The logical priority of the tree over characters and some of its consequences for taxonomy

The aim of the present paper is to explore the role of the character in phylogenetic systematics. I argue that too much emphasis is put on particular characters rather than congruence both in the choice of phylogenetic hypotheses and in taxonomic decisions. This means that the logical priority of the tree over the characters is neglected. To a large extent, this is a result of not paying enough attention to the individuality thesis which states that clades are historical individuals and hence contingent in nature.     

Phylogenetic relationships in the tribe Oxyptilini (Lepidoptera,Pterophoridae, Pterophorinae) based on morphological data of adults

The monophyly of the tribe Oxyptilini and phylogenetic relationships of the genera embraced in this tribe were examined using 171 (75 binary and 96 multistate) characters of adult morphology. The study material included 98 species of 30 genera, representing all previously recognized genera of Oxyptilini, together with the genera Sphenarches, Antarches, Diacrotricha, and Cosmoclostis, four species of Oidaematophorini, three species of Platyptiliini, as well as three and two other species belonging to Pterophorini and Exelastini respectively. Two Agdistis species were used as outgroups. The cladistic analysis resulted in six equally parsimonious trees. A majority of the recovered synapomorphic characters have previously been used in the taxonomy of the subfamily. However, 25 novel characters were found. The monophyly of Oxyptilini was supported, although only with homoplastic characters and low amounts of tree confidence; the genera Capperia, Procapperia, Paracapperia, Oxyptilus, Megalorhipida, and Trichoptilus were found to be nonmonophyletic; Sphenarches and Antarches were recovered as members of Oxyptilini; the two genera Cosmoclostis and Diacrotricha were placed out of Oxyptilini, inside the tribe Pterophorini; and close affinity of the genus Dejongia to Stangeia, Stenodacma, Megalorhipida, Trichoptilus, and Buckleria species was revealed. Four new combinations, Cosmoclostis lanceata (Arenberger) comb. nov. , Nippoptilia regulus (Meyrick) comb. nov. , Capperia tadzhica (Zagulajev) comb. nov. , and Buckleria negotiosus (Meyrick) comb. nov. are proposed; Capperia insomnis Townsend was considered as a senior synonym of Procapperia hackeri Arenberger syn. nov. , Buckleria negotiosus (Meyrick) as a senior synonym of Buckleria vanderwolfi Gielis syn. nov. , and Oxyptilus variegatus Meyrick syn. nov. as a junior synonym of Oxyptilus secutor Meyrick. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 484–547.     

Weintrauboa, a new genus of pimoid spiders from Japan and adjacent islands, with comments on the monophyly and diagnosis of the family Pimoidae and the genus Pimoa (Araneoidea, Araneae)

The spider genus Weintrauboa new genus (Araneae, Pimoidae) is described to place two species of pimoids from Japan and adjacent islands that were formerly classified in the linyphiid genus Labulla . Weintrauboa contortipes (Karsch) new comb., the type species, and W. chikunii (Oi) new comb. are redescribed. Parsimony analysis of morphological characters provides robust support for the monophyly of the genus Weintrauboa and corroborates the monophyly of Pimoa , Pimoidae, and the clade Linyphiidae plus Pimoidae. New diagnoses for Pimoa and Pimoidae are provided.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 139 , 261–281     

Phylogeny of the nominotypical subgenus of Culex (Diptera: Culicidae): insights from analyses of anatomical data into interspecific relationships and species groups in an unresolved tree

The aim of this study was to produce the first objective and comprehensive phylogenetic analysis of the speciose subgenus Culex based on morphological data. We used implied and equally weighted parsimony methods to analyse a dataset comprised of 286 characters of the larval, pupal, and adult stages of 150 species of the subgenus and an outgroup of 17 species. We determined the optimal support by summing the GC supports for each MPC, selecting the cladograms with the highest supports to generate a strict consensus tree. We then collapsed the branches with GC support < 1 to obtain the ‘best’ topography of relationships. The analyses largely failed to resolve relationships among the species and the informal groups in which they are currently placed based on morphological similarities and differences. All analyses, however, support the monophyly of genus Culex. With the exception of the Atriceps Group, the analyses failed to find positive support for any of the informal species groups (monophyly of the Duttoni Group could not be established because only one of the two species of the group was included in the analyses). Since the analyses would seem to include sufficient data for phylogenetic reconstruction, lack of resolution appears to be the result of inadequate or conflicting character data, and perhaps incorrect homology assessments. Molecular and other biological data are needed to gain insights into the evolution of subgenus Culex. Nevertheless, we discuss the placement of several taxa in the current morphology-based classification of the subgenus based on insights realized during the study.

