A revision of Benjaminia (Hymenoptera: Ichneumonidae, Campopleginae)

Abstract The campoplegine genus Benjuminia is herein revised, with ten species being described as new, in addition to the five previously known species. Phylogenetic relationships of Benjuminia to other genera are discussed, and a cladogram presented for the species. Two monophyletic species groups are recognized, one being confined to the Palearctic and the other to the Nearctic. Examination of larval morphology not only fails to reveal larval characters for distinguishing Benjuminia but leads to the conclusion that use of larval morphology to identify campoplegine species is a dubious practice.     

Pattern of Phylogenetic Relationships among Members of the Tribe Melitaeini (Lepidoptera: Nymphalidae) Inferred from Mitochondrial DNA Sequences

We report a cladistic analysis of 77 butterfly species of the tribe Melitaeini (Lepidoptera: Nymphalidae) based on mitochondrial DNA gene sequences. We sequenced ca. 536 bp from the 16S ribosomal DNA (rDNA) and a 1422-bp sequence from the cytochrome oxidase I gene. Alignments are critical to statements of homology, especially when aligning rDNA sequences. We aligned the 16S sequences using conventional methods and direct optimization. We found that direct optimization of the sequences produced the best alignments and our preferred phylogenetic hypothesis. Our results suggest that many of the previously proposed genera are paraphyletic and we conclude that there are four monophyletic groups of species in our cladogram: the Euphydryas group, the Phyciodes group, the Chlosyne group, and the Melitaea group. The following genera are found to be paraphyletic: Castilia, Chlosyne, Didymaeformia, Eresia, Melitaea , and Thessalia . In addition, recognition of the monophyletic genera Cinclidia, Mellicta , and Telenassa would render other genera paraphyletic. Our phylogenetic hypothesis indicates that the melitaeines originated in the Nearctic and have colonized the Neotropics three times and the Palaearctic twice.     

A new genus of phreatoicidean isopod (Crustacea) from the north Kimberley region, Western Australia

A new genus and species of phreatoicidean isopod, Crenisopus acinifer, has been collected from a freshwater spring in the northern Kimberley region of Western Australia. Empirical cladistic analysis of 10 exemplars of phreatoicidean genera found a single cladogram. The new genus and species assumed a basal position in the Phreatoicidea, placing it within the family Amphisopodidae sensu lata. This family, however, was not monophyletic in the preliminary cladogram, suggesting that the taxonomic structure of the suborder must be revised. The cladogram provided evidence for the monophyly of the Phreatoicidae and its New Zealand clade. The analysis suggested that clades of modern phreatoicideans diverged from one another during the Mesozoic Era after they entered fresh water, but prior to the fragmentation of East Gondwana.     

Reconciling taxonomy and phylogeny in the bristleworm family Eunicidae (polychaete,Annelida)

Eunicid annelids inhabit diverse marine habitats worldwide, have ecological and economic importance and have been pictured in the news as giant predator worms. They compose a traditional stable taxon recently supported as monophyletic but characterized by plesiomorphies. Most genera within the family have been recovered as paraphyletic in previous studies. We present a phylogenetic hypothesis for eunicid based on molecular (COI, 16S rDNA, 18S rDNA) and morphological data (213 characters), including an explicit attempt to account for serial homology. Eunicidae as well as monophyletic genera Marphysa sensu stricto and Lysidice is redefined based on synapomorphies. Nematonereis is synonymized to Lysidice. Leodice and Nicidion are resurrected to name monophyletic groups including species previously included in Eunice and Marphysa sensu lato. Traditional diagnostic characters such as the absence/presence of peristomial cirri, lateral antennae and branchiae are homoplasies and not informative at the generic level. Different coding of traditional characters (i.e. articulation of prostomial appendages) and novel characters of prostomial features and regionalization of the body support the monophyly of the family and genera level clades. Thus, the phylogenetic hypothesis presented here and the evolution of characters provided background information for taxonomic changes yielding evolutionary meaningful classification and diagnoses for the family and genera.     

