Evolution of the family Lumbriculidae

The structure of the male ducts of numerous representatives of the family Lumbriculidae testifies against the assumption of Stephenson (1930) of an archaism of this family and against the classification of the Oligochaeta based on this fallacy in the papers of Yamaguchi (1953) and Brinkhurst (1971). The genus Dorydrilus Piguet is confirmed as a member of the family Lumbriculidae.     

Phylogenetic systematics and character evolution in the angiosperm family Haloragaceae

The poorly known Haloragaceae R. Br. (Saxifragales) are highly diverse in habit (small trees to submerged aquatics) and labile in floral merosity (2-4), both uncommon among the core eudicots. This family has a cosmopolitan distribution, but taxonomic diversity is concentrated in Australia. An explicit phylogenetic approach has not previously been utilized to examine relationships or character evolution in this family. We used molecular evidence from nrDNA ITS and cpDNA trnK and matK regions under both Bayesian and parsimony analyses to address phylogenetic relationships. Combined molecular analyses defined a monophyletic Haloragaceae with the woody genera (Haloragodendron, Glischrocaryon) sister to the rest. Relationships among many genera were well resolved, with genera as currently delimited generally well supported, although there were notable exceptions; a new genus (Trihaloragis) is recognized, and the aquatic genus Meionectes is again distinct from Haloragis. Three new species combinations are also recognized. There are multiple (two or three) origins of the submerged aquatic habit in the family and potentially an intermediate reversal to the terrestrial habit, neither previously demonstrated in a core eudicot family using an explicit phylogenetic hypothesis. Ancestral character analyses suggest two origins of trimerous flowers and multiple reductions to dimerous flowers throughout Haloragaceae.     

Two atavistic characters of some Lumbriculidae and their importance for the classification of Oligochaeta

The rudimentary atria, and the posterior sperm funnels and sperm ducts, which occur in some species of the Lumbriculidae, are discussed. It is shown that the posterior location of funnels and sperm ducts is the result of a forward shift of the atria, which refutes Stephenson's supposition that the Lumbriculidae is the most archaic family of the present-day Oligochaeta.     

The family classification of the Anoplura

Fifteen families of Anoplura are recognized and defined, one with two sub-families, and a key is provided for their identification. The included genera are listed for each family, together with the relevant type-species as well as the mammalian hosts. Phylogenetic relationships between the families are discussed, and an extensive historical review and analysis of the available taxonomic characters is presented.     

18S rDNA phylogeny of the Tubificidae (Clitellata) and its constituent taxa: dismissal of the Naididae

The phylogeny of the Tubificidae, and of most of its subfamilies and some of its genera, is revisited, on the basis of sequences of 18S ribosomal DNA in a selection of species. Forty-six new 18S sequences of Naididae (6), Tubificidae (37), Phreodrilidae (1), Lumbriculidae (1), and Enchytraeidae (1) are reported and aligned together with corresponding sequences of 21 previously studied taxa. The 18S gene of Insulodrilus bifidus provides the first molecular evidence that phreodrilids are closely related to tubificids, corroborating previous conclusions based on morphology. The data further support the monophyletic status of Tubificidae, provided that the "Naididae" is regarded a part of this family; "naidids" may not even constitute a monophyletic group. It is thus suggested that the family name Naididae is formally suppressed as a junior synonym of the Tubificidae. The 18S gene also resolves a number of relationships within the tubificids. Among the subfamilies, Tubificinae is supported, Rhyacodrilinae and Phallodrilinae are revealed as nonmonophyletic, and Limnodriloidinae remains unresolved. Most tubificid genera tested for monophyly are corroborated by the data, only one (Tubifex) is refuted, and two (Tubificoides and Limnodriloides) are unresolved from other taxa. It is concluded that it will be valuable to expand the taxonomic sampling for 18S rDNA in clitellates, and in annelids in general, as this is likely to improve the resolution at many levels. However, it will be equally important to combine the annelid 18S data with other gene sequences and nonmolecular characters, to estimate the phylogeny of these common and diverse worms with greater precision.     

