A Total Evidence Phylogeny of the Arctoidea (Carnivora: Mammalia): Relationships Among Basal Taxa

A total evidence phylogenetic analysis was performed for 14 extant and 18 fossil caniform genera using a data matrix of 5.6 kbp of concatenated sequence data from six independent loci and 80 morphological characters from the cranium and dentition. Maximum parsimony analysis recovered a single most parsimonious cladogram (MPC). The topology of the extant taxa in the MPC agreed with previous molecular phylogenies. Phylogenetic positions for fossil taxa indicate that several taxa previously described as early members of extant families (e.g., Bathygale and Plesictis) are likely stem taxa at the base of the Arctoidea. Taxa in the “Paleomustelidae” were found to be paraphyletic, but a monophyletic Oligobuninae was recovered within this set of taxa. This clade was closely related to the extant genera Gulo and Martes, therefore, nested within the extant radiation of the family Mustelidae. This analysis provides a resolution to several discrepancies between phylogenies considering either fossil taxa or extant taxa separately, and provides a framework for incorporating fossil and extant taxa into comprehensive combined evidence analyses.     

Phylogeny of the Acanthocephala based on morphological characters

Only four previous studies of relationships among acanthocephalans have included cladistic analyses, and knowledge of the phylogeny of the group has not kept pace with that of other taxa. The purpose of this study is to provide a more comprehensive analysis of the phylogenetic relationships among members of the phylum Acanthocephala using morphological characters. The most appropriate outgroups are those that share a common early cell-cleavage pattern (polar placement of centrioles), such as the Rotifera, rather than the Priapulida (meridional placement of centrioles) to provide character polarity based on common ancestry rather than a general similarity likely due to convergence of body shapes. The phylogeny of 22 species of the Acanthocephala was evaluated based on 138 binary and multistate characters derived from comparative morphological and ontogenetic studies. Three assumptions of cement gland structure were tested: (i) the plesiomorphic type of cement glands in the Rotifera, as the sister group, is undetermined; (ii) non-syncytial cement glands are plesiomorphic; and (iii) syncytial cement glands are plesiomorphic. The results were used to test an early move of Tegorhynchus pectinarius to Koronacantha and to evaluate the relationship between Tegorhynchus and Illiosentis. Analysis of the data-set for each of these assumptions of cement gland structure produced the same single most parsimonious tree topology. Using Assumptions i and ii for the cement glands, the trees were the same length (length = 404 steps, CI = 0.545, CIX = 0.517, HI = 0.455, HIX = 0.483, RI = 0.670, RC = 0.365). Using Assumption iii, the tree was three steps longer (length = 408 steps, CI = 0.539, CIX = 0.512, HI = 0.461, HIX = 0.488, RI = 0.665, RC = 0.359). The tree indicates that the Palaeacanthocephala and Eoacanthocephala both are monophyletic and are sister taxa. The members of the Archiacanthocephala are basal to the other two clades, but do not themselves form a clade. The results provide strong support for the Palaeacanthocephala and the Eoacanthocephala and the hypothesis that the Eoacanthocephala is the most primitive group is not supported. Little support for the Archiacanthocephala as a monophyletic group was provided by the analysis. Support is provided for the recognition of Tegorhynchus and Illiosentis as distinct taxa, as well as the transfer of T. pectinarius to Koronacantha.     

HOMOPLOID RETICULATE EVOLUTION IN HELIANTHUS (ASTERACEAE): EVIDENCE FROM RIBOSOMAL GENES

Phylogenetic relationships among the 21 taxa comprising Helianthus sect. Helianthus and three outgroup species were assessed by restriction site mapping of the 18S-25S nuclear ribosomal RNA gene family. Wagner parsimony analysis of the 41 restriction site or length mutations observed produced a single 59-step most parsimonious tree. This tree was then compared to a cytoplasmic-based plastid phylogeny for this group. Several major discrepancies were observed between the two phylogenies suggesting both recent and ancient introgression. Furthermore, three cases of diploid hybrid speciation are unambiguously documented and a fourth case is suggested. These data are interpreted to suggest that evolution in Heliathus is reticulate rather than exclusively dichotomous and branching.     

New late Permian Chonetids (brachiopoda) from Southeastern Pamirs

New chonetid genera and species Subtilichonetes pamiricus gen. et sp. nov. (subfamily Undulellinae, family Rugosochonetidae) and Parvichonetes igrimiusensis gen. et sp. nov. (subfamily Tornquistiinae, family Anopliidae) are described from the Takhtabulak Formation, Changhsingian Stage of the Upper Permian of Southeastern Pamir.     

