are discussed. Interaction of morphological and molecular characters in combined analysis produced better resolved and supported topologies than when either was analysed separately. Combined analyses recovered a strongly supported Geophaginae that was closely related to Cichlasomatinae. Within Geophaginae, two sister clades included all geophagine genera. Acarichthyini (Acarichthys+Guianacara) was sister to the ‘B clade’, which contained the ‘Geophagus clade’ (‘Geophagus’steindachneri+Geophagus sensu stricto, and both sister to Gymnogeophagus) as sister to the ‘Mikrogeophagus clade’ (Mikrogeophagus+‘Geophagus’brasiliensis), and in turn, the Geophagus and Mikrogeophagus clades were sister to the crenicarine clade (Crenicara+Dicrossus) and Biotodoma. The second geophagine clade included the ‘Satanoperca clade’ (Satanoperca+Apistogramma and Taeniacara) as sister to the ‘Crenicichla clade’ (Crenicichla+Biotoecus). Several lineages were supported by unique morphological synapomorphies: the Geophaginae + Cichlasomatinae (5 synapomorphies), Geophaginae (1), Crenicichla clade (3), crenicarine clade (1), the sister relationship of Apistogramma and Taeniacara (4) and of Geophagus sensu stricto and‘Geophagus’steindachneri (1), and the cichlasomine tribe Heroini (1). Incorporation of Crenicichla in Geophaginae reconciles formerly contradictory hypotheses based on morphological and molecular data, and makes the subfamily the most diverse and ecologically versatile clade of cichlids outside the African great lakes. Results of this study support the hypothesis that morphological differentiation of geophagine lineages occurred rapidly as part of an adaptive radiation.     

Emergence of a superradiation: pselaphine rove beetles in mid‐Cretaceous amber from Myanmar and their evolutionary implications

Pselaphinae is an exceptionally species‐rich, globally distributed subfamily of minute rove beetles (Staphylinidae), many of which are inquilines of social insects. Deducing the factors that drove pselaphine diversification and their evolutionary predisposition to inquilinism requires a reliable timescale of pselaphine cladogenesis. Pselaphinae is split into a small and highly plesiomorphic supertribe, Faronitae, and its sister group, the ‘higher Pselaphinae’ – a vast multi‐tribe clade with a more derived morphological ground plan, and which includes all known instances of inquilinism. The higher Pselaphinae is dominated by tribes with a Gondwanan taxonomic bias. However, a minority of tribes are limited to the Nearctic and Palearctic ecozones, implying a potentially older, Pangaean origin of the higher Pselaphinae as a whole. Here, I describe fossils from mid‐Cretaceous (～99 million years old) Burmese amber that confirm the existence of crown‐group higher pselaphines on the Eurasian supercontinent prior to contact with Gondwanan landmasses. Protrichonyx rafifrons gen. et sp.n. is placed incertae sedis within the higher Pselaphinae. Boreotethys gen.n. , erected for B. grimaldii sp.n . and B. arctopteryx sp.n. , represents an extinct sister taxon and putative stem group of Bythinini, a Recent tribe with a primarily Holarctic distribution. The Laurasian palaeolocality of the newly described taxa implies that higher pselaphines are indeed probably of Jurassic, Pangaean extraction and that the Laurasian‐Gondwanan tribal dichotomy of this clade may have developed vicariantly following Pangaean rifting. Higher pselaphines probably predate the earliest ants. Their physically protective morphological ground plan may have been a preadaptation for myrmecophily when ants became diverse and ecologically ubiquitous, much later in the Cenozoic. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:36E3FE2A-B947-422D-89CA-0EF43B99C382 .