     

A cladistic analysis of Siboglinidae Caullery, 1914 (Polychaeta, Annelida): formerly the phyla Pogonophora and Vestimentifera

It has been proposed in recent years that the phyla Pogonophora and Vestimentifera are a derived clade of polychaete annelids. It has also been proposed that if this clade belongs among polychaetes, then the taxon name Pogonophora is misleading and should revert to a name first formulated for the group, Siboglinidae Caullery, 1914. This recommendation is adopted in this paper, and a cladistic study using terminals of 'generic' rank in the former Pogonophora (including Vestimentifera) is undertaken. The purpose of this is to assess which taxon names should now be used for clades within Siboglinidae, and to provide a revised taxonomy, based on phylogenetic principles. Another major aim is to assess the position of the vestimentiferan clade within Siboglinidae. The results show that Vestimentifera is the sister group to Sclerolinum, and this clade is then sister group to Frenulata, i.e. the remaining Siboglinidae. The results suggest that all taxa within Siboglinidae that are not genera or species are redundant, except for the following: Siboglinidae is defined as the first polychaete, and all its descendants, to have an gut occluded by expanded endoderm filled with chemoautotrophic bacteria, as seen in the holotype of Riftia pachyptila Jones, 1981. Monilifera can be defined based on apomorphy-based system such that it is the first siboglinid, and all its descendants, to have rings of chaetae (uncini) in the opisthosoma, as seen in the holotype of Sclerolinum magdalenae Southward, 1972. Vestimentifera can be denned as the first siboglinid and all its descendants to have a vestimentum as seen in the holotype of Riftia pachyptia. Frenulata is defined as the siboglinid, and all its descendants, to have a mid-trunk girdle, as seen in the holotype of Siboglinum weberi Caullery, 1914. The taxa of generic rank are not defined here since their monophyly was not investigated.     

Monophyly and phylogenetic placement of the spider genus Labulla Simon, 1884 (Araneae, Linyphiidae) and description of the new genus Pecado

The genus Labulla Simon is circumscribed in phylogenetic terms to include the species Labulla thoracica (Wider), L. flahaulti Simon and L. machadoi sp. nov. The genital anatomy of the genus is described in detail and the taxonomy of the genus is reviewed. The monophyly of Labulla is supported by numerous morphological apomorphies of the male palp and female epigynum. Based on a cladistic analysis, a new genus, Pecado gen. nov. , is erected to place Labulla impudica Denis, from Northern Africa. Lepthyphantes insularis Saito and ' Labulla ' nepula Tikader, both formerly included in Labulla , are not congeneric with the type species of Labulla .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 359–404.     

Nanoa, an enigmatic new genus of pimoid spiders from western North America (Pimoidae, Araneae)

The spider genus Nanoa gen. nov. (Araneae, Pimoidae) is described to place Nanoa enana , a new species of pimoids from Western North America. Parsimony analysis of morphological characters provides support for the monophyly of Pimoa plus Nanoa and corroborates the monophyly of Pimoidae and of the clade Linyphiidae plus Pimoidae. © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 249–262.     

