Breaking family ties: taxon sampling and molecular phylogeny of chromodorid nudibranchs (Mollusca,Gastropoda)

Johnson, R. F. (2010). Breaking family ties: taxon sampling and molecular phylogeny of chromodorid nudibranchs (Mollusca, Gastropoda). —Zoologica Scripta, 40, 137–157. Although researchers have debated the monophyly of the diverse chromodorid nudibranchs (Chromodorididae) for over 100 years, the monophyly of this family has not been properly tested. Recent morphological and molecular phylogenetic studies have added to the debate, but have not used appropriate methods to resolve this issue. I investigate how outgroup choice and taxon sampling influences tree topology and in turn the recovery of chromodorid monophyly. As a demonstration of these potential methodological problems, I then present phylogenies resulting from different taxon‐sampling schemes using the same molecular data. Taxon sampling has a strong influence on the resulting phylogenies. With comprehensive taxon sampling and outgroup selection, Cadlina is not a member of the Chromodorididae. The chromodorid nudibranchs without Cadlina are monophyletic and possibly sister to the Actinocyclidae. Additionally, I found, for the first time, support for most current family groupings in the Doridoidea. I propose a new classification in which Cadlina is not considered a member of the Chromodorididae. Instead, I resurrect the family name Cadlinidae to include the genera Cadlina and Aldisa.     

Basal chromodorid sperm ultrastructure (Nudibranchia, Gastropoda, Mollusca)

The relationship between three genera considered basal in the Chromodorididae (Cadlina, Tyrinna, Cadlinella) has not yet been resolved by traditional morphological means. Here we examined the sperm ultrastructure of Tyrinna nobilis, Tyrinna evelinae, Cadlina flavomaculata and Cadlina cf. nigrobranchiata, with the expectation of finding phylogenetically informative characters. No Tyrinna or Cadlina species showed sperm similarities to Cadlinella. Both Cadlina species and Tyrinna nobilis (but not T. evelinae) exhibited coarse striations in the acrosomal pedestal. The putative fibers that occurred between the coarse striations of the pedestal are condensed into a layer in Cadlina and Tyrinna, but not in other species that also have coarse striations (Gymnodoris), and may constitute evidence for a close relationship. Tyrinna evelinae possessed fine acrosomal striations, which was shared with other Chromodorididae, Actinocyclidae and the cryptobranchs Rostanga and Aphelodoris. We also examined the sperm ultrastructure of ‘Chromodoris’ ambiguus, an animal which has shown molecular affinities to species of Cadlina, and not Chromodoris. The sperm of ‘C.’ ambiguus did not exhibit the typical Cadlina characteristics, but also showed important differences to other investigated Chromodoris species.     

Can the morphology of the integumentary spicules be used to distinguish genera and species of phyllidiid nudibranchs (Porostomata: Phyllidiidae)?

Thirty-eight specimens belonging to four genera and 15 species of the nudibranch family Phyllidiidae were examined to investigate whether the morphology of their integumentary calcareous spicules and/or the occurrence of the spicules within the regions of the body could be used to distinguish genera and species. The spicules were studied separately from five regions of the body of each specimen—the foot, gills, mantle, dorsal pustules (or ridges in Reticulidia) and rhinophores. The mantle itself plus its pustules were found to possess the full complement of spicules in every individual. Four types of spicules were recorded overall—smooth diactines, centro-polytylote diactines, triactines and tetractines. Different regions of the body were found to possess different spicule types: (a) only smooth diactines in the gills, (b) both smooth diactines and triactines in the foot and (c) all of smooth diactines, centro-polytylote diactines and triactines in the mantle, dorsal pustules and the rhinophores. Among the genera, three types of spicules (smooth diactine, triactine, and tetractine) are present in Phyllidia, Phyllidiopsis and Reticulidia, but the form of the spicules is not diagnostic between these genera or between the constituent species. The fourth type of spicule (centro-polytylote diactine) is present exclusively in Phyllidiella, and is diagnostic for that genus. However, we failed to find any difference in spicule form, or composition, or location in the body between the three (closely related) species of Phyllidiella we investigated. Therefore, our key conclusion is that spicule morphology is an extremely important character to tell the genus Phyllidiella apart from all the other genera of the family, but it is not taxonomically informative at the level of species.     

