Ancient Divergence in Bathypelagic Lake Tanganyika Deepwater Cichlids: Mitochondrial Phylogeny of the Tribe Bathybatini

The cichlid species flock of Lake Tanganyika represents a polyphyletic assemblage of eight ancestral lineages, which colonized the emerging lake independently. Our study is focused on one of these lineages, the Bathybatini, a tribe of specialized piscivorous cichlids of the deep pelagic zone. By analyzing three mtDNA gene segments of all eight species of the tribe and two species of the closely related Trematocarini, we propose on the basis of a linearized tree analysis that the Bathybatini comprise two distinct lineages, the genera Hemibates and Bathybates, that seeded the primary lacustrine Tanganyika radiation independently. The genus Hemibates is likely to represent a distinct lineage that emerged simultaneously with the tribe Trematocarini and the genus Bathybates and should be therefore treated as a distinct tribe. Within the genus Bathybates, B. minor clearly represents the most ancestral split and is likely to have diverged from the remaining species in the course of the primary lacustrine Tanganyika radiation during which also the radiations of the Lamprologini and the H-lineage took place. The remaining large Bathybates species also diversified almost simultaneously and in step with the diversification of other Tanganyikan lineages—the Limnochromini and Cyprichromini—with B. graueri occupying the most ancestral branch, suggesting that these were induced by the same environmental changes. The lack of geographic color morphs suggests that competition and resource partitioning, rather than allopatric speciation, promoted speciation within the genus Bathybates.Reviewing Editor: Dr. Axel Meyer     

Phylogenetic relationships within the Archiloa genus complex (Proseriata,Monocelididae)

Of the seven genera which we have recognised within the Archiloa genus complex sensu Karling (1966) the cosmopolitan genus Archilina is the most primitive and is characterised only by plesiomorphic characters, and has to be considered paraphyletic. All other species of the Archiloa genus complex are hypothesized to be derived from Archilina-like ancestors through different evolutionary lineages. One lineage led to the genera Archiloa, Inaloa, Archilopsis and Monocelopsis, taxa found in the Atlantic and the Mediterranean. These genera are monophyletic and their relationships are analyzed. The genera Mesoda (Brazil) and Tajikina (Northern Pacific) can be considered as two other separate lineages. Similarly, within what we now consider as the genus Archilina different lineages can be recognized in different regions.     

Phylogenetic systematics of the colorful, cyanide-producing millipedes of Appalachia (Polydesmida, Xystodesmidae, Apheloriini) using a total evidence Bayesian approach

Here, we provide an exemplar-approach phylogeny of the xystodesmid millipede tribe Apheloriini with a focus on genus-group relationships-particularly of the genus Brachoria. Exemplars for the phylogenetic analysis were chosen to represent the maximum breadth of morphological diversity within all nominal genera in the tribe Apheloriini, and to broadly sample the genus Brachoria. In addition, three closely related tribes were used (Rhysodesmini, Nannariini, and Pachydesmini). Morphological and DNA sequence data were scored for Bayesian inference of phylogeny. Phylogenetic analysis resulted in polyphyletic genera Brachoria and Sigmoria, a monophyletic Apheloriini, and a "southern clade" that contains most of the tribal species diversity. We used this phylogeny to track morphological character histories and reconstruct ancestral states using stochastic character mapping. Based on the findings from the character mapping study, the diagnostic feature of the genus Brachoria, the cingulum, evolved independently in two lineages. We compared our phylogeny against prior classifications using Bayes factor hypothesis-testing and found that our phylogenetic hypothesis is inconsistent with the previous hypotheses underlying the most recent classification. With our preferred total-evidence phylogeny as a framework for taxonomic modifications, we describe a new genus, Appalachioria; supply phylogenetic diagnoses of monophyletic taxa; and provide a phylogeny-based classification for the tribe Apheloriini.     