Phylogeny and Cladistic Classification of the Paramonostiliferous Family Plectonemertidae (Phylum Nemertea)

Abstract— Numerical phylogenetic analysis of nemertean species from the genera Plectonemertes, Katechonemertes and Acteonementes (marine), Argonemertes, Antiponemertes and Leptonemertes (terrestrial), Potamonemertes and Campbellonemertes (freshwater) in the family Plectonemertidae was carried out with the following aims. (1) To estimate the evolutionary relationships between and among these species. (2) To lest the hypothesis of Moore and Gibson that the analysed freshwater species originated from a terrestrial ancestor. (3) To classify the species in monophyletic genera. The analysis did not confirm the hypothesis that the freshwater habitat has been colonised via land. Instead, it suggests that both terrestrial and freshwater species within this family have evolved from a marine ancestor. The cladogram suggests that Antiponemetes is paraphyletic, and it is combined with Argonemertes to form a monophyletic genus.     

A molecular phylogeny of Panicum (Poaceae: Paniceae): tests of monophyly and phylogenetic placement within the Panicoideae

Panicum L. is a cosmopolitan genus with approximately 450 species. Although the genus has been considerably reduced in species number with the segregation of many taxa to independent genera in the last two centuries, Panicum remains a heterogeneous assemblage, as has been demonstrated in recent years. The genus is remarkably uniform in its floral characters but exhibits considerable variation in anatomical, physiological, and cytological features. As a result, several classifications, and criteria of what the genus should really include, have been postulated in modern literature. The purpose of this research, based on molecular data of the chloroplast ndhF gene, is to test the monophyly of Panicum, to evaluate infrageneric classifications, and to propose a robust phylogenetic hypothesis. Based on the present results, previous morphological and molecular phylogenetic studies, and inferred diagnostic morphological characters, we restrict Panicum sensu stricto (s.s.) to the former subgenus Panicum and support recognition of Dichanthelium, Phanopyrum, and Steinchisma as distinct genera. We have transfered other species of Panicum to other genera of the Paniceae. Most of the necessary combinations have been made previously, so few nomenclatural changes have been required. The remaining species of Panicum sensu lato (s.l.) are included within Panicum incertae sedis representing isolated species or species grouped within monophyletic clades. Additionally, we explore the performance of the three codon position characters in producing the supported phylogeny.     

<Emphasis Type="Italic">Aethycteron robisoni</Emphasis> n. sp. (Monogenea: Ancyrocephalidae) from the sunburst darter, <Emphasis Type="Italic">Etheostoma mihileze</Emphasis> Mayden (Perciformes: Percidae), in Arkansas,USA

Aethycteron robisoni n. sp. is described from the sunburst darter, Etheostoma mihileze Mayden (Perciformes: Percidae), in the Arkansas River Drainage of the Ozark Region in the Central Highlands of Arkansas, USA. Aethycteron robisoni morphologically most closely resembles A. caerulei Suriano & Beverley-Burton, 1982, A. moorei (Mizelle, 1940) and A. nigrei Suriano & Beverly-Burton, 1982, by possessing a male copulatory organ with a distinct distal curvature and spiraling sheath. The haptoral sclerites of A. robisoni, with the exception of the hooks, are distinctly larger than those of the other three species. This is the first time a monogenean parasite has been reported from E. mihileze as well as the first time the genus Aethycteron Suriano & Beverley-Burton, 1982 has been reported from Arkansas, USA.     

Phylogenetic relationship among Libellula, Ladona and Plathemis (Odonata: Libellulidae) based on DNA sequence of mitochondrial 16S rRNA gene