石山苣苔属(苦苣苔科)花形态演化及分类学意义

石山苣苔属(苦苣苔科)约30种,主要分布于我国南部的石灰岩地区.目前该属已知物种数虽少但花形态极其多样,是该科中分类最为困难的类群之一.基于分子证据,其它8个属中花形态迥异的一些物种被并入石山苣苔属.然而,该属花形态的演化趋势缺乏系统性的研究,传统分类对属的界定与分子系统学研究结果相矛盾的原因,以及是否有形态特征支持新界定的石山苣苔属还不清楚.该研究中,总共编码了19种石山苣苔属植物和9种报春苣苔属植物的35个形态特征,其中包括26个花部形态特征,在分子系统树上追踪了它们的演化路径.结果表明:无论属内还是属间,多数花部形态特征,尤其以往属的分类界定特征,在演化过程中变化频繁且发生了高度同塑性演化,这是导致传统形态分类不自然的关键因素.此外,在观察研究的所有特征中,花丝和柱头的差异可能在石山苣苔属植物共同祖先中经历了演变,或可用于区分石山苣苔属与其姐妹报春苣苔属的大多数种类.因此,在苦苣苔科植物的分类学研究中应当慎用这些花部性状作为分类依据,而且应对形态特征进行广泛地观察研究,在密集的取样和分辨率更高、更可靠的系统树上追踪它们的演化规律.更为重要的是,需要进一步研究导致复杂形态性状演化的内在分子调控机理和外在的自然选择动力,最终更加深入地理解石山苣苔属等典型喀斯特植物的演化过程和机理.     

Taxonomic classification of the reef coral families Merulinidae,Montastraeidae, and Diploastraeidae (Cnidaria: Anthozoa: Scleractinia)

Modern coral taxonomy has begun to resolve many long‐standing problems in traditional systematics stemming from its reliance on skeletal macromorphology. By integrating examinations of colony, corallite, and subcorallite morphology with the molecular sequence data that have proliferated in the last decade, many taxa spread across the scleractinian tree of life have been incorporated into a rigorous classification underpinned by greater phylogenetic understanding. This monograph focuses on one of the most challenging clades recovered to date – its disarray epitomized by the informal name ‘Bigmessidae’. This group of predominantly Indo‐Pacific species previously comprised families Merulinidae, Faviidae, Pectiniidae, and Trachyphylliidae, but in a recent study these have been incorporated within Merulinidae. We studied 84 living merulinid species by examining morphological traits at three different scales of coral skeletal structure ? macromorphology, micromorphology, and microstructure ? to construct a morphological matrix comprising 44 characters. Data were analysed via maximum parsimony and also transformed onto a robust molecular phylogeny under the parsimony and maximum likelihood criteria. Comparisons amongst morphological character types suggest that although many characters at every scale are homoplastic, some to a greater extent than others, several can aid in distinguishing genus‐level clades. Our resulting trees and character analyses form the basis of a revised classification that spans a total of 139 species contained within 24 genera. The tree topologies necessitate the synonymization of Barabattoia as Dipsastraea, and Phymastrea as Favites. Furthermore, Astrea and Coelastrea are resurrected, and one new genus, P aramontastraea Huang & Budd gen. nov. , is described. All the genera in Merulinidae, along with the monotypic Montastraeidae and Diploastraeidae, are diagnosed based on the characters examined. The integrative classification system proposed here will form the framework for more accurate biodiversity estimates and guide the taxonomic placement of extinct species. © 2014 The Linnean Society of London     

Taxonomic classification of the reef coral family Lobophylliidae (Cnidaria: Anthozoa: Scleractinia)