Tribal assignment of <Emphasis Type="Italic">Heldreichia</Emphasis> Boiss. (Brassicaceae): evidence from nuclear ITS and plastidic <Emphasis Type="Italic">ndh</Emphasis>F markers

Heldreichia Boiss. is a monospecific genus mainly distributed in Anatolia and the Lebanon. Although morphological variation and infrageneric phylogenetic relationships were recently studied in detail, Heldreichia remained as one of the few orphan genera that have not yet been assigned to any tribe. In the current study, we used sequence data from the nuclear ITS and chloroplast ndhF regions of Heldreichia and representatives of main Brassicaceae lineages and tribes to determine its tribal affiliation. Bayesian-based phylogenetic analyses clearly show with high support that Heldreichia is a member of the recently expanded tribe Biscutelleae. Furthermore, we characterize the tribe Biscutelleae morphologically and provide a determination key for all its genera.     

Phylogeny of Mesoamerican freshwater mussels and a revised tribe‐level classification of the Ambleminae

Evolutionary and ecological hypotheses of the freshwater mussel subfamily Ambleminae are intensely geographically biased—a consequence of the complete exclusion of Mesoamerican taxa in phylogenetic reconstructions of the clade. We set out to integrate a portion of the Mesoamerican freshwater mussel assemblage into existing hypotheses of amblemine classification and evolution by generating a molecular phylogeny that includes four previously unsampled Mesoamerican genera and nine species endemic to that region. Given the traditionally hypothesized affinity to Nearctic mussels and the understanding that classification should reflect common ancestry, we predicted that (a) Mesoamerican genera would be recovered as members of the recognized tribes of the Ambleminae, and (b) genera would be supported as monophyletic. The mutilocus phylogeny (COI + 28S + 16S) reported herein does not fully support either of those hypotheses. Neither Cyrtonaias nor Psorula were supported as monophyletic and we predict several other Mesoamerica genera are also non‐monophyletic. The reconstructed phylogeny recovered four independent lineages of Mesoamerican freshwater mussels and these clades are distributed across the phylogeny of the Ambleminae, including the tribe Quadrulini (Megalonaias), Lampsilini (two lineages: Cyrtonaias explicata/Sphenonaias microdon, and Pachynaias), and a previously unrecognized, exclusively Mesoamerican and Rio Grande clade consisting of the genera Psoronaias, Psorula and Popenaias. The latter clade possesses several morphological characteristics that distinguish it from its sister taxon, tribe Lampsilini, and we recognize this newly identified Mesoamerican clade as a fifth tribe of the Ambleminae attributable to the Popenaiadini Heard & Guckert, 1970. This revised classification more completely recognizes the suprageneric diversity of the Ambleminae.     

Monophyly and relationships of the tribe Exaceae (Gentianaceae) inferred from nuclear ribosomal and chloroplast DNA sequences

Both chloroplast trnL (UAA) intron and nuclear ribosomal ITS sequences highly confirmed the monophyly of the tribes of the Gentianaceae defined by the recent classification, and revealed the tribe Exaceae as a basal clade just next to the basal-most lineage, the tribe Saccifolieae. Within the tribe Exaceae, Sebaea (except Sebaea madagascariensis) appeared as the most basal clade as the sister group to the rest of the tribe. The Madagascan endemic genera Gentianothamnus and Tachiadenus were very closely related to each other, together standing as sister to a clade comprising Sebaea madagascariensis, Ornichia, and Exacum. The saprophytic genus Cotylanthera nested deeply inside Exacum. Sebaea madagascariensis was shown closer to the Madagascan endemic genus Ornichia than to any other sampled Sebaea species. Exacum appeared as the most derived taxon within this tribe. The topology of the phylogenetic trees conform with the Gondwana vicariance hypothesis regarding the biogeography of Exaceae. However, no evidence for matching the older relationships within the family to the tectonic history could be corroborated with various divergence time analyses. Divergence dating estimated a post-Gondwana diverging of the Gentianaceae about 50 million years ago (MYA), and the tribe Exaceae as about 40 MYA. The Mozambique Channel land-bridge could have played an important role in the biogeographic history of the tribe Exaceae.     