Toward a global phylogeny of the Brassicaceae

The Brassicaceae is a large plant family (338 genera and 3,700 species) of major scientific and economic importance. The taxonomy of this group has been plagued by convergent evolution in nearly every morphological feature used to define tribes and genera. Phylogenetic analysis of 746 nrDNA internal transcribed spacer (ITS) sequences, representing 24 of the 25 currently recognized tribes, 146 genera, and 461 species of Brassicaceae, produced the most comprehensive, single-locus-based phylogenetic analysis of the family published to date. Novel approaches to nrDNA ITS analysis and extensive taxonomic sampling offered a test of monophyly for a large complement of the currently recognized tribes and genera of Brassicaceae. In the most comprehensive analysis, tribes Alysseae, Anchonieae plus Hesperideae, Boechereae, Cardamineae, Eutremeae, Halimolobeae, Iberideae, Noccaeeae, Physarieae, Schizopetaleae, Smelowskieae, and Thlaspideae were all monophyletic. Several broadly defined genera (e.g., Draba and Smelowskia) were supported as monophyletic, whereas others (e.g., Sisymbrium and Alyssum) were clearly polyphyletic. Analyses of ITS data identified several problematic sequences attributable to errors in sample identification or database submission. Results from parsimony ratchet and Bayesian analyses recovered little support for the backbone of the phylogeny, suggesting that many lineages of Brassicaceae have undergone rapid radiations that may ultimately be difficult to resolve with any single locus. However, the development of a preliminary supermatrix including the combination of 10 loci for 65 species provides an initial estimate of intertribal relations and suggests that broad application of such a method will provide greater understanding of relationships in the family.     

Floral structure and evolution in the Anacardiaceae

WANNAN, B. S. & QUINN, C. J, 1991. Floral structure and evolution in the Anacardiaceae. Carpel morphology and anatomy is investigated in 17 genera and carpellode morphology in 12 genera. There is an evolutionary sequence in the family from poorly differentiated, nearly apocarpous gynoecia towards syncarpous gynoecia with clearly defined stigmata, styles and ovaries. There has also been marked reduction culminating in pseudomonomery. The carpellodes of the male flowers appear more conservative, and provide evidence of affinities between genera with quite different fertile gynoecia. The characters have been polarized using Burseraceae as a sister group. Data from these sources, as well as from pericarp anatomy, wood anatomy and biflavonoid content indicate that the long standing intrafamilial classification into five tribes is artificial, and that the two small satellite families, Blepharocaryaceae and Julianiaceae should be included in the family. A large monophyletic group is recognized comprised of essentially four of the existing tribes (Anacardiëae, Dobineëae, Semecarpeae, Rhoëae), as well as the two satellite families. This group incorporates two subgroups of more closely allied genera. The remaining genera (mostly Spondiadeae) are very diverse, and for the present are placed in an artificial group characterised by a set of plesiomorphs. Relationships within this group must be resolved before a satisfactory taxonomy of the family can be achieved.     

Molecular phylogenetic study of the systematic position of Baikalian oligochaetes in Clitellata

Phylogenetic relationships within the class Clitellata were examined using partial nucleotide sequences of the nuclear 18S ribosomal RNA gene. In the analysis, the already determined sequences for the individual species representing the class members Hirudinea, Acanthobdellida, Branchiobdellida, and Oligochaeta were included. Furthermore, newly determined sequences of the thirteen representatives of the family Lumbriculidae, including 12 Baikalian endemic species, were analyzed. The hypothesis on the close relatedness of these four groups of Clitellata was supported. Leeches, branchiobdellids, and lumbriculids form three independent parallel branches of evolution. These results were consistent with the hypothesis on the role of the family Lumbriculidae as a connecting link, or the transition form between the parasitic and free-living groups of Clitellata. At the same time, these data refute the suggestion that Lumbriculidae could be the ancestral lineage of other Oligochaeta. Moreover, polymorphic group of Baikalian lumbriculids clustered independently from the other representatives of the family, pointing to the uniqueness of the Baikalian fauna of oligochaetes, which was formed within relatively closed system of this ancient lake.     

Molecular phylogenetic study of the systematic position of Baikalian oligochaetes in the system Clitellata

Phylogenetic relationships within the class Clitellata were examined using nucleotide sequences of nuclear 18S ribosomal RNA gene fragments. In the analysis, the already determined sequences for the individual species representing the class members Hirudinea, Acanthobdellida, Branchiobdellida, and Oligochaeta were included. Furthermore, newly determined sequences of the thirteen representatives of the family Lumbriculidae. including 12 Baikalian endemic species, were analyzed. The hypothesis on the close relatedness of these four groups of Clitellata was supported. Leeches, branchiobdellidans, and lumbriculides form three independent parallel branches of evolution. These results were consistent with the hypothesis on the role of the family Lumbriculidae as a connecting link, or the transition form between the parasitic and free-living groups of Clitellata. At the same time, these data refute the suggestion that Lumbriculidae could be the ancestral lineage of other Oligochaeta. Moreover, polymorphic group of Baikalian lumbriculides clustered independently from the other representatives of the family, pointing to the uniqueness of the Baikalian fauna of oligochaetes, which was formed within relatively closed system of this ancient lake.     

