Phylogeny of Syllidae (Polychaeta) based on morphological data

The phylogeny of Syllidae is assessed in two parsimony analyses of 107 morphological characters. The first analysis included one species of each of the 71 genera of the Syllidae, as well as members of other close families. In the second analysis, 23 poorly known genera were excluded. Character information is based on the examination of available types, additional non-types and newly collected material. Syllidae, except Bollandia Glasby, 1994 is monophyletic. Both analyses supported three of the four traditional subfamilies (Exogoninae, Syllinae and Autolytinae) as monophyletic, whereas Eusyllinae was clearly a polyphyletic group. The genera Anoplosyllis Claparède, 1868, Astreptosyllis Kudenov & Dorsey, 1982 , Streptosyllis Webster & Benedict, 1884, Streptospinigera Kudenov, 1983 and Syllides Örsted, 1845 comprise a well-supported monophyletic group, which we classified as a new subfamily: Anoplosyllinae n. subfam. Our results indicated high levels of homoplasy in the morphological characteristics that traditionally used to differentiate groups, such as the fusion of palps and the presence of nuchal epaulettes. Considering the reproductive modes, schizogamy has appeared twice in the family as the derived condition evolving from epigamy, and Exogoninae may be divided into two monophyletic groups based on the brood system.     

Phylogeny and reproduction in Syllidae (Polychaeta)

The phylogeny of 12 representatives of the four syllid subfamilies is analysed using 44 morphological characters and 96 character states. The analysis is designed to answer two questions: are the four subfamilies monophyletic, and how are the different reproductive strategies related? Syllids show two modes of reproduction, epigamy and schizogamy. Epigamy is mainly constrained to two of the four subfamilies, Eusyllinae and Exogoninae but occurs also in some members of Autolytinae. Schizogamy can be of two types–scissiparity and gemmiparity–and both types are found in Syllinae and Autolytinae. Several modes of brood protection also occur in the family; in some epigamous species the eggs and even juveniles are attached to the main animal, whereas in brooding schizogamous species the stolons care for the young. Brood protection appears in Exogoninae, a few species of Eusyllinae, and some Autolytinae. The resulting tree indicates that members exhibiting brood protection are constrained to one clade, and that it is uncertain whether epigamy or schizogamy is the plesiomorphic reproductive mode. Three of the four subfamilies, Autolytinae, Exogoninae and Syllinae, are supported in the analysis whereas Eusyllinae is found to be polyphyletic.     

Systematics and evolution of syllids (Annelida,Syllidae)

A large, combined phylogenetic analysis (including morphological and molecular data from 18S rDNA, 16S rDNA and cytochrome c oxidase subunit I), with the highest number of species and genera of Syllidae studied to date (213 terminals), is examined. The data were explored with different parameters and optimality criteria (parsimony, likelihood, and bayesian inference). The monophyly of Syllidae and most of the traditional subfamilies is supported. The subfamily Eusyllinae is polyphyletic, as currently delineated, but it is herein reorganized and its diagnosis modified to be a valid group. Additional well supported clades arise. The phylogenetic relationships of the well known and established genera, as well as several enigmatic genera (e.g. Anguillosyllis, Paraopisthosyllis and Parahaplosyllis), the position of which in syllid taxonomy was uncertain or dubious to date, are clarified. The results corroborate previous hypotheses about the evolution of the reproductive and brooding modes. Within Syllinae, the nature of the stolon is phylogenetically informative. The classification of the whole family is revised and discussed on the basis of this phylogenetic hypothesis. © The Willi Hennig Society 2011.     

Autolytinae, Eusyllinae an Exogoninae (Syllidae: Polychaeta) from Coiba National Park, Panama)