Phylogeny and taxonomy of casebearer moths (Lepidoptera,Coleophoridae) based on morphological and molecular genetic data. 1. Reconstruction of phylogeny of coleophoridae using analysis of <Emphasis Type="Italic">COI</Emphasis> gene variability

A phylogenetic study of representatives of the family Coleophoridae was conducted using a comprehensive approach, including methods of morphological and molecular genetic analyses. The existent data on the family system were compared with the results of phylogenetic analysis of the COI mitochondrial gene sequences. Four of the five studied subfamilies (Coleophorinae, Ischnophaninae, Augasminae, and Tolleophorinae) corresponded to their location on the phylogram; representatives of Metriotinae were part of Coleophorinae. According to the aggregate data from molecular phylogeny and morphology, the most numerous subfamily of casebearers, Coleophorinae, is polyphyletic within its current boundaries. The results of our analysis of COI molecular divergence does not refute the monophyly of the tribes Casignetellini, Carpochenini, Klinzigedini, Goniodomini, Casasini, and Atractulini from the subfamily Coleophorinae. The allocation of the tribes Aporipturini and Sistrophoecini within the subfamily does not correspond to the molecular data. Monophyly of the genera Ecebalia, Perygra, and Casignetella was confirmed. These genera are well isolated, which reflects the evolutionary significance of the morphological characters chosen for their taxonomic division. The boundaries of the cluster containing these genera correspond to those of the tribe Casignetellini, justifying the allocation of this tribe within the subfamily. The existence of monophyletic tribes Goniodomini (genus Goniodoma) and Carpochenini (genera Ionescumia, Carpochena, and Falkmisa) was also supported. The exceptions were the genera Kasyfia, Tollsia, and Agapalsa, whose monophyly was not confirmed by our results. The distribution of the sequences of species of these genera indicated a paraphyletic origin of Kasyfia and Tollsia and a polyphyletic origin of Agapalsa.     

A comprehensive molecular phylogeny of tiger beetles (Coleoptera,Carabidae, Cicindelinae)

Tiger beetles are a remarkable group that captivates amateur entomologists, taxonomists and evolutionary biologists alike. This diverse clade of beetles comprises about 2300 currently described species found across the globe. Despite the charisma and scientific interest of this lineage, remarkably few studies have examined its phylogenetic relationships with large taxon sampling. Prior phylogenetic studies have focused on relationships within cicindeline tribes or genera, and none of the studies have included sufficient taxon sampling to conclusively examine broad species patterns across the entire subfamily. Studies that have attempted to reconstruct higher‐level relationships of Cicindelinae have yielded conflicting results. Here, we present the first taxonomically comprehensive molecular phylogeny of Cicindelinae to date, with the goal of creating a framework for future studies focusing on this important insect lineage. We utilized all available published molecular data, generating a final concatenated dataset including 328 cicindeline species, with molecular data sampled from six protein‐coding gene fragments and three ribosomal gene fragments. Our maximum‐likelihood phylogenetic inferences recover Cicindelinae as sister to the wrinkled bark beetles of the subfamily Rhysodinae. This new phylogenetic hypothesis for Cicindelinae contradicts our current understanding of tiger beetle phylogenetic relationships, with several tribes, subtribes and genera being inferred as paraphyletic. Most notably, the tribe Manticorini is recovered nested within Platychilini including the genera Amblycheila Say, Omus Eschscholtz, Picnochile Motschulsky and Platychile Macleay. The tribe Megacephalini is recovered as paraphyletic due to the placement of the monophyletic subtribe Oxycheilina as sister to Cicindelini, whereas the monophyletic Megacephalina is inferred as sister to Oxycheilina, Cicindelini and Collyridini. The tribe Collyridini is paraphyletic with the subtribes Collyridina and Tricondylina in one clade, and Ctenostomina in a second one. The tribe Cicindelini is recovered as monophyletic although several genera are inferred as para‐ or polyphyletic. Our results provide a novel phylogenetic framework to revise the classification of tiger beetles and to encourage the generation of focused molecular datasets that will permit investigation of the evolutionary history of this lineage through space and time.     