Phylogeny and classification of Ochlerotatus and allied taxa (Diptera: Culicidae: Aedini) based on morphological data from all life stages

The phylogenetic relationships and generic assignments of ‘Ochlerotatus’ and related taxa of uncertain taxonomic position in the classification of Aedini previously proposed by the authors in 2004 and 2006 are explored using 297 characters from eggs, fourth‐instar larvae, pupae, adults and immature habitat coded for 158 exemplar species. The ingroup comprises 54 species and the outgroup includes four non‐aedine species and 100 aedine species, 21 of which were previously classified as incertae sedis. Data are analysed in a total‐evidence approach using implied weighting. The analysis produced 158 most parsimonious cladograms. The strict consensus tree (SCT) corroborates the monophyly of the 30 generic‐level taxa recognized previously that are included in the analysis. Overall, the results show remarkable congruence with those obtained previously despite differences in the taxa and morphological characters analysed in this and the two previous studies. All species of Ochlerotatus s.s., subgenus ‘Ochlerotatus’sensu auctorum, Geoskusea, Levua, Pseudoskusea and Rhinoskusea included in the analysis fall within a single clade that is treated as genus Ochlerotatus; thus, the last four taxa are restored to their previous subgeneric rank within this genus. Nine additional subgenera, of which four are new, are proposed for monophyletic clades of Ochlerotatus species based on the strength of character support and application of the principle of equivalent rank. Acartomyia stat. nov. , Culicelsa stat. nov. , Gilesia stat. nov. , Protoculex stat. nov. and Chrysoconops stat. nov. are resurrected from synonymy with Ochlerotatus; and Empihals subgen. nov. (type species: Culex vigilax Skuse), Pholeomyia subgen. nov. (type species: Aedes calcariae Marks), Buvirilia subgen. nov. (type species: Aedes edgari Stone & Rosen) and Sallumia subgen. nov. (type species: Aedes hortator Dyar & Knab) are described as new. The sister group of Ochlerotatus includes a number of species that were previously regarded as incertae sedis in ‘Oc. (Finlaya)’ and ‘Oc. (Protomacleaya)’. Based on previous observations, refined relationships and new character support, three additional genera are recognized for species previously included in ‘Finlaya’, i.e. Danielsia stat. nov . (type species: Danielsia albotaeniata Leicester), Luius gen. nov. (type species: Aedes fengi Edwards) and Hopkinsius gen. nov. (type species: Aedes ingrami Edwards). Additionally, Alloeomyia subgen. nov. (type species: Culex pseudotaeniatus Giles) and Yamada subgen. nov. (type species: Aedes seoulensis Yamada) are introduced as subgenera of Collessius and Hopkinsius, respectively. As is usual with generic‐level groups of Aedini, the newly recognized genera and subgenera are polythetic taxa that are diagnosed by unique combinations of characters. The analysis corroborates the previous observation that ‘Oc. (Protomacleaya)’ is a polyphyletic assemblage of species. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 29–114.     

Phylogeny and classification of tribe Aedini (Diptera: Culicidae)