Refining molluscan characters: morphology, character coding and a phylogeny of the Caenogastropoda

Midgut morphology of gastropod molluscs has been underutilized as a resource of characters for phylogenetic analysis. The exclusion of these features reflects the inference that they will be uninformative in determining phylogenetic relationships because they are functionally correlated. In general, it has been hypothesized that the style sac form of midgut is an adaptation to microphagy and becomes secondarily simplified in taxa that have adopted a macrophagous/carnivorous habit with a corresponding increase in extracellular digestion (i.e. radular trituration, gizzards and/or foregut glands). This assumption has resulted in the formulation of adaptive scenarios concerning gastropod alimentary systems, which are mapped onto phylogenetic hypotheses derived from other characters. However, any conclusions regarding phylogenetic utility, and therefore homology, must be realized within a cladistic context. For this analysis, a multi‐organ system anatomical data set was assembled for 16 caenogastropods and two outgroups. The data matrix comprises 64 characters and includes many systems poorly represented in previous broad‐based comparative surveys, such as the alimentary and reno‐pericardial systems. In addition, several taxa were included for which no comprehensive anatomical studies have been available (Cyclophoroidea, Ptenoglossa). Phylogenetic analysis with NONA 1.6 resulted in two most‐parsimonious trees with length 188, CI = 0.53 and RI = 0.63, differing only in the placement of Prunum apicinum and Conus jaspideus. The topology of the strict consensus (Macleaniella Theodoxus((Neocyclotus Marisa)((Lampanella((Petaloconchus Strombus)(Crepidula Bithyia)))(Littorina(Neverita(Cypraea(Nitidiscala(Panarona(Prunum Conus(Ilyanassa Urosalpinx)))))))))), is largely congruent with several independent estimates based on both morphological and molecular data, supporting caenogastropod monophyly and monphyly of the Architaenioglossa, Sorbeoconcha and Neogastropoda. To evaluate phylogenetic utility of the midgut, a broad sampling of taxa was included representing a diversity of feeding modes, food preferences and alimentary morphologies. Character optimization revealed that the evolution of midgut structure is highly mosaic, cutting broadly across patterns of feeding, diet and foregut complexity, to a degree previously unappreciated. In addition, features from the foregut (subradular organ, oesophageal folding) and midgut (position of gastric shield) are broadly distributed among large clades of caenogastropods, providing critical basal synapomorphies within the group. This demonstrates that the gut is an unexploited resource for important and informative characters in higher order systematics of caenogastropods. © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 137 , 447–554.     

Phantoms of Gondwana?—phylogeny of the spider subfamily Mynogleninae (Araneae: Linyphiidae)

This is the first genus‐level phylogeny of the subfamily Mynogleninae. It is based on 190 morphological characters scored for 44 taxa: 37 mynoglenine taxa (ingroup) representing 15 of the 17 known genera and seven outgroup taxa representing the subfamilies Stemonyphantinae, Linyphiinae (Linyphiini and Micronetini), and Erigoninae, and a representative of the family Pimoidae, the sister‐group to Linyphiidae. No fewer than 147 of the morphological characters used in this study are new and defined for this study, and come mainly from male and female genitalia. Parsimony analysis with equal weights resulted in three most parsimonious trees of length 871. The monophyly of the subfamily Mynogleninae and the genera Novafroneta, Parafroneta, Laminafroneta, Afroneta, Promynoglenes, Metamynoglenes, and Haplinis are supported, whereas Pseudafroneta is paraphyletic. The remaining seven mynoglenine genera are either monotypic or represented by only one taxon. Diagnoses are given for all genera included in the analysis. The evolution of morphological traits is discussed and we summarize the diversity and distribution patterns of the 124 known species of mynoglenines. The preferred topology suggests a single origin of mynoglenines in New Zealand with two dispersal events to Africa, and does not support Gondwana origin.     

Chloroplast DNA variation in the cultivated and wild olive taxa of the genus Olea L.