Abstract. The type genus for the dragonfly family Libellulidae is Libellula. At present, Libellula s.l. includes twenty-nine species, whose distribution is largely Nearctic. Whether two other libellulid taxa, Ladona and Plathemis, should be considered synonyms of Libellula, subgenera of Libellula, or separate genera, has been a subject of intermittent debate for over a century. Earlier proposals concerning Ladona and Plathemis were based on a limited number of morphological characters and lacked rigorous phylogenetic analyses. Therefore, we used the DNA sequence of a portion of the mitochondrial 16S rRNA gene and parsimony, maximum likelihood and neighbour-joining analyses to explore whether Ladona and Plathemis are monophyletic lineages distinct from Libellula. We obtained ≈ 415 bp of DNA sequence from twenty-three taxa including thirteen species of Libellula s.s., all three recognized species of Ladona, the two species of Plathemis and representatives of four other libellulid genera. Tetragoneuria williamsoni (Odonata: Corduliidae) was included as the outgroup. Parsimony analysis suggested that Ladona and Plathemis are monophyletic lineages distinct from Libellula s.s. with a sister group relationship between Libellula and Ladona. The monophyly of Ladona, Plathemis and Libellula was supported in > 90% of bootstrap replications and in trees five to ten steps longer than the most parsimonious trees. Relationships inferred from maximum likelihood and neighbour-joining analyses also supported the monophyly of Ladona and Plathemis. The four other libellulid genera included in the study formed a monophyletic clade distinct from Libellula, Ladona and Plathemis. Based on our analysis, we propose that Ladona and Plathemis be considered either genera or subgenera within Libellulidae.     

A preliminary phylogenetic analysis of the New World Helopini (Coleoptera,Tenebrionidae, Tenebrioninae) indicates the need for profound rearrangements of the classification

Helopini is a diverse tribe in the subfamily Tenebrioninae with a worldwide distribution. The New World helopine species have not been reviewed recently and several doubts emerge regarding their generic assignment as well as the naturalness of the tribe and subordinate taxa. To assess these questions, a preliminary cladistic analysis was conducted with emphasis on sampling the genera distributed in the New World, but including representatives from other regions. The parsimony analysis includes 30 ingroup species from America, Europe and Asia of the subtribes Helopina and Cylindrinotina, plus three outgroups, and 67 morphological characters. Construction of the matrix resulted in the discovery of morphological character states not previously reported for the tribe, particularly from the genitalia of New World species. A consensus of the 12 most parsimonious trees supports the monophyly of the tribe based on a unique combination of characters, including one synapomorphy. None of the subtribes or the genera of the New World represented by more than one species (Helops Fabricius, Nautes Pascoe and Tarpela Bates) were recovered as monophyletic. Helopina was recovered as paraphyletic in relation to Cylindrinotina. One Nearctic species of Helops and one Palearctic species of Tarpela (subtribe Helopina) were more closely related to species of Cylindrinotina. A relatively derived clade, mainly composed by Neotropical species, was found; it includes seven species of Tarpela, seven species of Nautes, and three species of Helops, two Nearctic and one Neotropical. Our results reveal the need to deeply re-evaluate the current classification of the tribe and subordinated taxa, but a broader taxon sampling and further character exploration is needed in order to fully recognize monophyletic groups at different taxonomic levels (from subtribes to genera).     

A cladistic analysis of wax scales (Hemiptera: Coccoidea: Coccidae: Ceroplastinae)

Seven genera of wax scales (Ceroplastinae) have been accepted as valid by some or all coecidologists: Ceroplastes Gray, Ceroplastidia Cockerell, Cerostegia De Lotto, Gascardia Targioni Tozzetti, Paracerostegia Tang, Vinsonia Signoret and Waxiella De Lotto. The status of most these genera is controversial. The present study reviews the taxonomic history of wax scales and presents a new classification based on cladistic analysis of morphological data from the adult females of eighty-three species. The analysis indicates that the wax scales form a monophyletic group, that Waxiella is monophyletic but that Ceroplastidia, Gascardia and Vinsonia are not, and fails to support the generic status of Cerostegia and Paracerostegia. Apart from the type species of Vinsonia , which was placed as a sister to the rest of wax scales, the only other species assigned to Vinsonia and most species of all other genera fall within Ceroplastes in the cladogram. Recognition of any of these genera renders Ceroplastes paraphyletic or polyphyletic. Thus, cladistically, the wax scales should be classified into one genus, Ceroplastes.     