Molecular systematics of the Cactaceae

Bayesian, maximum‐likelihood, and maximum‐parsimony phylogenies, constructed using nucleotide sequences from the plastid gene region trnK‐matK, are employed to investigate relationships within the Cactaceae. These phylogenies sample 666 plants representing 532 of the 1438 species recognized in the family. All four subfamilies, all nine tribes, and 69% of currently recognized genera of Cactaceae are sampled. We found strong support for three of the four currently recognized subfamilies, although relationships between subfamilies were not well defined. Major clades recovered within the largest subfamilies, Opuntioideae and Cactoideae, are reviewed; only three of the nine currently accepted tribes delimited within these subfamilies, the Cacteae, Rhipsalideae, and Opuntieae, are monophyletic, although the Opuntieae were recovered in only the Bayesian and maximum‐likelihood analyses, not in the maximum‐parsimony analysis, and more data are needed to reveal the status of the Cylindropuntieae, which may yet be monophyletic. Of the 42 genera with more than one exemplar in our study, only 17 were monophyletic; 14 of these genera were from subfamily Cactoideae and three from subfamily Opuntioideae. We present a synopsis of the status of the currently recognized genera.
© The Willi Hennig Society 2011.     

Multiple origin of the tropical forest tree family Icacinaceae

Analyses of DNA sequences from four genes (ndhF, rbcL, atpB, and 18S rDNA) and morphological data show that the members of the tropical forest tree family Icacinaceae do not have a common origin. All of the genera earlier placed in Icacinaceae are euasterids but placed in the following three different orders: Garryales, Aquifoliales, and Apiales. Icacina and related genera are members of Garryales and, pending more data, are still best treated as Icacinaceae (sensu stricto). The genera related to Aquifoliales are placed in Cardiopteridaceae and a new family, Stemonuraceae. The genus Pennantia is a member of Apiales and the family Pennantiaceae is recognized. Morphological characters delimiting these groups are discussed. Twenty-six new ndhF sequences were obtained for the study (25 from former Icacinaceae and 1 from Cardiopteris).     

A comprehensive generic-level phylogeny of the sunflower family: Implications for the systematics of Chinese Asteraceae

The sunflower family (Asteraceae) is the largest and the most diverse flowering plant family, comprising 24 000–30 000 species and 1600–1700 genera. In China, Asteraceae are also the largest family, with approximately 2336 indigenous species in 248 genera. In the past two decades, molecular phylogenetic analyses has contributed greatly to our understanding of the systematics of Asteraceae. Nevertheless, the large-scale analyses and knowledge about the relationships of Chinese Asteraceae at the generic level as a whole are far from complete due to difficulties in sampling. In this study, we presented a three-marker (rbcL, ndhF, and matK) phylogeny of Asteraceae, including 506 genera (i.e., approximately one-third of Asteraceae genera). The study sampled 200 Chinese genera (i.e., approximately 80% of Chinese Asteraceae genera). The backbones of the new phylogeny were largely congruent with earlier studies, with 13 subfamilies and 45 tribes recognized. Chinese Asteraceae were distributed in 7 subfamilies (Mutisioideae, Wunderlichioideae, Carduoideae, Pertyoideae, Gymnarrhenoideae, Cichorioideae, and Asteroideae) and 22 tribes (Mutiseae, Hyalideae, Cardueae, Pertyeae, Gymnarrheneae, Vernonieae, Cichorieae, Doroniceae, Senecioneae, Astereae, Anthemideae, Gnaphalieae, Calenduleae, Inuleae, Athroismeae, Helenieae, Coreopsideae, Neurolaeneae, Tageteae, Millieae, Eupatorieae, and Heliantheae). Chinese Asteraceae lacked 6 basal subfamilies and 23 tribes. Several previously ambiguous relationships were clarified. Our analyses also resolved some unplaced genera within Chinese Asteraceae. Finally, our phylogenetic tree was used to revise the classification for all genera of Chinese Asteraceae. In total, 255 genera, 22 tribes, and 7 subfamilies in China are recognized.     