Molecular phylogeny of the mega‐diverse rove beetle tribe Staphylinini (Insecta,Coleoptera, Staphylinidae)

Chatzimanolis, S., Cohen, I. M., Schomann, A. & Solodovnikov, A. (2010). Molecular phylogeny of the mega‐diverse rove beetle tribe Staphylinini (Insecta, Coleoptera, Staphylinidae). —Zoologica Scripta, 39, 436–449. Phylogeny of the rove beetle tribe Staphylinini is explored by parsimony and Bayesian analyses of sequences of four genes (COI, wingless, Topoisomerase I, and 28S) for 43 ingroup (various genera of Staphylinini) and eight outgroup (two genera of Paederinae, six genera of other tribes of Staphylininae) taxa. Analyses were conducted for each gene independently and for the concatenated data set. Results of the most robust combined analyses were compared with the morphology‐based phylogenies of Staphylinini (‘test phylogeny’), and with the conventional classification of this tribe. Molecular results were congruent with the ‘test phylogeny’ in the following: ancestors of Staphylinini were ‘Quediina‐like’ lineages; formal subtribe Quediina mixes at least two relatively basal groups, ‘Quediina propria’ and ‘southern Quediina’; specialized subtribe Amblyopinina is an internal clade within ‘southern Quediina’; a relatively deeply nested ‘Staphylinini propria’ that unites current subtribes Staphylinina, Eucibdelina, Anisolinina, Xanthopygina and Philonthina is well supported as a monophyletic group. In strong contrast with morphology, molecular data place the tribes Othiini and Xantholinini nested within Staphylinini. Molecular results strongly conflict with morphology by uniting morphologically very different genera Holisus and Atanygnathus in one clade that has uncertain position within Staphylinini. Consistently with the most congruent areas of the morphology‐ and molecular‐based phylogenies, taxonomic changes are implemented for the formal subtribes Quediina and Amblyopinina.     

A molecular phylogeny of the tribe Aphidini (Insecta: Hemiptera: Aphididae) based on the mitochondrial tRNA/COII, 12S/16S and the nuclear EF1α genes

Abstract A phylogeny of the tribe Aphidini (Hemiptera: Aphididae) was reconstructed from three gene fragments: two mitochondrial regions, partial tRNA‐leucine + cytochrome oxidase II (tRNA/COII), partial 12S rRNA + tRNA‐valine + 16S rRNA (12S/16S) and one nuclear gene, the elongation factor‐1 alpha (EF1α). Bayesian phylogenetic (BP) analyses were performed on each individual dataset of tRNA/COII, 12S/16S and EF1α, and maximum parsimony (MP), Bremer support test, maximum likelihood (ML) and BP analysis were performed on the combined dataset. After comparing our molecular phylogenetic results with the classic classification based on morphological and ecological data, we analysed three main issues: the monophyletic relationships among tribes and subtribes, the validities of the latest taxonomic positions of genera and species and the status of certain Aphis species groups. Our results indicate that 36 of the species analysed, with the exception of Cryptosiphum artemisiae, are clustered within the clade of Aphidini. Also, the 28 species representative of the subtribe Aphidina were separated from the eight species representative of Rhopalosiphina; each monophyletic subtribe was supported by significant P‐values in the combined analysis. According to our results, Cryptosiphum should be moved to Macrosiphini because it is more closely related to the genera Lipaphis and Brevicoryne. The genus Toxoptera was recovered as non‐monophyletic. In Rhopalosiphina, three genera, Hyalopterus, Rhopalosiphum and Schizaphis, were relatively closer to each other than to the genus Melanaphis. In the relationships between species‐groups among Aphis, most species were separated into two main lineages; the fabae group seemed to be more closely related to the spiraecola and craccivora group rather than to the gossypii group.     