Polymorphism in the lengths of restriction fragments of the whole cpDNA molecule were studied in 15 taxa (species or subspecies) of the genus Olea. From restriction analysis using nine endonucleases, 28 site mutations and five length polymorphisms were identified, corresponding to 12 distinct chlorotypes. From a phenetic analysis based on a Nei’s dissimilarity matrix and a Dollo parsimony cladistic analysis using, as an outgroup, a species of the genus Phillyrea close to Olea, the ten taxa of section Olea were distinguished clearly from the five taxa of section Ligustroides which appear to posses more ancestral cpDNA variants. Within the section Ligustroides, the tropical species from central-western Africa, Olea hochtetteri, showed a chlorotype which differed substantially from those of the other four Olea taxa growing in southern Africa, supporting a previous assessment according to which O. hochtetteri may have been subjected to a long period of geographical isolation from the other Olea taxa. Within the Olea section, three phyla were identified corresponding to South and East Africa taxa, Asiatic taxa, and a group including Saharan, Macaronesian and Mediteranean taxa, respectively. On the basis of cpDNA variation, the closest Olea taxa to the single Mediterranean species, Olea europaea, represented by its very predominant chlorotype, observed in both wild and cultivated olive, were found to be Olea laperrinei (from the Sahara), Olea maroccana (from Maroccan High Atlas) and Olea cerasiformis (from Macaronesia). These three taxa, which all share the same chlorotype, may have a common maternal origin. Received: 5 December 1999 / Accepted: 30 December 1999     

Morphological phylogeny of the variable fly family Stratiomyidae (Insecta,Diptera)

Brammer, C. A. & von Dohlen, C. D. (2010). Morphological phylogeny of the variable fly family Stratiomyidae (Insecta, Diptera). —Zoologica Scripta, 39, 363–377. Stratiomyidae is a dipteran family distributed worldwide and containing 2800 species classified into 12 subfamilies. Previous phylogenetic work on the Stratiomyidae consisted of a 20‐character morphological analysis of the subfamilies [ World Catalog of the Stratiomyidae (Insect: Diptera). Leiden: Backhuys Publishers, 2001 ], and a molecular study using 69 taxa and two gene regions [ Molecular Phylogenetics and Evolution, 43, 2007, 660 ]. In this study, we present an expanded morphological cladistic analysis using 92 characters and 80 taxa, representing 36 of 39 described genera and all 12 Stratiomyidae subfamilies, as well as Xylomyidae and Pantophthalmidae outgroups. Data are analysed under maximum parsimony with all characters unordered and weighted equally; nodal support is assessed with the bootstrap and Bremer index. The strict consensus of all shortest trees is well resolved, and many of the deeper nodes are supported, although the root is ambiguous. Antissinae, Stratiomyinae, Sarginae and the diverse Clitellariinae are not monophyletic. Clitellariinae are positioned across several lineages, with most species grouped into a single, unsupported clade. Many of the well‐supported relationships are consistent with several aspects of the previous studies. The position of Exodontha remains elusive. Character support for subfamilies and other major clades is discussed.     

The cytochrome b gene as a phylogenetic marker: the limits of resolution for analyzing relationships among cichlid fishes

The mitochondrial cytochrome b (cyt-b) gene is widely used in systematic studies to resolve divergences at many taxonomic levels. The present study focuses mainly on the utility of cyt-b as a molecular marker for inferring phylogenetic relationship at various levels within the fish family Cichlidae. A total of 78 taxa were used in the present analysis, representing all the major groups in the family Cichlidae (72 taxa) and other families from the suborders Labroidei and Percoidei. Gene trees obtained from cyt-b are compared to a published total evidence tree derived from previous studies. Minimum evolution trees based on cyt-b data resulted in topologies congruent with all previous analyses. Parsimony analyses downweighting transitions relative to transversions (ts1:tv4) or excluding transitions at third codon positions resulted in more robust bootstrap support for recognized clades than unweighted parsimony. Relative rate tests detected significantly long branches for some taxa (LB taxa) which were composed mainly by dwarf Neotropical cichlids. An improvement of the phylogenetic signal, as shown by the four-cluster likelihood mapping analysis, and higher bootstrap values were obtained by excluding LB taxa. Despite some limitations of cyt-b as a phylogenetic marker, this gene either alone or in combination with other data sets yields a tree that is in agreement with the well-established phylogeny of cichlid fish. Received: 11 October 2000 / Accepted: 26 February 2001     

Molecular Phylogeny of Grey Mullets (Teleostei: Mugilidae) in Greece: Evidence from Sequence Analysis of mtDNA Segments