This is the second paper on the Syllidae (Polychaeta) from hard substrata of the Coiba National Park (Pacific coast of Panama) (the first paper treated Syllinae). On this second paper nineteen species belonging to eleven genera of the subfamilies Autolytinae, Eusyllinae and Exogoninae (Syllidae: Polychaeta) are reported. The samples were collected by SCUBA diving, either by removing 4 kg blocks of dead coral (Pocillopora spp.) or scraping off 25 x 25 cm quadrats of Telesto multiflora or algae (Dyctiota cf. flavellata, Padina cf. durvillaei and another currently unidentified species). They were collected during four expeditions carried out between 1996 and 1998. Four species are newly reported for the Eastern Pacific: Eusyllis lamelligera Marion & Bobretzky, 1875, Exogone exmouthensis Hartmann-Schr?der, 1980, Proceraea cf. cornuta (Agassiz, 1863) and Sphaerosyllis (S.) magnidentata Perkins, 1981. Ten species are reported for the first time from the Central America Pacific coast: Amblyosyllis cf. granosa Ehlers, 1897 (sensu Westheide 1974), Amblyosyllis speciosa Izuka, 1912, Autolytus multidenticulatus Westheide, 1974, Exogone naidinoides Westheide, 1974, Grubeosyllis concinna (Westheide, 1974), Odontosyllis fulgurans (Audouin & Milne Edward, 1833), Opisthodonta mitchelli Kudenov & Harris, 1995, Pionosyllis articulata Kudenov & Harris, 1995, Sphaerosyllis (S.) californiensis Hartman, 1966 and Syllides cf. reishi Dorsey, 1978. The family Syllidae is represented in the Coiba National Park by 14% of cosmopolitan species, 14% of circumtropical species, 22% of Atlantic and Pacific species and 25% of Pacific species restricted to warm waters. In samples of dead coral the dominant species are S. armillaris (Muller, 1771) and S. gracilis (Campoy, 1982); in samples of telestaceans Grubeosyllis concinna dominates.     

Exogoninae (Polychaeta: Syllidae) from the Mexican Caribbean region with a key for the Gran Caribbean species]

This paper identifies the Exogoninae (Syllidae) from the Mexican Caribbean coasts and includes a key to identify all the species recorded from the Grand Caribbean Sea. The classification of the family and the composition of Exogoninae are briefly examined; the correct names of the subfamilies are Syllinae Grube, 1850, Eusyllinae Malaquin, 1893, Autolytinae Malaquin, 1893 and Exogoninae Langerhans, 1879. Exogoninae includes Anguillosyllis Day, 1963, Brania de Quatrefages, 1866, Braniella Hartman, 1963, Exogone ?rsted, 1845, Exogonella Hartman, 1961, Exogonoides Day, 1963, Parapionosyllis Fauvel, 1923, Psammosyllis Westheide, 1990, Spermosyllis Claparède, 1864, and Sphaerosyllis Claparède, 1863. Pseudexogone Augener, 1922, formerly included in the group, is not a syllid; it belongs to Pilargidae. We collected 814 specimens belonging to 3 genera, 3 subgenera and 13 species as Brania (4), Exogone (4) and Sphaerosyllis (5); five new species are described: Brania russelli n. sp, Brania uebelackerae n. sp, Brania westheidei n. sp., Exogone (Exogone) bondi n. sp. and Exogone (Parexogone) sanmartini n. sp. For each species, selected references, diagnostic features, observations on morphological variability, distribution and illustrations are provided; new species also have an english diagnosis. Most abundant species were B. uebelackerae n. sp. (295), S. taylori Perkins (169), E. (E.) dispar Webster (76), and E. (E.) bondi n sp. (72).     

Phylogeny and evolution of reproductive modes in Autolytinae (Syllidae,Annelida)

The phylogeny of 31 autolytine taxa (Syllidae, Polychaeta, and Annelida) was estimated based on 16S rDNA and 18S rDNA sequences. Outgroups included 12 non-autolytine syllids and four other annelids from related groups. The phylogeny was used to trace the evolution of the various reproductive strategies (i.e., epigamy, anterior and posterior scissiparity, and gemmiparity) within the group, and it will also serve as a basis for a forthcoming revision of autolytine taxonomy. The two genes were analysed both separately and in combination using parsimony, maximum likelihood, and Bayesian inference. Regardless of method used the combined analysis supported a division of Autolytinae into three major clades: one with epigamous Autolytus; a second comprising Autolytus and Myrianida with posterior scissiparity and gemmiparity; and a third containing Proceraea, Procerastea, and Virchowia with anterior scissiparity. The relationship between these three groups is uncertain. Ancestral reproductive states were reconstructed with parsimony and maximum likelihood, and the results unequivocally support epigamy as the plesiomorphic reproductive mode in Syllidae, and that schizogamy in Syllinae and Autolytinae are separate events. The evolution of reproductive traits is ambiguous within Autolytinae, and either of the different reproductive modes could represent the ancestral state.     