The cladistics and higher classification of the Pimpliformes (Hymenoptera: Ichneumonidae)

The monophyly of the ichneumonid clade Pimpliformes is established and the phylogenetic relationships of the eight component subfamilies are resolved. The clade (Acaenitinae + (Diacritinae + (Cylloceriinae + (Diplazontinae + Orthocentrinae)))) is the sister-lineage to the clade (Pimplinae + (Rhyssinae + Poemeniinae)). The Nearctic genus Cressonia Dasch is transferred to the Diacritinae from the Orthocentrinae. Tribes are not recognized in the Acaenitinae as the Coleocentrini (sensu Townes, 1971) is paraphyletic with respect to the Acaenitini. The Cylloceriinae is recognized as comprising three genera, Cylloceria Schiødte, Allomacrus Förster and Sweaterella gen.n. The Orthocentrinae, including the Helictinae of authors, is shown to be monophyletic, but the latter is clearly shown to be paraphyletic if the Orthocentrus genus-group is excluded. The Pimplinae comprises four monophyletic tribes: the Delomeristini, consisting of Delomerista Förster and Atractogaster Kriechbaumer; the Perithoini trib.n., which includes only Perithous Holmgren (= Hybomischos Baltazar syn.n.); the Pimplini, which includes the Theronia genus-group as well as the Pimpla genus-group; and the Ephialtini, which includes the Polysphinctini syn.n., a monophyletic group that previously rendered the restricted Ephialtini paraphyletic. The tribe Delomeristini is the sister-group to the clade (Ephialtini + (Perithoini + Pimplini)). The subfamily Poemeniinae is recognized as comprising three tribes: the Pseudorhyssini (trib.n.) which includes the single Holarctic genus Pseudorhyssa Merrill; the Rodrigamini (trib.n.) which includes only the Costa Rican genus Rodrigama Gauld; and the Poemeniini. The tribe Pseudorhyssini is the sister-group to the clade (Rodrigamini + Poemeniini). The phylogenetic inter-relationships of the genera of Poemeniini are resolved. A new genus from South Africa, Guptella (gen.n.) is described, and Achorocephalus Kriechbaumer is shown to be a synonym of Eugalta Cameron (syn.n.). The evolution of biological traits within the Pimpliformes is discussed with reference to the elucidated phylogeny, and zoogeographic patterns are outlined.     

Multiple origin of flightlessness in Phaneropterinae bushcrickets and redefinition of the tribus Odonturini (Orthoptera: Tettigonioidea: Phaneropteridae)

The possession of wings and ability to fly are a unifying character of higher insects, but secondary loss of wings is widespread. Within the bushcrickets, the subfamily Phaneropterinae (Orthoptera: Tettigonioidea) comprises more than 2000 predominantly long-winged species in the tropics. However, the roughly 300 European representatives are mainly short-winged. The systematics of these radiations have been unclear, leading to their unreliable formal treatment, which has hindered analysis of the evolutionary patterns of flight loss. A molecular phylogeny is presented for 42 short-winged species and members of all European long-winged genera based on the combined data from three nuclear gene sequences (18S, H3, ITS2). We found four phylogenetic lineages: (i) the first included the short-wing species of the genus Odontura; (ii) a further branch is represented by the South-American short-winged Cohnia andeana; (iii) an assemblage of long-wing taxa with a deep branching pattern includes the members of the tribes Acrometopini, Ducetiini, Phaneropterini, and Tylopsidini; (iv) a large group contained all short-winged taxa of the tribe Barbitistini. Phaneropterinae flightlessness originated twice in the Western Palaearctic, with a number of mainly allo- and parapatrically distributed species of the Barbistini in Southeastern Europe, and the Middle East and a limited number of Odontura species in Northern Africa and Southwestern Europe. Both short-winged lineages are well separated, which makes it necessary to restrict the tribe Odonturini to the West-Palaearctic genus Odontura. Other flightless genera previously included in the Odonturini are placed as incertae sedis until their phylogenetic position can be established.     