The phylogeny and classification of tribe Aedini are delineated based on a cladistic analysis of 336 characters from eggs, fourth‐instar larvae, pupae, adult females and males, and immature stage habitat coded for 270 exemplar species, including an outgroup of four species from different non‐aedine genera. Analyses of the data set with all multistate characters treated as unordered under implied weights, implemented by TNT version 1.1, with values of the concavity constant K ranging from 7 to 12 each produced a single most parsimonious cladogram (MPC). The MPCs obtained with K values of 7–9 were identical, and that for K = 10 differed only in small changes in the relationships within one subclade. Because values of K < 7 and > 10 produced large changes in the relationships among the taxa, the stability of relationships exemplified by the MPC obtained from the K = 9 analysis is used to interpret the phylogeny and classification of Aedini. Clade support was assessed using parsimony jackknife and symmetric resampling. Overall, the results reinforce the patterns of relationships obtained previously despite differences in the taxa and characters included in the analyses. With two exceptions, all of the groups represented by two or more species were once again recovered as monophyletic taxa. Thus, the monophyly of the following genera and subgenera is corroborated: Aedes, Albuginosus, Armigeres (and its two subgenera), Ayurakitia, Bothaella, Bruceharrisonius, Christophersiomyia, Collessius (and its two subgenera), Dahliana, Danielsia, Dobrotworskyius, Downsiomyia, Edwardsaedes, Finlaya, Georgecraigius (and its two subgenera), Eretmapodites, Geoskusea, Gilesius, Haemagogus (and its two subgenera), Heizmannia (and subgenus Heizmannia), Hopkinsius (and its two subgenera), Howardina, Hulecoeteomyia, Jarnellius, Kenknightia, Lorrainea, Macleaya, Mucidus (and its two subgenera), Neomelaniconion, Ochlerotatus (subgenera Chrysoconops, Culicelsa, Gilesia, Pholeomyia, Protoculex, Rusticoidus and Pseudoskusea), Opifex, Paraedes, Patmarksia, Phagomyia, Pseudarmigeres, Rhinoskusea, Psorophora (and its three subgenera), Rampamyia, Scutomyia, Stegomyia, Tanakaius, Udaya, Vansomerenis, Verrallina (and subgenera Harbachius and Neomacleaya), Zavortinkius and Zeugnomyia. In addition, the monophyly of Tewarius, newly added to the data set, is confirmed. Heizmannia (Mattinglyia) and Verrallina (Verrallina) were found to be paraphyletic with respect to Heizmannia (Heizmannia) and Verrallina (Neomacleaya), respectively. The analyses were repeated with the 14 characters derived from length measurements treated as ordered. Although somewhat different patterns of relationships among the genera and subgenera were found, all were recovered as monophyletic taxa with the sole exception of Dendroskusea stat. nov. Fifteen additional genera, three of which are new, and 12 additional subgenera, 11 of which are new, are proposed for monophyletic clades, and a few lineages represented by a single species, based on tree topology, the principle of equivalent rank, branch support and the number and nature of the characters that support the branches. Acartomyia stat. nov. , Aedimorphus stat. nov. , Cancraedes stat. nov. , Cornetius stat. nov. , Geoskusea stat. nov. , Levua stat. nov. , Lewnielsenius stat. nov. , Rhinoskusea stat. nov. and Sallumia stat. nov., which were previously recognized as subgenera of various genera, are elevated to generic status. Catageiomyia stat. nov. and Polyleptiomyia stat. nov. are resurrected from synonymy with Aedimorphus, and Catatassomyia stat. nov. and Dendroskusea stat. nov. are resurrected from synonymy with Diceromyia. Bifidistylus gen. nov. (type species: Aedes lamborni Edwards) and Elpeytonius gen. nov. (type species: Ochlerotatus apicoannulatus Edwards) are described as new for species previously included in Aedes (Aedimorphus), and Petermattinglyius gen. nov. (type species: Aedes iyengari Edwards) and Pe. (Aglaonotus) subgen. nov. (type species: Aedes whartoni Mattingly) are described as new for species previously included in Aedes (Diceromyia). Four additional subgenera are recognized for species of Ochlerotatus, including Oc. (Culicada) stat. nov. (type species: Culex canadensis Theobald), Oc. (Juppius) subgen. nov. (type species: Grabhamia caballa Theobald), Oc. (Lepidokeneon) subgen. nov. (type species: Aedes spilotus Marks) and Oc. (Woodius) subgen. nov. (type species: Aedes intrudens Dyar), and seven are proposed for species of Stegomyia: St. (Actinothrix) subgen. nov. (type species: Stegomyia edwardsi Barraud), St. (Bohartius) subgen. nov. (type species: Aedes pandani Stone), St. (Heteraspidion) subgen. nov. (type species: Stegomyia annandalei Theobald), St. (Huangmyia) subgen. nov. (type species: Stegomyia mediopunctata Theobald), St. (Mukwaya) subgen. nov. (type species: Stegomyia simpsoni Theobald), St. (Xyele) subgen. nov. (type species: Stegomyia desmotes Giles) and St. (Zoromorphus) subgen. nov. (type species: Aedes futunae Belkin). Due to the unavailability of specimens for study, many species of Stegomyia are without subgeneric placement. As is usual with generic‐level groups of Aedini, the newly recognized genera and subgenera are polythetic taxa that are diagnosed by unique combinations of characters. The analysis corroborates the previous observation that ‘Oc. (Protomacleaya)’ is a polyphyletic assemblage of species.     