A new genus of Syllidae (Polychaeta) from Western Australia

During a study carried out on the subfamily Exogoninae (Syllidae) from Australia, several specimens of a new genus and species were found in samples of dead coral substrate from Western Australia. They have long palps, fused except for a terminal notch, long median and two short lateral antennae, a single pair of short tentacular cirri, and short dorsal cirri, somewhat longer than the parapodial lobes. These characters resemble those of the genus Exogone Örsted, 1845. However, all these appendages are articulated. The chaetae are very similar to those of several species of Syllis Lamarck, 1818, having coarse spines on the margin of compound chaetal blades and truncated dorsal simple chaetae. Furthermore, the pharynx begins in chaetiger 3, posterior to the peristomium, as in many species of the genus Syllis; this condition does not occur in any described species of Exogone. The new genus is provisionally proposed to belong to the subfamily Syllinae, although it has some characters typical of the Exogoninae. Examination under the SEM shows another peculiar feature, the nuchal organs are distinctly laterally located. Within the Syllinae, only Paratyposyllis Hartmann-Schröder, 1962 has a single pair of tentacular cirri, but in that genus, the palps are only basally fused.     

Infrastructural study of photoreceptor organs in the larvae of two Syllidae (Annelida, Polycheta)

The ultrastructure of the larval eyes of two Syllidae: Syllis amica (subfamily: Syllinae) and Odontosyllis ctenostoma (subfamily: Eusyllinae) were studied. These eyes are composed of a pigmented supportive cell and one or two sensory cells. A dense body, corresponding to a secreted lens, can be observed in the ocular cavity formed by the supportive cell. Such a structure present in O. ctenostoma adult eyes has never been described in S. amica adult eyes.     

A phylogeny of ranid frogs (Anura: Ranoidea: Ranidae), based on a simultaneous analysis of morphological and molecular data

During the last two decades, major taxonomic rearrangements were instituted in the anuran family Ranidae. Most of these changes were not based on phylogenetic analysis, and many are controversial. Addressing the phylogeny of Ranidae requires broader taxon sampling within the superfamily Ranoidea, the phylogenetic relationships and higher classification of which are also in a state of flux. No comprehensive attempt has yet been made to reconstruct ranid phylogeny using both morphological and molecular data. In the present contribution, data from 178 organismal characters were collated for 74 exemplar species representing the families Arthroleptidae, Hemisotidae, Hyperoliidae, Mantellidae Microhylidae, Petropedetidae, Rhacophoridae, Sooglossidae, and most subfamilies of Ranidae. These were combined with ～1 kb of DNA sequence from the mitochondrial 12S rDNA and 16S rDNA gene regions in a simultaneous parsimony analysis with direct optimization. Results support the classification of Hemisus with the brevicipitine microhylids, confirm that Arthroleptidae (and its two component subfamilies Astylosterninae and Arthroleptinae) are monophyletic, and advocate the recognition of Leptopelidae. Monophyly of Ranidae is compromised by recognition of Petropedetidae, Rhacophoridae and Mantellidae, which should be recognized as subfamilies of Ranidae at present. Furthermore, Petropedetidae was found to be grossly paraphyletic, comprising three clades which are all considered separate subfamilies of Ranidae, i.e., Petropedetinae, Phrynobatrachinae and Cacosterninae. Three well defined subfamilies of Ranidae were consistently retrieved as monophyletic in a sensitivity analysis, i.e., Tomopterninae, Ptychadeninae and Pyxicephalinae. However, Ptychadeninae and Pyxicephalinae were embedded in Raninae and Dicroglossinae, respectively. Ceratobatrachinae is removed from Dicroglossinae. Dicroglossinae is synonymized with Pyxicephalinae. A new subfamily Strongylopinae is proposed. Raninae should be conservatively treated as a “metataxon” (sensu Ford and Cannatella, 1993 ) until such time as it is fully revised. Tomopterninae is removed from synonymy with Cacosterninae. Morphological synapomorphies are reported for major monophyletic clades retrieved in the simultaneous analysis with equal weights. The present study found that many Old World clades appear to contain both African and Asian taxa, contrary to the findings of some recent biogeographical analyses. This study demonstrates the value of broad taxonomic sampling in ranid phylogeny, and highlights the immense contribution that can be made from detailed morphological data. © The Willi Hennig Society 2005.     

A new genus and species of the Syllidae (Annelida, Polychaeta) from south India

A syllid species new to science is described from intertidal sandy sediments on the cast coast of India, for which a new genus is erected: Psammosyllis aliceae gen. et sp.n. Incorporation into either the Eusyllinae or the Exogoninae is unresolved, because it possesses characters of both of these poorly defined subfamilies.     