Lobophylliidae is a family‐level clade of corals within the ‘robust’ lineage of Scleractinia. It comprises species traditionally classified as Indo‐Pacific ‘mussids’, ‘faviids’, and ‘pectiniids’. Following detailed revisions of the closely related families Merulinidae, Mussidae, Montastraeidae, and Diploastraeidae, this monograph focuses on the taxonomy of Lobophylliidae. Specifically, we studied 44 of a total of 54 living lobophylliid species from all 11 genera based on an integrative analysis of colony, corallite, and subcorallite morphology with molecular sequence data. By examining coral skeletal features at three distinct levels – macromorphology, micromorphology, and microstructure – we built a morphological matrix comprising 46 characters. Data were analysed via maximum parsimony and transformed onto a robust molecular phylogeny inferred using two nuclear (histone H3 and internal transcribed spacers) and one mitochondrial (cytochrome c oxidase subunit I) DNA loci. The results suggest that micromorphological characters exhibit the lowest level of homoplasy within Lobophylliidae. Molecular and morphological trees show that Symphyllia, Parascolymia, and Australomussa should be considered junior synonyms of Lobophyllia, whereas Lobophyllia pachysepta needs to be transferred to Acanthastrea. Our analyses also lend strong support to recent revisions of Acanthastrea, which has been reorganized into five separate genera (Lobophyllia, Acanthastrea, Homophyllia, Sclerophyllia, and Micromussa), and to the establishment of Australophyllia. Cynarina and the monotypic Moseleya remain unchanged, and there are insufficient data to redefine Oxypora, Echinophyllia, and Echinomorpha. Finally, all lobophylliid genera are diagnosed under the phylogenetic classification system proposed here, which will facilitate the placement of extinct taxa on the scleractinian tree of life.     

A molecular phylogeny of marine amphipods in the herbivorous family Ampithoidae

Ampithoid amphipods dominate invertebrate assemblages associated with shallow‐water macroalgae and seagrasses worldwide and represent the most species‐rich family of herbivorous amphipod known. To generate the first molecular phylogeny of this family, we sequenced 35 species from 10 genera at two mitochondrial genes [the cytochrome c oxidase subunit I (COI) and the large subunit of 16 s (LSU)] and two nuclear loci [sodium–potassium ATPase (NAK) and elongation factor 1‐alpha (EF1)], for a total of 1453 base pairs. All 10 genera are embedded within an apparently monophyletic Ampithoidae (Amphitholina, Ampithoe, Biancolina, Cymadusa, Exampithoe, Paragrubia, Peramphithoe, Pleonexes, Plumithoe, Pseudoamphithoides and Sunamphitoe). Biancolina was previously placed within its own superfamily in another suborder. Within the family, single‐locus trees were generally poor at resolving relationships among genera. Combined‐locus trees were better at resolving deeper nodes, but complete resolution will require greater taxon sampling of ampithoids and closely related outgroup species, and more molecular characters. Despite these difficulties, our data generally support the monophyly of Ampithoidae, novel evolutionary relationships among genera, several currently accepted genera that will require revisions via alpha taxonomy and the presence of cryptic species.     

An assessment of the status of Polycirridae genera (Annelida: Terebelliformia) and evolutionary transformation series of characters within the family

A phylogenetic analysis was performed to determine the monophyly of non‐monotypic genera of the terebelliform family Polycirridae, i.e. Polycirrus, Amaeana, Lysilla, and Hauchiella, and the evolution of characters among members of this clade. The monotypic genera, Enoplobranchus and Biremis, were also included, together with members of both known species in Hauchiella. Representative species were included for remaining genera: 14 species of Polycirrus, six species of Amaeana, and six species of Lysilla. Out‐groups consisted of representatives of Spionidae, Cirratulidae, and Sabellariidae, as well as several species of Telothelepodidae. A total of 40 in‐ and out‐group species were coded for 50 subjects (‘characters’) and 117 subject–predicate relationships (‘states’). Although results are consistent with recent phylogenetic studies within Terebelliformia that suggest Polycirridae monophyly, only Hauchiella was found to be monophyletic, albeit part of the more inclusive clade comprising remaining polycirrid genera. Evolutionary transformation series are discussed for selected characters in relation to the non‐monophyly of Polycirrus, Lysilla, and Amaeana. Implications for the use of supraspecific taxa as ‘taxonomic surrogates’ are highlighted. The definition of Polycirridae is emended. © 2015 The Linnean Society of London     