Mitochondrial DNA sequence analysis has been used to explore genetic differentiation and phylogenetic relationships among five species of the Mugilidae family, Mugil cephalus, Chelon labrosus, Liza aurata, Liza ramada, and Liza saliens. DNA was isolated from samples originating from the Messolongi Lagoon in Greece. Three mtDNA segments (12s rRNA, 16s rRNA, and CO I) were PCR amplified and sequenced. Sequencing analysis revealed that the greatest genetic differentiation was observed between M. cephalus and all the other species studied, while C. labrosus and L. aurata were the closest taxa. Dendrograms obtained by the neighbor-joining method and Bayesian inference analysis exhibited the same topology. According to this topology, M. cephalus is the most distinct species and the remaining taxa are clustered together, with C. labrosus and L. aurata forming a single group. The latter result brings into question the monophyletic origin of the genus Liza.     

The Postcranial Morphology of Ptilocercus lowii (Scandentia, Tupaiidae): An Analysis of Primatomorphan and Volitantian Characters

The eutherian orders Scandentia, Primates, Dermoptera, and Chiroptera have been grouped together by many morphologists, using various methods and data sets, into the cohort Archonta. Molecular evidence, however, has supported a clade (called Euarchonta) that includes Scandentia, Primates, and Dermoptera, but not Chiroptera. Within Archonta, some systematists have grouped Dermoptera and Chiroptera in Volitantia, while others have grouped Dermoptera and Primates in Primatomorpha. The order Scandentia includes the single family Tupaiidae, with two subfamilies, Ptilocercinae and Tupaiinae. Ptilocercinae is represented only by Ptilocercus lowii, which has been said to be the taxon most closely approximating the ancestral tupaiid. However, most researchers working on archontan phylogeny typically do not treat the order Scandentia as being polymorphic. They usually use Tupaia to represent Scandentia, despite the fact that Ptilocercus is quite distinct from Tupaia and has been argued to be the more plesiomorphic of the two taxa. In this study, a character analysis was performed on postcranial features that have been used to support the competing Primatomorpha and Volitantia hypotheses. In recognition of the polymorphic nature of Scandentia, taxonomic sampling within Scandentia was increased to include Ptilocercus. The postcranium of Ptilocercus was compared to that of tupaiines, euprimates, plesiadapiforms, dermopterans, and chiropterans. Several character states used to support either Primatomorpha or Volitantia, while not found in Tupaia, were found in Ptilocercus. While these features may have evolved independently in Ptilocercus, it is perhaps more likely that they represent features that first evolved in the ancestral archontan and were then lost in one of the extant orders. This character analysis greatly reduces the supportive evidence for the Primatomorpha hypothesis.     

A simple parameter of dispersion index that serves as an adjunct to karyotype asymmetry

In order to refine the measure of karyotype asymmetry a new chromosomal parameter of dispersion index is proposed that has the potential to decipher even the minor karyotypic variations, thus permitting further evolutionary gradations to the karyotype asymmetry classes of Stebbins. The higher the dispersion index, the more specialized would be the karyotype. The dispersion index takes into account the variance lor gradual change in chromosome size within a complementvis-a-vis variance for the position of centromere in a karyotypic totality. The dispersion index is calculated as the proportionate measure of centromeric gradient to the coefficient of variation for chromosome length; wherin centromeric gradient = length of median short arm — length of median chromosome. Thus, the three most important karyotypic criteriaviz., differences in: absolute chromosome size, position of centromere and relative chromosome size, are all covered in the proposed parameter. The effectiveness of dispersion index has been tested on a plant taxa,Papaver L., where karyomorphological details, nuclear DNA content, and morphotaxonomic parameters have been amply elucidated from an evolutionary stand point. It is hoped that dispersion index would find immense utility in delimiting species interrelationships particularly in the closely related taxa, when applied in conjunction with other systematic parameters.     