Sperm ultrastructure of three Syllinae (Annelida, Phyllodocida) species with considerations on syllid phylogeny and Syllis vittata reproductive biology

Phylogeny of Syllidae is under debate due to new studies based on molecular and morphological data. The noticeable taxonomic diversity of syllids (about 700 listed species) is also mirrored in the array of reproductive strategies as well as in sperm morphology, counting a display of forms already supposed to reflect phylogenetic relationships between the species. The sperm ultrastructure of Syllis gerlachi, S. prolifera and S. vittata is herein presented and compared to the Syllinae species studied previously. Moreover, the egg structure and the gamete allocation within stolons of S. vittata are particularly investigated. Both male germinal cells at different level of maturation and oocytes were found in the same individual of S. vittata, suggesting simultaneous hermaphroditism. The ultrastructural analysis revealed that the observed spermatozoa belong to the ect-aquasperm type resembling those of the similar studied species (Syllis sp., S. pigmentata and S. krohni). Differences in the acrosome structure and nucleus shape are in accordance with a recent phylogenetic reconstruction and suggest a trend in the evolution of spermatozoa in Syllinae toward the development of the apical part. However, further molecular and ultrastructural analyses are needed to support this hypothesis. This is the first record of simultaneous hermaphroditism within Syllinae.     

Combined molecular and morphological phylogeny of Eulophidae (Hymenoptera: Chalcidoidea), with focus on the subfamily Entedoninae

A new combined molecular and morphological phylogeny of the Eulophidae is presented with special reference to the subfamily Entedoninae. We examined 28S D2–D5 and CO1 gene regions with parsimony and partitioned Bayesian analyses, and examined the impact of a small set of historically recognized morphological characters on combined analyses. Eulophidae was strongly supported as monophyletic only after exclusion of the enigmatic genus Trisecodes. The subfamilies Eulophinae, Entiinae (=Euderinae) and Tetrastichinae were consistently supported as monophyletic, but Entedoninae was monophyletic only in combined analyses. Six contiguous bases in the 3e′ subregion of the 28S D2 rDNA contributed to placement of nominal subgenus of Closterocerus outside Entedoninae. In all cases, Euderomphalini was excluded from Entiinae, and we suggest that it be retained in Entedoninae. Opheliminae n. stat. is raised from tribe to subfamily status. Trisecodes is removed from Entedoninae but retained as incertae sedis in Eulophidae until its family placement can be determined new placement . The genera Neochrysocharis stat. rev. and Asecodes stat. rev. are removed from synonymy with Closterocerus because strong molecular differences corroborate their morphological differences. Closterocerus (Achrysocharis) germanicus is transferred to the genus Chrysonotomyia n. comb. based on molecular and morphological characters.     

Phylogenetic analysis of the minute brown scavenger beetles (Coleoptera: Latridiidae), and recognition of a new beetle family,Akalyptoischiidae fam.n. (Coleoptera: Cucujoidea)

We infer the first phylogenetic hypothesis for Latridiidae Erichson (Coleoptera: Cucujoidea). Portions of seven genes (18S ribosomal DNA, 28S ribosomal DNA, 12S ribosomal DNA, 16S ribosomal DNA, cytochrome c oxidase I and II and histone III) were analysed. Twenty‐seven latridiid species were included, representing both subfamilies and more than half of the currently recognized genera. Eight outgroup taxa from other families of Cucujoidea were included. Parsimony and partitioned Bayesian analyses were performed on the combined dataset. In both phylogenetic analyses, the enigmatic Akalyptoischion Andrews (Latridiinae) was recovered outside of Latridiidae. The subfamilies Corticariinae and Latridiinae (without Akalyptoischion) were each recovered as monophyletic in both analyses. A new family, Akalyptoischiidae fam.n. is erected based on the results of the phylogenetic study and further support from adult morphology, key features of which are illustrated.     