The Beothukis/Culmofrons problem and its bearing on Ediacaran macrofossil taxonomy: evidence from an exceptional new fossil locality

The late Ediacaran siliciclastic successions of eastern Newfoundland, Canada, are renowned for their fossils of soft‐bodied macro‐organisms, which may include some of the earliest animals. Despite the potential importance of such fossils for evolutionary understanding, the taxonomic framework within which Ediacaran macrofossils are described is not clearly defined. Rangeomorphs from a newly discovered fossil surface on the Bonavista Peninsula, Newfoundland, require us to reconsider contemporary use of morphological characters to distinguish between genera and species within Ediacaran taxa. The new surface exhibits remarkable preservational fidelity, resolving features smaller than 0.1 mm in dimension in both frondose and non‐frondose taxa. Such preservation permits the recognition of rarely observed fourth‐ and fifth‐order rangeomorph branching, offering unparalleled opportunities to investigate the fine‐scale construction of rangeomorph taxa including Culmofrons plumosa Laflamme et al., 2012. Our observations enable resolution of taxonomic issues relating to rangeomorphs, specifically overlap between the diagnoses of the frondose genera Beothukis Brasier and Antcliffe, 2009 and Culmofrons. We propose a taxonomic framework for all Ediacaran macrofossils whereby gross architecture, the presence/absence of discrete morphological characters and consideration of growth programme are used to distinguish genera, whereas morphometric or continuous characters define taxa at the species level. On the basis of its morphological characters, Culmofrons plumosa is herein synonymized to a species (Beothukis plumosa comb. nov.) within the genus Beothukis. This discussion emphasizes the need to standardize the taxonomic approach used to describe Ediacaran macrofossil taxa at both the genus and species levels, and raises important considerations for future formulation of higher‐level taxonomic groups.     

Phylogenetic relationships of Microdontinae (Diptera: Syrphidae) based on molecular and morphological characters

The intrasubfamilial classification of Microdontinae Rondani (Diptera: Syrphidae) has been a challenge: until recently more than 300 out of more than 400 valid species names were classified in Microdon Meigen. We present phylogenetic analyses of molecular and morphological characters (both separate and combined) of Microdontinae. The morphological dataset contains 174 characters, scored for 189 taxa (9 outgroup), representing all 43 presently recognized genera and several subgenera and species groups. The molecular dataset, representing 90 ingroup species of 28 genera, comprises sequences of five partitions in total from the mitochondrial gene COI and the nuclear ribosomal genes 18S and 28S. We test the sister‐group relationship of Spheginobaccha with the other Microdontinae, attempt to elucidate phylogenetic relationships within the Microdontinae and discuss uncertainties in the classification of Microdontinae. Trees based on molecular characters alone are poorly resolved, but combined data are better resolved. Support for many deeper nodes is low, and placement of such nodes differs between parsimony and Bayesian analyses. However, Spheginobaccha is recovered as highly supported sister group in both. Both analyses agree on the early branching of Mixogaster, Schizoceratomyia, Afromicrodon and Paramicrodon. The taxonomical rank in relation to the other Syrphidae is discussed briefly. An additional analysis based on morphological characters only, including all 189 taxa, used implied weighting. A range of weighting strengths (k‐values) is applied, chosen such that values of character fit of the resulting trees are divided into regular intervals. Results of this analysis are used for discussing the phylogenetic relationships of genera unrepresented in the molecular dataset.     