Phylogeny and classification of Aedini (Diptera: Culicidae), based on morphological characters of all life stages

Higher‐level relationships within Aedini, the largest tribe of Culicidae, are explored using morphological characters of eggs, fourth‐instar larvae, pupae, and adult females and males. In total, 172 characters were examined for 119 exemplar species representing the existing 12 genera and 56 subgenera recognized within the tribe. The data for immature and adult stages were analysed separately and in combination using equal (EW) and implied weighting (IW). Since the classification of Aedini is based mainly on adult morphology, we first tested whether adult data alone would support the existing classification. Overall, the results of these analyses did not reflect the generic classification of the tribe. The tribe as a whole was portrayed as a polyphyletic assemblage of Aedes and Ochlerotatus within which eight (EW) or seven (IW) other genera were embedded. Strict consensus trees (SCTs) derived from analyses of the immature stages data were almost completely unresolved. Combining the adult and immature stages data resulted in fewer most parsimonious cladograms (MPCs) and a more resolved SCT than was found when either of the two data subsets was analysed separately. However, the recovered relationships were still unsatisfactory. Except for the additional recovery of Armigeres as a monophyletic genus, the groups recovered in the EW analysis of the combined data were those found in the EW analysis of adult data. The IW analysis of the total data yielded eight MPCs consisting of three sets of two mutually exclusive topologies that occurred in all possible combinations. We carefully studied the different hypotheses of character transformation responsible for each of the alternative patterns of relationship but were unable to select one of the eight MPCs as a preferred cladogram. Overall, the relationships within the SCT of the eight MPCs were a significant improvement over those found by equal weighting. Aedini and all existing genera except Ochlerotatus and Aedes were recovered as monophyletic. Ochlerotatus formed a polyphyletic assemblage basal to Aedes. This group included Haemagogus and Psorophora, and also Opifex in a sister‐group relationship with Oc. (Not.) chathamicus. Aedes was polyphyletic relative to seven other genera, Armigeres, Ayurakitia, Eretmapodites, Heizmannia, Udaya, Verrallina and Zeugnomyia. With the exception of Ae. (Aedimorphus), Oc. (Finlaya), Oc. (Ochlerotatus) and Oc. (Protomacleaya), all subgenera with two or more species included in the analysis were recovered as monophyletic. Rather than leave the generic classification of Aedini in its current chaotic state, we decided a reasonable and conservative compromise classification would be to recognize as genera those groups that are ‘weighting independent’, i.e. those that are common to the results of both the EW and IW analyses of the total data. The SCT of these combined analyses resulted in a topology of 29 clades, each comprising between two and nine taxa, and 30 taxa (including Mansonia) in an unresolved basal polytomy. In addition to ten genera (Armigeres, Ayurakitia, Eretmapodites, Haemagogus, Heizmannia, Opifex, Psorophora, Udaya, Verrallina and Zeugnomyia), generic status is proposed for the following: (i) 32 existing subgenera of Aedes and Ochlerotatus, including nine monobasic subgenera within the basal polytomy, i.e. Ae. (Belkinius), Ae. (Fredwardsius), Ae. (Indusius), Ae. (Isoaedes), Ae. (Leptosomatomyia), Oc. (Abraedes), Oc. (Aztecaedes), Oc. (Gymnometopa) and Oc. (Kompia); (ii) three small subgenera within the basal polytomy that are undoubtedly monophyletic, i.e. Ae. (Huaedes), Ae. (Skusea) and Oc. (Levua), and (iii) another 20 subgenera that fall within the resolved part of the SCT, i.e. Ae. (Aedes), Ae. (Alanstonea), Ae. (Albuginosus), Ae. (Bothaella), Ae. (Christophersiomyia), Ae. (Diceromyia), Ae. (Edwardsaedes), Ae. (Lorrainea), Ae. (Neomelaniconion), Ae. (Paraedes), Ae. (Pseudarmigeres), Ae. (Scutomyia), Ae. (Stegomyia), Oc. (Geoskusea), Oc. (Halaedes), Oc. (Howardina), Oc. (Kenknightia), Oc. (Mucidus), Oc. (Rhinoskusea) and Oc. (Zavortinkius). A clade consisting of Oc. (Fin.) kochi, Oc. (Fin.) poicilius and relatives is raised to generic rank as Finlaya, and Downsiomyia Vargas is reinstated from synonymy with Finlaya as the generic name for the clade comprising Oc. (Fin.) leonis, Oc. (Fin.) niveus and their relatives. Three other species of Finlaya?Oc. (Fin.) chrysolineatus, Oc. (Fin.) geniculatus and Oc. (Fin.) macfarlanei? fall within the basal polytomy and are treated as Oc. (Finlaya) incertae sedis. Ochlerotatus (Ochlerotatus) is divided into three lineages, two of which, Oc. (Och.) atropalpus and Oc. (Och.) muelleri, are part of the basal polytomy. The remaining seven taxa of Oc. (Ochlerotatus) analysed, including the type species, form a reasonably well‐supported group that is regarded as Ochlerotatus s.s. Ochlerotatus (Rusticoidus) is retained as a subgenus within Ochlerotatus s.s. Ochlerotatus (Nothoskusea) is recognized as a subgenus of Opifex based on two unique features that support their sister‐group relationship. A new genus, Tanakaius gen. nov. , is proposed for Oc. (Fin.) togoi and the related species Oc. (Fin.) savoryi. The taxonomic status and generic placement of all currently valid species of Aedini are listed in an appendix. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 142 , 289?368.     