Phylogeny, character evolution, and classification of Sapotaceae (Ericales)

We present the first cladistic study of the largely tropical family Sapotaceae based on both morphological and molecular data. The data were analyzed with standard parsimony and parsimony jackknife algorithms using equally and successive weighted characters. Sapotaceae are confirmed to constitute two main evolutionary lineages corresponding to the tribes Isonandreae‐Mimusopeae‐Sideroxyleae and Chrysophylleae‐Omphalocarpeae. The Sideroxyleae are monophyletic, Isonandreae are polyphyletic as presently circumscribed, and as suggested by the analyses, the subtribe Mimusopeae‐Mimusopinae has evolved within the Mimusopeae‐Manilkarinae, which hence is also paraphyletic. Generic limits must be altered within Sideroxyleae with the current members Argania, Nesoluma and Sideroxylon. Argania cannot be maintained at a generic level unless a narrower generic concept is adopted for Sideroxylon. Nesoluma cannot be upheld in a narrow or broad generic concept of Sideroxylon. The large tribe Chrysophylleae circumscribes genera such as Chrysophyllum, Pouteria, Synsepalum, and Xantolis, but the tribe is monophyletic only if the taxa from Omphalocarpeae are also included. Neither Chrysophyllum nor Pouteria are monophyletic in their current definitions. The results indicate that the African taxa of Pouteria are monophyletic and distinguishable from the South American taxa. Resurrection of Planchonella, corresponding to Pouteria section Oligotheca, is proposed. The African genera Synsepalum and Englerophytum form a monophyletic group, but their generic limits are uncertain. Classification of the Asian genus Xantolis is particularly interesting. Morphology alone is indecisive regarding Xantolis relationships, the combined unweighted data of molecules and morphology indicates a sister position to Isonandreae‐Mimusopeae‐Sideroxyleae, whereas molecular data alone, as well as successive weighted combined data point to a sister position to Chrysophylleae‐Omphalocarpeae. An amended subfamily classification is proposed corresponding to the monophyletic groups: Sarcospermatoideae (Sarcosperma), Sapotoideae (Isonandreae‐Mimusopeae‐Sideroxyleae) and Chrysophylloideae (Chrysophylleae‐Omphalocarpeae), where Sapotoideae circumscribes the tribes Sapoteae and Sideroxyleae as well as two or three as yet unnamed lineages. Morphological characters are often highly homoplasious and unambiguous synapomorphies cannot be identified for subfamilies or tribes, which we believe are the reason for the variations seen between different classifications of Sapotaceae. © The Willi Hennig Society 2005.     

PHYLOGENETIC RELATIONSHIPS AMONG LINEAGES OF THE CERAMIACEAE (CERAMIALES,RHODOPHYTA) BASED ON NUCLEAR SMALL SUBUNIT rDNA SEQUENCE DATA1

Phylogenetic relationships among 69 species of the Ceramiales (51 Ceramiaceae, six Dasyaceae, seven Delesseriaceae, and five Rhodomelaceae) were determined based on nuclear SSU rDNA sequence data. We resolved five strongly supported but divergent lineages among the included Ceramiaceae: (i) the genus Inkyuleea, which weakly joins other orders of the Rhodymeniophycidae rather than the Ceramiales in our analyses; (ii) the tribe Spyridieae, which is sister to the remainder of the included ceramialean taxa; (iii) the subfamily Ceramioideae, weakly including the tribe Warrenieae; (iv) the subfamily Callithamnioideae; and (v) the subfamily Compsothamnioideae, which emerges as sister to the Dasyaceae/Delesseriaceae/Rhodomelaceae complex, thus rendering the Ceramiaceae sensu lato unequivocally paraphyletic, as has been argued separately on anatomical grounds by Kylin and Hommersand. Our data support a restricted concept of the Ceramiaceae that includes only one of the five lineages (Ceramioideae) that we have resolved. In addition to failing to ally with the Ceramiales in our molecular analyses, species of Inkyuleea differ substantially from other Ceramiaceae sensu lato in details of pre‐ and postfertilization development. The genus Inkyuleea is here assigned to the Inkyuleeaceae fam. nov., which we provisionally retain in the Ceramiales. Species of Spyridia also differ from the remaining Ceramiaceae in their postfertilization development, and, in light of our molecular data, the genus Spyridia is assigned to the Spyridiaceae. The Callithamnioideae is strongly monophyletic (100% in all analyses), which, in combination with key anatomical differences, supports elevation to family status for this lineage as the Callithamniaceae. Similarly, the Compsothamnioideae is solidly monophyletic in our molecular trees and has a unique suite of defining anatomical characters that supports family status for a complex that we consider to include the tribes Compsothamnieae, Dasyphileae, Griffithsieae, Monosporeae, Ptiloteae, Spermothamnieae, Sphondylothamnieae, Spongoclonieae, and Wrangelieae, for which the reinstated family name Wrangeliaceae is available.     