Phylogeny of the tribe Abrotrichini (Cricetidae,Sigmodontinae): integrating morphological and molecular evidence into a new classification

The tribe Abrotrichini (five genera and 14 living species) is a small clade within the speciose subfamily Sigmodontinae (Rodentia, Cricetidae), representing one of the extant successful radiations of mammals at southern high latitudes of the Neotropics. Its distribution is mostly Andean, reaching its greatest diversity in southern Argentina and Chile. We evaluate the phylogenetic relationships within this tribe through parsimony and Bayesian approaches based on 99 morphological characters (including 19 integumental characters, 38 skull characters, 31 dental characters, three postcranial skeletal characters, seven from the male accessory glands and phallus and one from the digestive system) and six molecular markers (one mitochondrial and five nuclear). We include representatives of all, except one, of the currently recognized species of living Abrotrichini plus one fossil form. Based on total evidence, we recovered a primary division between the genus Abrothrix and a group including the long‐clawed Abrotrichini, Chelemys, Geoxus, Notiomys and Pearsonomys. Both clades are recognized and named here as subtribes. The large degree of morphological variation observed within Abrothrix suggests that species in the genus fall into four groups, which we recognize as subgenera. In addition, the two known species of Chelemys do not form a monophyletic group, and Geoxus was recovered as paraphyletic with respect to Pearsonomys. To reconcile classification and phylogenetics, we describe a new genus for Chelemys macronyx and include Pearsonomys as a junior synonym of Geoxus. Our results highlight the importance of both morphology and molecules in resolving the phylogenetic relationships within this tribe. Based on biogeographical analyses, we hypothesize that Abrotrichini originated in south‐western South America by vicariance and then diversified mostly by successive dispersal events.     

Genera and species of Baetidae in Japan: <Emphasis Type="Italic">Nigrobaetis,Alainites, Labiobaetis</Emphasis>, and <Emphasis Type="Italic">Tenuibaetis</Emphasis> n. stat. (Ephemeroptera)

We describe morphological characters of the genera Nigrobaetis, Alainites, Labiobaetis, and Tenuibaetis n. stat. and provide generic situations of six Japanese species: Nigrobaetis chocoratus n. comb., N. sacishimensis n. comb., Alainites atagonis, A. florens, A. yoshinensis, and Tenuibaetis pseudofrequentus. To evaluate the polarities of the morphological characters and the monophyly of Nigrobaetis, Alainites, Labiobaetis, and Tenuibaetis, character states of these four genera were compared with the genus Cloeon as an outgroup. Labiobaetis is considered to be a monophyletic group supported by a wide paraglossa. Tenuibaetis is a monophyletic group that is distinguishable from the related genera by robust setae with a medial ridge on the dorsomedian surface of the nymphal femur. We did not find any synapomorphic characters of Nigrobaetis or Alainites. Although we tentatively treat Nigrobaetis and Alainites as distinct genera, they are considered to be paraphyletic taxa.     

The taxonomic utility of micromorphology in Lepidaploa (Vernonieae: Asteraceae)

Lepidaploa belongs to tribe Vernonieae, one of the most complex tribes of Asteraceae, and the relationships within Lepidaploa and among related genera are poorly understood. Microcharacters may be of taxonomic value and may be used in the identification of taxa at different ranks. To evaluate the reliability of microcharacters as taxonomic markers in this group, we analysed the micromorphology of phyllaries, florets and cypselae in detail in 23 species of Lepidaploa .The species were studied using stereo, light, and scanning electron microscopy. Eight trichome types (eglandular and glandular) were observed on phyllaries, florets and cypselae, in addition to crystals, idioblasts and other microstructures. The results demonstrates that the ocurrence of different combinations of trichome types and crystals, presence of a stylar basal node, idioblasts and glandular apical anther appendages are highly useful to differentiate between related species of Lepidaploa and a diagnostic key using these characters is presented. However, these characters are not of much use to distinguish between closely related genera of Vernonieae since most characters appear homeoplasic and are found in representatives of different genera.     