Molecular phylogenetics of the family Cyprinidae (Actinopterygii: Cypriniformes) as evidenced by sequence variation in the first intron of S7 ribosomal protein-coding gene: further evidence from a nuclear gene of the systematic chaos in the family

The family Cyprinidae is the largest freshwater fish group in the world, including over 200 genera and 2100 species. The phylogenetic relationships of major clades within this family are simply poorly understood, largely because of the overwhelming diversity of the group; however, several investigators have advanced different hypotheses of relationships that pre- and post-date the use of shared-derived characters as advocated through phylogenetic systematics. As expected, most previous investigations used morphological characters. Recently, mitochondrial DNA (mtDNA) sequences and combined morphological and mtDNA investigations have been used to explore and advance our understanding of species relationships and test monophyletic groupings. Limitations of these studies include limited taxon sampling and a strict reliance upon maternally inherited mtDNA variation. The present study is the first endeavor to recover the phylogenetic relationships of the 12 previously recognized monophyletic subfamilies within the Cyprinidae using newly sequenced nuclear DNA (nDNA) for over 50 species representing members of the different previously hypothesized subfamily and family groupings within the Cyprinidae and from other cypriniform families as outgroup taxa. Hypothesized phylogenetic relationships are constructed using maximum parsimony and Basyesian analyses of 1042 sites, of which 971 sites were variable and 790 were phylogenetically informative. Using other appropriate cypriniform taxa of the families Catostomidae (Myxocyprinus asiaticus), Gyrinocheilidae (Gyrinocheilus aymonieri), and Balitoridae (Nemacheilus sp. and Beaufortia kweichowensis) as outgroups, the Cyprinidae is resolved as a monophyletic group. Within the family the genera Raiamas, Barilius, Danio, and Rasbora, representing many of the tropical cyprinids, represent basal members of the family. All other species can be classified into variably supported and resolved monophyletic lineages, depending upon analysis, that are consistent with or correspond to Barbini and Leuciscini. The Barbini includes taxa traditionally aligned with the subfamily Cyprininae sensu previous morphological revisionary studies by Howes (Barbinae, Labeoninae, Cyprininae and Schizothoracinae). The Leuciscini includes six other subfamilies that are mainly divided into three separate lineages. The relationships among genera and subfamilies are discussed as well as the possible origins of major lineages.     

Phylogeny and historical biogeography of Agabinae diving beetles (Coleoptera) inferred from mitochondrial DNA sequences

The Agabinae, with more than 350 species, is one of the most diverse lineages of diving beetles (Dytiscidae). Using the mitochondrial genes 16S rRNA and cytochrome oxidase I we present a phylogenetic analysis based on 107 species drawn mostly from the four main Holarctic genera. Two of these genera (Ilybius and Ilybiosoma) are consistently recovered as monophyletic with strong support, Platambus is never recovered as monophyletic, and Agabus is found paraphyletic with respect to several of the species groups of Platambus. Basal relationships among the main lineages are poorly defined, although within each of them relationships are in general robust and very consistent across the parameter space, and in agreement with previous morphological analyses. In each of the two most diverse lineages (Ilybius and Agabus including part of Platambus) there is a basal split between Palearctic and Nearctic clades, estimated to have occurred in the late Eocene. The Palearctic clade in turn splits into a Western Palearctic clade and a clade containing mostly Eastern Palearctic species, and assumed to be ancestrally Eastern Palearctic but with numerous transitions to a Holarctic or Nearctic distribution. These results suggest an asymmetry in the colonization routes, as there are very few cases of transcontinental range expansions originating from the Nearctic or the Western Palearctic. According to standard clock estimates, we do not find any transcontinental shift during the Pliocene, but numerous speciation events within each of the continental or subcontinental regions.     

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.     

中国蹄盖蕨属的研究I─属下分类

蹄盖蕨属是蕨类植物中分类最困难的大属之一,中国、喜马拉雅和日本为其分布中心。隶属于本属的种实际上是由蹄盖蕨科或广义蹄盖蕨属中其他属被划分出去以后剩余下来的种类集合而成的“大杂烩”,因此,它们被放在一起并非是一个单系起源的自然类群;属下不同类群分化的程度相差也很大,有的类群进化活跃,近缘种很多,有的类群则很孤立,所以属下单位所包括的种数相差悬殊;孢子周壁表面褶皱的有无在本属的属下分类中占有重要的位置。已发表的中国蹄盖蕨属植物名称约有３００多个,它们初步被处理为１１７种和一些变种和杂种,本文建议把这些种分为１４个组和１５个系。     