Phylogeny and classification of Muscini (Diptera, Muscidae)

Worldwide in distribution, the tribe Muscini comprises 21 accepted genera and about 350 species. In the present study, a cladistic analysis based upon adult morphological characters is carried out in order to discuss the monophyly of the tribe and its genera, the intergeneric relationships and, in some cases, also the intrageneric relationships. As a result, Muscini is supported as a monophyletic tribe sister-group of Stomoxyini. Except for Morellia Robineau-Desvoidy, Curranosia Paterson, and Eudasyphora Townsend, all the remaining genera are monophyletic. The results are dubious for Polietes Rondani, which was then provisionally kept unchanged. Morellia was broadened to include the Neotropical endemic genera Parapyrellia Townsend, Trichomorellia Stein, and Xenomorellia Malloch. Therefore, a new classification is proposed for Morellia in which it is divided into four subgenera: Morellia s.s. , Parapyrellia , Trichomorellia , and Xenomorellia . Furthermore, the previously proposed subgenus Dasysterna Zimin is given new status as a genus; however, as it is preoccupied by Dasysterna Dejean, the new replacement name Ziminellia nom. nov. is proposed herewith. Eudasyphora was found to be a paraphyletic group relative to Dasyphora Robineau-Desvoidy; both genera are hence synonymized, and Dasyphora is classified in three subgenera: Dasyphora s.s. , Eudasyphora , and Rypellia Malloch. The analysis demonstrated that the traditional classification of Musca Linnaeus into subgenera is artificial and, moreover, that the use of characters from male genitalia could be strongly informative for classifying the genus in phylogeny-supported species groups. Finally, the new classification proposal for Muscini recognizes 18 genera and, furthermore, two undescribed genus-ranked taxa are indicated.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 493–532.     

A Family Level Analysis of Tardigrade Phylogeny

In the present study a character data set suitable for cladistic analysis at the family level was developed. A data matrix consisting of 50 morphological characters from 15 families of tardigrades was analyzed by maximum parsimony. Kinorhynchs, loriciferans, and gastrotrichs were used as outgroups. The results agree with the currently accepted hypothesis that Eutardigrada and Heterotardigrada are distinct monophyletic groups. Among the eutardigrades, Eohypsibiidae was found to be a sister group to Macrobiotidae+Hypsibiidae, while Milnesiidae was the basal eutardigrade family. The basal heterotardigrade family was found to be Oreellidae. Echiniscoideans grouped with some traditional Arthrotardigrada (Renaudarctidae, Coronarctidae+Batillipedidae) suggesting that the arthrotardigrades are not monophyletic. The 18S rRNA gene sequence of Batillipes mirus Richters, 1909 and Calohypsibius schusteri Nelson & McGlothlin, 1996 were obtained and their addition to a previously published dataset supports the monophyly of Heterotardigrada and of Parachela versus Apochela within the Eutardigrada.     