Global interrelationships of Plesiosauria (Reptilia,Sauropterygia) and the pivotal role of taxon sampling in determining the outcome of phylogenetic analyses

Previous attempts to resolve plesiosaurian phylogeny are reviewed and a new phylogenetic data set of 66 taxa (67% of ingroup taxa examined directly) and 178 characters (eight new) is presented. We recover two key novel results: a monophyletic Plesiosauridae comprising Plesiosaurus dolichodeirus, Hydrorion brachypterygius, Microcleidus homalospondylus, Occitanosaurus tournemirensis and Seeleyosaurus guilelmiimperatoris; and five plesiosaurian taxa recovered outside the split between Plesiosauroidea and Pliosauroidea. These taxa are Attenborosaurus conybeari, ‘Plesiosaurus’macrocephalus and a clade comprising Archaeonectrus rostratus, Macroplata tenuiceps and BMNH 49202. Based on this result, a new name, Neoplesiosauria, is erected for the clade comprising Plesiosauroidea and Pliosauroidea. Taxon subsamples of the new dataset are used to simulate previous investigations of global plesiosaurian relationships. Based on these simulations, most major differences between previous global phylogenetic hypotheses can be attributed to differences in taxon sampling. These include the position of Leptocleididae and Polycotylidae and the monophyly or paraphyly of Rhomaleosauridae. On this basis we favour the results recovered by our, larger analysis. Leptocleididae and Polycotylidae are sister taxa, forming a monophyletic clade within Plesiosauroidea, indicating that the large‐headed, short‐necked ‘pliosauromorph’ body plan evolved twice within Plesiosauria. Rhomaleosauridae forms the monophyletic sister taxon of Pliosauridae within Pliosauroidea. Problems are identified with previous phylogenetic definitions of plesiosaurian clades and new, stem‐based definitions are presented that should maintain their integrity over a range of phylogenetic hypotheses. New, rank‐free clade names Cryptoclidia and Leptocleidia are erected to replace the superfamilies Cryptoclidoidea and Leptocleidoidea. These were problematic as they were nested within the superfamily Plesiosauroidea. The incongruence length difference test indicates no significant difference in levels of homoplasy between cranial and postcranial characters.     

Plastomic data shed new light on the phylogeny,biogeography, and character evolution of the family Crassulaceae

Crassulaceae is a mid-sized family of angiosperms, most species of which are herbaceous succulents, usually with 5-merous flowers and one or two whorls of stamens. Although previous phylogenetic studies revealed seven major “clades” in Crassulaceae and greatly improved our understanding of the evolutionary history of the family, relationships among major clades are still contentious. In addition, the biogeographic origin and evolution of important morphological characters delimiting infrafamilial taxa have not been subject to formal biogeographic and character evolution analyses based on a well-supported phylogeny backbone. In this study, we used plastomic data of 52 species, representing all major clades revealed in previous studies to reconstruct a robust phylogeny of Crassulaceae, based on which we unraveled the spatiotemporal framework of diversification of the family. We found that the family may originate in southern Africa and then dispersed to the Mediterranean, from there to eastern Asia, Macaronesia, and North America. The crown age of Crassulaceae was dated at ca. 63.93 million years ago, shortly after the Cretaceous–Paleogene (K-Pg) boundary. We also traced the evolution of six important morphological characters previously used to delimit infrafamilial taxa and demonstrated widespread parallel and convergent evolution of both vegetative (life form and phyllotaxis) and floral characters (number of stamen whorls, petals free or fused, and flower merism). Our results provide a robust backbone phylogeny as a foundation for further investigations, and also some important new insights into biogeography and evolution of the family Crassulaceae.     

Molecular phylogeny of grapsoid crabs (Decapoda, Brachyura) and allies based on two mitochondrial genes and a proposal for refraining from current superfamily classification

Grapsoid and ocypodoid crabs receive a lot of attention in the literature due to their predominance and important role as primary and secondary consumers in intertidal as well as supratidal marine habitats. They are especially species‐rich in the tropics, where they have been found to repeatedly invade terrestrial and freshwater habitats. However, the systematics of the crabs belonging to these two superfamilies is still not settled, despite recent steps clarifying phylogenetic relationships and introducing new taxa. In this study, a molecular phylogeny of grapsoid crabs primarily based on East African representatives is constructed based on DNA sequences of the mitochondrial small and large ribosomal subunits (12S and 16S rRNA), thus complementing previous molecular taxonomic studies that had been carried out with the American and East Asian fauna. In addition, selected representatives of all ocypodoid families and subfamilies were included. The monophyly of Grapsidae, Ocypodidae (sensu stuctu), Sesarmidae and Varunidae is well confirmed, if the genera Cyclograpsus, Helice are considered Varunidae and Euchirograpsus a Plagusiidae, as previously suggested. The monophyly of the family Gecarcinidae cannot be supported with our data. The family Plagusiidae in its present composition is polyphyletic. Special attention was given to the large family Sesarmidae, which has many endemic genera in the Indo‐West Pacific. According to this study, two of the most speciose genera, Chiromantes and Parasesarma, are not monophyletic and need to be redefined. On the higher taxonomic level, it becomes evident that both superfamilies, Grapsoidea and Ocypodoidea, are not monophyletic in their current composition, as exemplified by a proposed sister group relationship of Varunidae and Macrophthalmidae. These results confirm those from previous molecular studies and we therefore propose to refrain from the traditional use of the Grapsoidea and Ocypodoidea as monophyletic superfamilies and treat the constituent families separately.     