Family‐level relationships of the spittlebugs and froghoppers (Hemiptera: Cicadomorpha: Cercopoidea)

The spittlebug superfamily Cercopoidea (Hemiptera: Cicadomorpha) comprises approximately 3000 phytophagous species (including some economically important pests of grass crops) classified among the families Cercopidae, Aphrophoridae, Epipygidae, Clastopteridae and Machaerotidae. However, the monophyly of these taxa has never been tested and the evolutionary relationships among these major lineages are unknown. Presented here are the results of the first ever phylogenetic investigation of the higher‐level relationships within Cercopoidea, based on DNA nucleotide sequence data from six loci (18S rDNA, 28S rDNA, histone 3, wingless, cytochrome oxidase I and cytochrome oxidase II) generated from exemplars of 109 spittlebug species representing all five described families, seven of eight subfamilies and 61 genera (eight additional exemplars, representing a selection of other Auchenorrhyncha taxa, were included as outgroups). The resulting topologies are used to evaluate the monophyly of each cercopoid family, and further to calculate divergence date estimates to examine the chronological origins and historical diversification of Cercopoidea. The results of this investigation suggest that: (i) four of the five described families are monophyletic; Epipygidae was recovered consistently as originating within Aphrophoridae; (ii) the exclusively Old World Machaerotidae is the most anciently diversified family of extant spittlebugs; (iii) New World Cercopidae (i.e. Ischnorhininae) constitute a derived monophyletic lineage; (iv) the genus Microsargane Fowler, classified currently within Aphrophoridae, actually belongs within Cercopidae; and (v) the origins of the major spittlebug lineages probably coincided with the breakup of Pangaea and, subsequently, Gondwana, as well as major floristic diversification such as the rise of angiosperms.     

Is a robust phylogeny of the enterobacterial plant pathogens attainable?

Sequences from gapA, gyrA and ompA were used to evaluate the relationships of the enterobacterial plant pathogens, and assess whether a robust phylogeny can be ascertained using this group of housekeeping genes. Up to 48 taxa were included in a combined phylogenetic analysis to explore the evolutionary distribution of plant pathogenic species across the family Enterobacteriaceae. Phylogenies were reconstructed from gapA, gyrA and ompA gene sequences using maximum parsimony and maximum likelihood algorithms, and phylogenetic congruence was evaluated by the incongruence length difference test and the partition addition bootstrap alteration approach. The resulting gene trees were found to be incongruent, with gapA supporting a monophyletic origin for the plant pathogenic species. In contrast, gyrA and ompA supported multiple polyphyletic origins of Erwinia, Brenneria, Pectobacterium and Pantoea in conjunction with a previously published 16S rDNA phylogeny. However, none of the trees (not even the published 16S rDNA gene tree) supports the current taxonomic classification of these genera into four clades, with Pantoea forming the only monophyletic group in the gapA, gyrA and 16S rDNA trees. Finally, the gapA, gyrA and previously published 16S rDNA phylogenies differ in the taxonomic placement of several bacterial strains which are separated in the three trees. The observed incongruence among the four gene histories is likely to be the result of horizontal transfer events, confounding the search for a robust set of housekeeping genes with a shared evolutionary history that could be used to confidently characterize the relationships of the plant pathogenic enterobacteria. © The Willi Hennig Society 2010.     

A phylogenetic analysis of relationships among genera of Conopidae (Diptera) based on molecular and morphological data

Members of the family Conopidae (Diptera) have been the focus of little targeted phylogenetic research. The most comprehensive test of phylogenetic support for the present subfamily classification of Conopidae is presented here using 66 specimens, including 59 species of Conopidae and seven outgroup taxa. Relationships among subfamily clades are also explored. A total of 6824 bp of DNA sequence data from five gene regions (12S ribosomal DNA, cytochrome c oxidase subunit I, cytochrome b, 28S ribosomal DNA and alanyl‐tRNA synthetase) are combined with 111 morphological characters in a combined analysis using both parsimony and Bayesian methods. Parsimony analysis recovers three shortest trees. Bayesian analysis recovers a nearly identical tree. Five monophyletic subfamilies of Conopidae are recovered. The rarely acknowledged Zodioninae is restored, including the genera Zodion and Parazodion. The genus Sicus is removed from Myopinae. Morphological synapomorphies are discussed for each subfamily and inter‐subfamily clade, including a comprehensive review of the character interpretaions of previous authors. Included are detailed comparative illustrations of male and female genitalia of representatives of all five subfamilies with new morphological interpretation.     