Primary classification and phylogeny of the Polemoniaceae, with comments on molecular cladistics

The system of classification of the Polemoniaceae currently in use was published by Grant in 1959. Much new evidence concerning relationships in the family has been obtained by numerous workers since 1959, and the old system is in need of revision. A revised system down to the genus level, based on conventional and unconventional characters, including molecular evidence, is presented here. Nineteen genera are grouped into eight tribes and two subfamilies. Three new tribes are described: Acanthogilieae, Loeselieae, and Leptodactyloneae. Several genera are transferred to new groups. The phylogeny of the family is discussed in the light of both the older and new evidence. The approach used in constructing both the 1959 and new systems is that of evolutionary systematics. Two recent (1996, 1997) family-wide surveys of cpDNA and rDNA use cladistic methods of analysis to arrive at sets of major groups. Some of this molecular evidence has been adopted for the present revised system. However, much incongruence still exists between the new sets of clades, on the one hand, and the present revised system or the still-viable parts of the 1959 system on the other hand. The incongruences call for an examination and comparison of the contrasting methods of evolutionary systematics and molecular cladistics. A fundamental flaw in the 1996 and 1997 treatments is the attempt to classify plants on the basis of single-gene gene trees.     

木霉菌及其系统分类学研究回顾

从木霉菌(Trichodermaspp.)的分类历史、主要分类系统、分子生物学在木霉菌分类研究中的应用、我国木霉菌分类研究状况等几个方面,对木霉菌的分类研究进展进行了简要回顾。近些年来国际上对木霉菌的分类研究取得了显著进展,陆续发表了许多新种,目前木霉属已报道有3组60种和2个变型。另外,对木霉分类研究中存在的问题也进行了讨论。     

Circularity,evolution, systematics … and circularity

Many have argued strongly that incorporation of evolutionary theory into systematics is dangerously circular, while others have maintained that such an integrated approach increases the accuracy of phylogenetic inference. Here, it is demonstrated that such blanket statements regarding exclusion or inclusion of evolutionary principles in systematics fail to distinguish between two very different types of principles. ‘Phylogeny-neutral’ evolutionary principles are those inferred without any recourse to specific phylogenetic hypotheses (e.g. via developmental genetics, biomechanics). In contrast, ‘phylogeny-dependent’ principles are those which can only be inferred on the basis of specific phylogenetic hypotheses (e.g. character associations detected via ‘comparative methods’). Inclusion of phylogeny-neutral principles in systematic studies as a priori assumptions can be justified, since these principles have (often strong) external empirical support from other spheres of investigation. However, inclusion of phylogeny-dependent principles in systematic studies is circular, since such principles have no external empirical support but are themselves derived from systematic studies. Advocating inclusion or exclusion of all (or as many as possible) evolutionary principles from phylogenetic analysis is therefore misguided. Rather, phylogeny-neutral principles are independently supported and can be included, while phylogeny-dependent principles are unjustified assumptions and should be excluded to avoid circularity. However, integration of complex phylogeny-neutral principles in systematics can create operational problems, even though there are no methodological reasons against their inclusion.     

Fish species – how and why

It is argued, with selected examples from freshwaterfish systematics, that species should be viewed as anexpression of self-perpetuated clustered variation innature, conforming to the phylogenetic speciesconcept. The importance of species lies in thefunctional and structural significance of theirdiagnostic characters. Species can be nested by theircharacters into a tree diagram (phylogeny) orhierarchical alignment structure (classification) ofcharacter distribution, which may be taken to reflectevolution, the unifying theory of organismaldiversification. The phylogenetic species concept,which emphasizes recognition of a pattern ofvariation, describes better than any other proposedconcept the units called species by systematists.Other concepts are based on processes and normally donot permit recognition of particular taxa. Specieshave unique histories, and speciation may proceed bydifferent mechanisms. Whereas it may be postulatedthat speciation entails an irreversible change in thegenetic structure of taxa, recognized by phenotypicexpression and apparently also maintained to a largeextent by selection for a particular phenotype,species recognition must remain independent ofassumptions about species history and spatialdistribution. Species are monophyletic taxa and thespecies category does not differ significantly inphylogenetic regard from other systematic categories.Species as such are not necessarily evolutionaryunits. It is recommended to apply species names withreference to the diagnostic characters of the speciesand to abandon the type specimen described by theInternational Code of Zoological Nomenclature as anomenclatural reference unit.