The Upper Triassic fishes from Polzberg bei Lunz, Austria

Carnian (Upper Triassic) fishes from Polzberg bei Lunz have been known since 1886 but no comprehensive account has been published. Eleven species are described nine of which, Saurichthys calcaratus, Polzbergia brochatus, Peltopleurus dirumptus, Habroichthys gregarius, Nannolepis elegans, Phaidrosoma lunzensis, Elpistoichthys pectinatus, E. striolatus and Pholidophoretes salvus are new, and two others, Thoracopterus niederristi Bronn and Gigantopterus telleri Abel, previously little-known.
New supraspecific taxa defined are:the order Polzbergiiformes, the family Thoracopteridae and the genera Polzbergia, Nannolepis, Phaidrosoma, Elpistoichthys and Pholidophoretes.
Habroichthys, Thoracopterus, Gigantopterus and Nannolepis show an unusual skull-roof pattern and are included in the re-defined order Luganoiiformes.
Two new ichthyokentemids considerably extend the known time-range of this family.
The genus Pholidophoretes is intermediate between the Archaeomenidae Goodrich 1909, and the Pholidophoridae sensu stricto Nybelin 1966.
The Polzberg assemblage was probably mainly marine with a small freshwater contribution; it shows less similarity to the Besano and Raibl assemblages than these do to each other.
The Luganoiiformes are probably, but not certainly, monophyletic; relationships within the order are analyzed and a cladogram constructed.
The Platysiagiformes, Peltopleuriformes, Luganoiiformes and Cephaloxeniformes could all have been derived from a common ancestor at the Perleidus level and are probably offshoots of the perleidid radiation.     

The history of a nearctic colonization: molecular phylogenetics and biogeography of the Nearctic toads (Bufo)

Previous hypotheses of phylogenetic relationships among Nearctic toads (Bufonidae) and their congeners suggest contradictory biogeographic histories. These hypotheses argue that the Nearctic Bufo are: (1) a polyphyletic assemblage resulting from multiple colonizations from Africa; (2) a paraphyletic assemblage resulting from a single colonization event from South America with subsequent dispersal into Eurasia; or (3) a monophyletic group derived from the Neotropics. We obtained approximately 2.5 kb of mitochondrial DNA sequence data for the 12S, 16S, and intervening valine tRNA gene from 82 individuals representing 56 species and used parametric bootstrapping to test hypotheses of the biogeographic history of the Nearctic Bufo. We find that the Nearctic species of Bufo are monophyletic and nested within a large clade of New World Bufo to the exclusion of Eurasian and African taxa. This suggests that Nearctic Bufo result from a single colonization from the Neotropics. More generally, we demonstrate the utility of parametric bootstrapping for testing alternative biogeographic hypotheses. Through parametric bootstrapping, we refute several previously published biogeographic hypotheses regarding Bufo. These previous studies may have been influenced by homoplasy in osteological characters. Given the Neotropical origin for Nearctic Bufo, we examine current distributional patterns to assess whether the Nearctic-Neotropical boundary is a broad transition zone or a narrow boundary. We also survey fossil and paleogeographic evidence to examine potential Tertiary and Cretaceous dispersal routes, including the Paleocene Isthmian Link, the Antillean and Aves Ridges, and the current Central American Land Bridge, that may have allowed colonization of the Nearctic.     

Molecular evolution of southern North American Cyprinidae (Actinopterygii), with the description of the new genus Tampichthys from central Mexico

Most of the recognized species of the genus Dionda inhabit drainages of the Gulf of Mexico from central Mexico to central Texas, USA, and have been considered a monophyletic group based on morphological, osteological, and allozyme investigations. Phylogenetic relationships of 15 species of Dionda and 34 species from closely related genera were inferred from one mitochondrial (cytb) and three nuclear gene sequences (S7, Rhodopsin, Rag1) totaling 4487 nucleotides. Separate analyses of all four genes yield congruent phylogenies; however the 15 putative species of Dionda evaluated were never recovered as a monophyletic group when species from nine related genera were included in the analyses. Among the ingroup taxa, one well-supported and highly divergent clade is consistently recognized and consists of six recognized and three undescribed northern species currently recognized in the genus Dionda. These nine species inhabit present or past tributaries of the Rio Grande basin of northern Mexico and southern USA, and were recovered as a basal clade in all analyses. Another large, also strongly supported clade, consisting of seven genera, include five southern recognized species currently in the genus Dionda, forming the sister group to the Codoma clade. These five species comprise the "Southern Dionda clade" and inhabit headwaters of the Pánuco-Tamesí drainage and some adjacent coastal rivers in the Tampico Embayment. The consistent and repeated identification of eight different clades recovered in most of the separate gene analyses strongly supports a division of the non-natural genus Dionda. A new genus, Tampichthys, is proposed for the clade of species endemic to east-central Mexico and formerly in Dionda. Tampichthys and the putative monotypic genus Codoma are more related to Mexican species of the genera Cyprinella and Notropis than to other species referred to Dionda sensu stricto.     