On the phylogenetic position and systematics of extant and fossil Aclopinae (Coleoptera: Scarabaeidae)

The Aclopinae is a small subfamily within the family Scarabaeidae. It currently comprises five extant genera with 28 species, and eight fossil genera with 25 species. The systematic position of Aclopinae within the family Scarabaeidae is uncertain, particularly because representative species of Aclopinae have been absent in previous phylogenetic studies. Here we performed phylogenetic analyses using morphological and molecular data to investigate the phylogenetic position of fossil and extant Aclopinae. For this objective, we expanded and revised a former morphological data matrix (composed of 68 characters) including all extant genera of Aclopinae. We complemented our morphological investigations with a molecular phylogenetic analysis based on four genes of several extant taxa of Aclopinae and a wide sample of diverse Scarabaeoidea. Our phylogenetic analyses show that all the type species of the fossil genera formerly included within Aclopinae do not belong within the extant Aclopinae clade and support both the exclusion of those fossil taxa and the monophyly of the extant genera of Aclopinae: Aclopus Erichson, Desertaclopus Ocampo & Mondaca, Gracilaclopus Ocampo & Mondaca, Neophanaeognatha Allsopp and Phanaeognatha Hope. Our results also show that the fossil taxa Prophaenognatha robusta Bai et al. and Ceafornotensis archratiras Woolley are closely related to Ochodaeidae, while the remaining type species of fossils formerly included in Aclopinae (Cretaclopus longipes (Ponomarenko), Holcorobeus vittatus Nikritin, Juraclopus rodhendorfi Nikolajev, Mesaclopus mongolicus (Nikolajev), and Mongolrobeus zherikhini Nikolajev) belong to a distinct lineage closely related to Diphyllostomatidae. Based on these results, the subfamily Aclopinae appears monophyletic and sister to the ‘pleurostict’ lineage. Consequently, we propose the following changes to the current classification of the fossil taxa: Holcorobeus monreali (Gómez‐Pallerola) belongs to Carabidae (incertae sedis) as proposed by the original author, and we place Ceafornotensis Woolley, Cretaclopus Nikolajev, Holcorobeus Nikritin, Juraclopus Nikolajev, Mesaclopus Nikolajev, Mongolrobeus Nikolajev and Prophaenognatha Bai et al. in Scarabaeoidea (incertae sedis). Furthermore, we provide an identification key to, and diagnoses of, the genera, illustrations of diagnostic characters and checklists of their included species. The evolutionary perspective presented provides new insights into the evolution of the pleurostict condition in Scarabaeoidea and the biogeography of this group, which is now regarded as Gondwanan, probably evolving during the Cretaceous and not from the upper Jurassic as previously assumed.     

Morphological phylogeny of the Tegulinae (Mollusca: Vetigastropoda) reinforces a Turbinidae position

A cladistic analysis of the Tegulinae (Turbinidae) is presented using 132 morphological characters and 41 taxa. Tegulinae is recovered and is sister to Prisogaster niger (Prisogasterinae) within the family Turbinidae. This scenario, with Tegulinae as a subfamily within Turbinidae, corroborates with the most molecular analyses. Tegulinae comprises >40 extant species, belonging to eight genera. Morphological studies have not resolved the placement of Tegulinae within Trochoidea sufficiently, and the systematic positions of the genera have never been investigated as a primary objective. The present morphology-based analysis of genus-level relationships within Tegulinae provides a robust, phylogenetic diagnosis of each group, rooted on a firm hypothesis of evolutionary relationships. An additional search was performed to include the tegulines Omphalius nigerrimus and Carolesia blakei terminals using unweighted and implied weighting. Our morphological data provide a solid foundation for ensuing systematic research on Tegulinae, as well as Trochoidea, and evidence facilitating the diagnosis of generic and suprageneric groups.     

Bio-sintering processes in hexactinellid sponges: Fusion of bio-silica in giant basal spicules from Monorhaphis chuni

The two sponge classes, Hexactinellida and Demospongiae, comprise a skeleton that is composed of siliceous skeletal elements (spicules). Spicule growth proceeds by appositional layering of lamellae that consist of silica nanoparticles, which are synthesized via the sponge-specific enzyme silicatein. While in demosponges during maturation the lamellae consolidate to a solid rod, the lamellar organization of hexactinellid spicules largely persists. However, the innermost lamellae, near the spicule core, can also fuse to a solid axial cylinder. Similar to the fusion of siliceous nanoparticles and lamella, in several hexactinellid species individual spicules unify during sintering-like processes. Here, we study the different stages of a process that we termed bio-sintering, within the giant basal spicule (GBS) of Monorhaphis chuni. During this study, a major GBS protein component (27 kDa) was isolated and analyzed by MALDI-TOF-MS. The sequences were used to isolate and clone the encoding cDNA via degenerate primer PCR. Bioinformatic analyses revealed a significant sequence homology to silicatein. In addition, the native GBS protein was able to mediate bio-silica synthesis in vitro. We conclude that the syntheses of bio-silica in M. chuni, and the subsequent fusion of nanoparticles to lamellae, and finally to spicules, are enzymatically-driven by a silicatein-like protein. In addition, evidence is now presented that in hexactinellids those fusions involve sintering-like processes.     