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.     

Revised systematics and higher classification of pierid butterflies (Lepidoptera: Pieridae) based on molecular data

The butterfly family Pieridae comprises approximately 1000 described species placed in 85 genera, but the higher classification has not yet been settled. We used molecular data from eight gene regions (one mitochondrial and seven nuclear protein‐coding genes) comprising a total of ~6700 bp from 96 taxa to infer a well‐supported phylogenetic hypothesis for the family. Based on this hypothesis, we revise the higher classification for all pierid genera. We resurrect the tribe Teracolini stat. rev. in the subfamily Pierinae to include the genera Teracolus, Pinacopteryx, Gideona, Ixias, Eronia, Colotis and most likely Calopieris. We transfer Hebomoia to the tribe Anthocharidini and assign the previously unplaced genera Belenois and Dixeia to the subtribe Aporiina. Three lineages near the base of Pierinae (Leptosia, Elodina and Nepheronia + Pareronia) remain unplaced. For each of these, we describe and delineate new tribes: Elodinini Braby tribus nova, Leptosiaini Braby tribus nova and Nepheroniini Braby tribus nova. The proposed higher classification is based on well‐supported monophyletic groups and is likely to remain stable even with the addition of more data.     

基于28S rDNA D2基因片段与形态特征的矛茧蜂亚科系统发育研究（膜翅目：茧蜂科）

本研究选取矛茧蜂亚科Doryctinae（昆虫纲Insecta：膜翅目Hymenoptera：茧蜂科Braconidae）的6族15属18种做内群,茧蜂科其它7亚科11属11种做外群,首次结合同源核糖体28S rDNA D2基因序列片段和100个形态学和解剖学特征对该亚科进行了系统发育学研究。利用“非圆口类"的小腹茧蜂亚科Microgastrinae为根,以PAUP*4.0和MrBayes 3.0B4软件分别应用最大简约法（MP）和贝叶斯法对矛茧蜂亚科的分子数据和分子数据与非分子数据的结合体进行了运算分析;并以PAUP*4.0对矛茧蜂亚科的28S rDNA D2基因序列片段的碱基组成与碱基替代情况进行了分析。结果表明：矛茧蜂亚科的28S rDNA D2基因序列片段的GC含量在39.33%～48.28%之间变动,而对于碱基替代情况来讲,矛茧蜂亚科各成员间序列变异位点上颠换（transversion）大于转换（transition）。不同的分析算法所产生的系统发育树都表明矛茧蜂亚科是一个界限分明的单系群;在矛茧蜂亚科内,除了吉丁茧蜂族Siragrini为单系群外,其他族（矛茧蜂族Doryctini和方头茧蜂族Hecabolini）都是并系群。对于矛茧蜂亚科内各属之间的相互亲缘关系,不同算法所得的系统发育树的拓扑结构不完全一致,表明矛茧蜂亚科内（属及族）的系统发育关系还有待于进一步研究。     

The tropical African legume Scorodophloeus clade includes two undescribed Hymenostegia segregate genera and Micklethwaitia,a rare,monospecific genus from Mozambique

Legume subfamily Caesalpinioideae accommodates approximately 2250 species in 171 genera which traditionally are placed in four tribes: Caesalpinieae, Cassieae, Cercideae and Detarieae. The monophyletic tribe Detarieae includes the Amherstieae subclade which contains about 55 genera. Our knowledge of the relationships among those genera is good in some cases but for many other genera phylogenetic relationships have been unclear. The non-monophyletic nature of at least two amherstioid genera, Cynometra and Hymenostegia has also complicated the picture. During the course of a multi-disciplinary study of Hymenostegia sensu lato, which includes phylogenetic analyses based on matK and trnL data, we have recovered the “Scorodophloeus clade”, an exclusively tropical African clade of four genera which includes the eponymous genus Scorodophloeus, two undescribed generic segregates of Hymenostegia sensu lato, and the previously unsampled rare monospecific genus Micklethwaitia from Mozambique. Zenkerella is suggested as a possible sister genus to the Scorodophloeus clade. A distribution map is presented of the seven species that belong to the Scorodophloeus clade.     