A molecular phylogeny of the groupers of the subfamily Epinephelinae (Serranidae) with a revised classification of the Epinephelini

The phylogenetic relationships among the fishes in the perciform tribe Epinephelini (Serranidae) have long been poorly understood, in large part because of the numerous taxa that must be considered and the large, circumtropical distribution of the group. In this study, genetic data from two nuclear (Tmo-4C4 and histone H3) and two mitochondrial (16S and 12S) genes were gathered from 155 serranid and acanthomorph species as a means of developing a phylogenetic hypothesis using both maximum-likelihood and -parsimony criteria. The maximum-parsimony analysis recovered 675 most parsimonious trees of length 5703 steps (CI = 0.2523, HI = 0.7477, RI = 0.6582), and the maximum-likelihood analysis recovered 1 tree at −lnLikelihood = 28279.58341. These phylogenetic hypotheses are discussed in light of previous morphological evidence to evaluate the evolutionary history of the group and their implications for the currently recognized taxonomy. Our results question the monophyly of the Serranidae, as well as the genera Cephalopholis, Epinephelus, and Mycteroperca as currently defined. The Serranidae is monophyletic only with the exclusion of the genera Acanthistius and Niphon. We propose a revised classification of the tribe Epinephelini that reflects the hypothesized shared ancestry of the group and recognizes 11 genera: Alphestes, Cephalopholis, Dermatolepis, Epinephelus, Gonioplectrus, Hyporthodus (which is resurrected for 11 species of deep-bodied groupers), Mycteroperca (including 7 species heretofore allocated to Epinephelus), Plectropomus, Saloptia, Triso, and Variola.     

Mouthpart structure of Stylogymnusa subantarctica Hammond, 1975 (Coleoptera: Staphylinidae: Aleocharinae) with a reanalysis of the phylogenetic position of the genus

Structure, particularly of the mouthparts, of the unusual aleocharine staphylinid Stylogymnusa subantarctica Hammond was reevaluated. The mouthparts are described in detail, and drawings of their structure are provided. Several features were found to be misinterpreted in previous accounts of the structure of this aleocharine. In particular, the stylate structures found on the prementum are not a highly modified, hollow, stylate glossa, as had been previously suggested. Instead, these structures represent highly modified labial palpi; the segments are completely fused, all signs of typical setae of the labial palpi are missing, and the palpi are very long and slender. In addition, there is no sign that they are hollow. Small, slender, membranous lobes at the base of, and between, the palpi are interpreted as the true glossae (=ligula). Reinterpretation of these characters, and greatly improved phylogenetic techniques in the 20 years since previous phylogenies had been proposed, allowed testing of previous hypotheses of the phylogenetic position of Stylogymnusa and the relationships of the tribes Gymnusini and Deinopsini within the Aleocharinae. Phylogenetic analysis of 7 out‐group and 12 in‐group taxa, based on 84 characters and 231 character states, produced 3 equally most‐parsimonious trees (tree length = 253, C.I. = 0.557, R.I. =0.674, rescaled C.I. = 0.376). These trees provide strong support for a monophyletic lineage consbting of Stylogymnusa, Gymnusa and the Deinopsini but only weak support for a monophyletic Gymnusini (Gymnusa + Stylogymnusa). Bootstrap analysis based on 1000 resampling repetitions showed the following monophyletic clades to be well supported by the dataset: Aleocharinae (79%), Stylogymnusa+Gymnusa+ Deinopsini (99%), Deinopsis+Adinopsis (100%), and representatives of Myllaenini + Athetini + Aleocharini + Homalotini + Oxypodini (the “higher Aleocharinae”) (88%). Weakly supported clades in the bootstrap analysis were: Paraconosoma+ Gymnusini + Deinopsini (56%), Anacyptus+‘higher Aleocharinae’ (54%), and Athetini + Oxypodini + Homalotini + Aleocharini (52%).