Phylogeny of the hyper‐diverse rove beetle subtribe Philonthina with implications for classification of the tribe Staphylinini (Coleoptera: Staphylinidae)

With 71 genera and over 2700 described species, Philonthina is the most speciose subtribe of rove beetle tribe Staphylinini and forms a major component of the largest remaining higher systematics challenge in Staphylinini, the ‘Staphylinini propria’ clade. A related systematics issue concerns the position of the genus Holisus (Hyptiomina), which was recovered within the Neotropical philonthine lineage in several recent analyses of morphology. With the aims of resolving the phylogeny of Philonthina and the position and, thus, validity of Hyptiomina, we performed phylogenetic analyses of the tribe Staphylinini based on molecular (six genes, 4471 bp) and morphological (113 characters) data including 138 taxa from all relevant lineages of Staphylinini. We found that ‘Staphylinini propria’ is a monophylum consisting of six lineages: current subtribes Anisolinina, Philonthina, Staphylinina and Xanthopygina; and two new subtribes, Algonina Schillhammer and Brunke and Philothalpina Chatzimanolis and Brunke. While the previously hypothesized Neotropical lineage of Philonthina was corroborated, Holisus was recovered as a separate subtribe, outside of Philonthina, within an informal ‘Southern Hemisphere clade’. Based on our analyses, we propose tentative new concepts of the polyphyletic genera Belonuchus and Philonthus. We propose the following taxonomic changes: synonymy of the subtribes Staphylinina Latreille (valid name) and Eucibdelina Sharp; resurrection of genera Barypalpus Cameron and Trapeziderus Motschulsky from synonymy with Rientis Sharp and Belonuchus Nordmann, respectively; transfer of 38 Belonuchus species, 16 Hesperus Fauvel species and one Philonthus Stephens species to Trapeziderus as new combinations; transfer of two Hesperus species to Eccoptolonthus Bernhauer as new combinations; transfer of one Belonuchus species to Paederomimus Sharp as a new combination; and transfer of Pridonius Blackwelder new status from its position as a subgenus of Quedius (subtribe Quediina) to Philonthina as a genus, and new combinations for its two described species.     

Reinstatement of Lophocoleaceae (Jungermanniopsida) based on chloroplast gene rbcL data: exploring the importance of female involucres for the systematics of Jungermanniales

Maximum likelihood analysis of 113 rbcL sequences leads to a well resolved phylogeny of Jungermanniales. All species with perigynia or marsupia are found in one clade, whereas species with coelocaules are placed in several lineages. The broadly circumscribed Geocalycaceae (including Lophocoleaceae) of most recent authors are resolved as polyphyletic. Geocalycaceae genera which develop female involucres without involvement of stem tissue (Chiloscyphus, Heteroscyphus, Leptoscyphus, Physotheca) form a robust clade which is placed sister to Plagiochilaceae whereas the genera with involucres originating at least partly from stem tissue (Geocalycaceae s.str., Geocalyx, Harpanthus, Saccogyna) are nested within the paraphyletic Jungermanniaceae. This topology leads to the exclusion of the strictly perianth-bearing species from Geocalycaceae and the reinstatement of Lophocoleaceae. Campanocolea is nested within Chiloscyphus. Physotheca and Chiloscyphus breutelii are placed within an unsupported clade with several accessions of Leptoscyphus. Heteroscyphus forms a paraphyletic grade at the base of Chiloscyphus.