Duthieeae,a new tribe of grasses (Poaceae) identified among the early diverging lineages of subfamily Pooideae: molecular phylogenetics,morphological delineation,cytogenetics and biogeography

The phylogeny of Pooideae, one of the largest subfamilies of grasses, has been intensively studied during the past years. To investigate the early evolutionary splits in Pooideae we used a broad sample of genera with uncertain placement, some of which have not been studied in molecular phylogenetics before, complemented by representatives from other lineages of this subfamily. Morphological, cytogenetic and biogeographical analyses were added to the molecular sequence work on chloroplast matK–3’trnK and nuclear ITS. According to chloroplast DNA data, a new and well-supported lineage was identified among the early branches. It consisted of Phaenosperma and a larger group of genera encompassing Anisopogon, Danthoniastrum, Duthiea, Metcalfia, Pseudodanthonia (inclusion resting on ITS and morphology), Sinochasea and Stephanachne. Based on structural characters we suggest to keep Phaenosperma under the monotypic tribe Phaenospermateae and to accommodate the other genera under a new tribe Duthieeae, which is morphologically well-defined by synapomorphic spikelet features. Megalachne and Podophorus were not part of the early diverging Pooideae lineages but belong to the Aveneae/Poeae complex. Morphological characteristics of Duthieeae are discussed with respect especially to Stipeae and reveal consistent differences between both tribes. The genera of Duthieeae and the major lineages of Stipeae are keyed. A cytogenetic survey of exemplary taxa corroborates high chromosome base numbers as prevailing within the early diverging lineages of Pooideae, but chromosome sizes are more highly varied than previously reported. Ecogeographical analyses point to warm and humid conditions as the ancestral bioclimatic niche of Phaenosperma and Duthieeae, whereas adaptation to cold and drought occurred only in a part of Duthieeae but was obviously less successful than in the widespread and much more species-rich tribe Stipeae. The distribution of Duthieeae with species-poor or monotypic genera in mountains of the northern hemisphere and Anisopogon as an outlier in Australia suggests relict character.     

A molecular view of pinniped relationships with particular emphasis on the true seals

Phylogenetic analysis of conservative nucleotide substitutions in 18 complete sequences of the mitochondrial cytochrome gene of Phocidae (true seals), Odobenidae (walruses), and Otariidae (sea lions and fur seals), plus three ursid and three felid sequences, identified the pinnipeds as monophyletic with Otariidae and Odobenidae on a common evolutionary branch. Analysis of total nucleotide differences separated the evolutionary lineages of northern and southern phocids. Both lineages are distinct from the most ancestral phocid genus, Monachus (monk seals), represented by the Hawaiian monk seal. The inclusion of the Hawaiian monk seal in the subfamily Monachinae makes the subfamily paraphyletic. Among the northern phocids, the hooded seal (genus Cystophora, chromosome number 2n = 34) is sister taxon to the Phoca complex. The Phoca complex, which is characterized by the chromosome number 2n = 32, includes genus Phoca and the monotypic genus Halichoerus (grey seal). The comparison does not support a generic distinction of Halichoerus within the Phoca complex. The present data suggest that Cystophora and Phoca separated 6 million years ago. Among the southern phocids the close molecular relationship of the Weddell and leopard seals relative to their morphological distinction exemplifies rapid adaptation to different ecological niches. This result stands in contrast to the limited morphological differentiation relative to the pronounced molecular distinctions that may occur within the Phoca complex.Correspondence to: Ú. Árnason