Morphology-based phylogeny of the treehopper family Membracidae (Hemiptera: Cicadomorpha: Membracoidea)

A parsimony‐based phylogenetic analysis of eighty‐three morphological characters of adults and immatures of seventy representatives of the tribes and subfamilies of Membracidae and two outgroup taxa was conducted to evaluate the status and relationships of these taxa. Centrotinae apparently gave rise to Nessorhinini and Oxyrhachini (both formerly treated as subfamilies, now syn.n. and syn.reinst., respectively, of Centrotinae). In contrast to previous analyses, a clade comprising Nicomiinae, Centronodinae, Centrodontinae, and the unplaced genera Holdgatiella Evans, Euwalkeria Goding and Antillotolania Ramos was recovered, but relationships within this clade were not well resolved. Nodonica bispinigera, gen.n. and sp.n., is described and placed in Centrodontini based on its sister‐group relationship to a clade comprising previously described genera of this tribe. Membracinae and Heteronotinae were consistently monophyletic. Neither Darninae nor Smiliinae, as previously defined, was monophyletic on the maximally parsimonious cladograms, but constraining both as monophyletic groups required only one additional step. The monophyly of Stegaspidinae, including Deiroderes Ramos (unplaced in Membracidae), was supported on some but not all equally parsimonious cladograms. More detailed analyses of individual subfamilies, as well as morphological data on the undescribed immatures of several membracid tribes and genera, will be needed to elucidate relationships among tribes and genera. A key to the subfamilies and tribes is provided.     

Searching for natural lineages within the Cerylonid Series (Coleoptera: Cucujoidea)

Phylogenetic relationships within the diverse beetle superfamily Cucujoidea are poorly known. The Cerylonid Series (C.S.) is the largest of all proposed superfamilial cucujoid groups, comprising eight families and representing most of the known cucujoid species diversity. The monophyly of the C.S., however, has never been formally tested and the higher-level relationships among and within the constituent families remain equivocal. Here we present a phylogenetic study based on 18S and 28S rDNA for 16 outgroup taxa and 61 C.S. ingroup taxa, representing seven of the eight C.S. families and 20 of 39 subfamilies. We test the monophyly of the C.S., investigate the relationships among the C.S. families, and test the monophyly of the constituent families and subfamilies. Phylogenetic reconstruction of the combined data was achieved via standard static alignment parsimony analyses, Direct Optimization using parsimony, and partitioned Bayesian analysis. All three analyses support the paraphyly of Cucujoidea with respect to Tenebrionoidea and confirm the monophyly of the C.S. The C.S. families Bothrideridae, Cerylonidae, Discolomatidae, Coccinellidae and Corylophidae are supported as monophyletic in all analyses. Only the Bayesian analysis recovers a monophyletic Latridiidae. Endomychidae is recovered as polyphyletic in all analyses. Of the 14 subfamilies with multiple terminals in this study, 11 were supported as monophyletic. The corylophid subfamily Corylophinae and the coccinellid subfamilies Chilocorinae and Scymninae are recovered as paraphyletic. A sister grouping of Anamorphinae+Corylophidae is supported in all analyses. Other taxonomic implications are discussed in light of our results.     

Molecular phylogenetics of Erebidae (Lepidoptera,Noctuoidea)

As a step towards understanding the higher‐level phylogeny and evolutionary affinities of quadrifid noctuoid moths, we have undertaken the first large‐scale molecular phylogenetic analysis of the moth family Erebidae, including almost all subfamilies, as well as most tribes and subtribes. DNA sequence data for one mitochondrial gene (COI) and seven nuclear genes (EF‐1α, wingless, RpS5, IDH, MDH, GAPDH and CAD) were analysed for a total of 237 taxa, principally type genera of higher taxa. Data matrices (6407 bp in total) were analysed by parsimony with equal weighting and model‐based evolutionary methods (maximum likelihood), which revealed a well‐resolved skeleton phylogenetic hypothesis with 18 major lineages, which we treat here as subfamilies of Erebidae. We thus present a new phylogeny for Erebidae consisting of 18 moderate to strongly supported subfamilies: Scoliopteryginae, Rivulinae, Anobinae, Hypeninae, Lymantriinae, Pangraptinae, Herminiinae, Aganainae, Arctiinae, Calpinae, Hypocalinae, Eulepidotinae, Toxocampinae, Tinoliinae, Scolecocampinae, Hypenodinae, Boletobiinae and Erebinae. Where possible, each monophyletic lineage is diagnosed by autapomorphic morphological character states, and within each subfamily, monophyletic tribes and subtribes can be circumscribed, most of which can also be diagnosed by morphological apomorphies. All additional taxa sampled fell within one of the four previously recognized quadrifid families (mostly into Erebidae), which are now found to include two unusual monobasic taxa from New Guinea: Cocytiinae (now in Erebidae: Erebinae) and Eucocytiinae (now in Noctuidae: Pantheinae).