中国蹄盖蕨属的研究Ⅰ-属下分类

蹄盖蕨属是蕨类植物中分类最困难的大属之一,中国、喜马拉雅和日本为其分布中心。隶属于本属的种实际上是由蹄盖蕨科或广义蹄盖蕨属中其他属被划分出去以后剩余下来的种类集合而成的"大杂烩",因此,它们被放在一起并非是一个单系起源的自然类群;属下不同类群分化的程度相差也很大,有的类群进化活跃,近缘种很多,有的类群则很孤立,所以属下单位所包括的种数相差悬殊;孢子周壁表面褶皱的有无在本属的属下分类中占有重要的位置。已发表的中国蹄盖蕨属植物名称约有300多个,它们初步被处理为117种和一些变种和杂种,本文建议把这些种分为14个组和15个系。     

Iridoid glucosides substitution patterns inVerbenaceae and their taxonomic implication

The familiar limits ofVerbenaceae are controversial, ranging from a sensu lato arrangement of seven subfamilies until a sharply defined sensu stricto group consisting of only some genera from subfam.Verbenoideae. Iridoid contents of genera belonging toVerbenaceae s. l. were examined through a cladistic analysis, using outgroup analysis to evaluate character polarity within the ingroup. The cladogram obtained shows the distribution of the species according to their iridoid content.     

Theraphosidae phylogeny: relationships of the ‘Ischnocolinae’ genera (Araneae,Mygalomorphae)

The mygalomorph spider subfamily ‘Ischnocolinae’ was originally established as a group based on the presence of divided tarsal scopula. Later, the divided condition of the scopula was considered the plesiomorphic state, which could not support the monophyly of ‘Ischnocolinae’. In Raven 1985, the subfamily was considered paraphyletic, pending a phylogenetic analysis to reinvestigate monophyletic groups. This study comprises such a phylogenetic analysis, based on morphological data, that includes representatives of all genera currently included in ‘Ischnocolinae’ as well as representatives of all other nine Theraphosidae subfamilies (Thrigmopoeinae, Ornithoctoninae, Eumenophorinae, Stromatopelminae, Harpactirinae, Selenogyrinae, Theraphosinae, Aviculariinae and Selenocosmiinae). The family Theraphosidae is considered monophyletic and expanded to include three additional genera previously considered as possible Barychelidae, namely Brachionopus (as Harpactirinae), Trichopelma and Reichlingia (as Ischnocolinae sensu stricto) while ‘Ischnocolinae’ as previously defined does not appear as monophyletic. However, two monophyletic groups were defined as subfamilies to include some former ‘Ischnocolinae’ representatives. The first group includes Acanthopelma rufescens, Trichopelma nitidum, Reichlingia annae, Ischnocolus spp., Holothele rondoni, Holothele culebrae and Holothele aff culebrae and is hereby named as Ischnocolinae (sensu stricto). The other subgroup comprises Sickius longibulbi, Holothele incei, Holothele aff incei, Guyruita spp., Schismatothele lineata, Hemiercus modestus, Holothele colonica and Holothele sp., together established as Schismatothelinae subfam. nov. Several genera included in former ‘Ischnocolinae’ appear as monophyletic (Catumiri, Oligoxystre, Heterothele, Nesiergus, Chaetopelma, Ischnocolus, Guyruita and Plesiophrictus). However, the genera Holothele, Schismatothele and Hemiercus deserve more attention in order to evaluate their intrarelationships and inclusion of species.