Studies on the Lumbriculidae (Oligochaeta) in Britain

Seven species of Lumbriculidae, a family of fresh water Oligochaeta, found in Britain are described and a key is erected for their identification. Two species of Trichodrilus , separated by Beddard are redescribed, resulting in a new synonymy. Trichodrilus Hrabêi sp. n. from Denbighshire is denned. Dorydrilus michaeheni is recorded for the first time from Great Britain.     

Revision of the amphipod (Crustacea) family Stegocephalidae

The amphipod (Crustacea) family Stegocephalidae Dana, 1852 is revised, and the results of a phylogenetic analysis of the family are presented. The morphological information, obtained mainly through direct examination of the species, has been transformed into 200 characters. 91 stegocephalid species (91% of all recognized species) are included in the analysis, in addition to six outgroup taxa. Based upon this analysis, the family is divided into five subfamilies and 26 genera. Four new subfamilies (Andaniexinae, Andaniopsinae, Bathystegocephalinae & Parandaniinae) and ten new genera (Alania, Austrocephaloides, Austrophippsia, Bouscephalus, Gordania, Medi-terexis, Pseudo, Schellenbergia, Stegonomadia & Stegomorphia) are erected. Five genera are put into synonymy (Andaniella with Andaniopsis; Phippsiella and Stegocephalopsis with Stegocephalus; Stegophippsiella with Stego-cephalina; Euandania with Parandania).     

Phylogenetic relationships and the temporal context for the diversification of African characins of the family Alestidae (Ostariophysi: Characiformes): evidence from DNA sequence data

Phylogenetic relationships within the family Alestidae were investigated using parsimony, maximum likelihood, and Bayesian approaches based on a molecular dataset that included both nuclear and mitochondrial markers. Multiple representatives of all but two of the recognized alestid genera were included, which allowed for testing previous hypotheses of intergeneric relationships and the monophyly of several genera. The phylogenetic position of the Neotropical genus Chalceus with respect to the family Alestidae was also examined. In order to understand the temporal context of alestid diversification, Bayesian methods of divergence time estimation using fossil data in the form of calibration priors were used to date the nodes of the phylogenetic tree. Our results rejected the monophyly of the family as currently recognized (Alestidae sensu lato) and revealed several instances of poly- and paraphyly among genera. The genus Chalceus was recovered well nested within Neotropical characiforms, thus rejecting the hypothesis that this taxon is the most basal alestid. The estimated mean divergence time for the alestid clade (Alestidae sensu stricto) was 54 Mya with a 95% credibility interval of 63-49 Mya. These results are incongruent with the hypothesis that the origin of the family Alestidae predates the African-South American Drift-Vicariance event.     

Phylogenetic classification of the family Heteroderidae (Nematoda: Tylenchida)

Summary The family Heteroderidae is revised. On the basis of shared, derived characters sister groups are established and arranged in a phylogenetic tree. A hypothetical, primitive ancestor for the family is defined. The genus Verutus has a large equatorial vulval slit and is considered to be the most primitive form. The genus Meloidodera developed by a reduction in vulval size. Genera which developed later exhibit a subterminally located vulval slit and progressively lost the annulation of the female cuticle. In this process four evolutionary lines emerge: (i) a posterior shift of the vulva and the formation of more or less distinct vulval lips gave rise to the genera Zelandodera and Cryphodera; (ii) changes in the lip configuration of the second-stage juvenile gave rise to the genera Hylonema, Afrodera n.g., Heterodera and Bidera; (iii) changes in the composition of the female cuticle resulted in the genera Thecavermiculatus, Atalodera, Sherodera, Sarisodera and Bellodera n.g. and; (iv) a reduction in vulval slit size led to the development of the genera Dolichodera, Globodera, Cactodera and Punctodera. The genera Ephippiodera and Rhizonema are synonymized with Bidera and Sarisodera respectively. Verutus and Meloidodera are recognized as subfamilies Verutinae and Meloidoderinae and the genera in the four evolutionary lines are recognized as subfamilies Cryphoderinae, Heteroderinae, Ataloderinae and Punctoderinae respectively.Two new genera, Afrodera and Bellodera, are erected for species originally described in Sarisodera and Cryphodera. Both new genera are characterized by a depressed vulval slit and the anus located on the dorsal side of the vulval cone. Differences in lip configuration of the infective juvenile and a postulated difference in female cuticle justifies their placement in different subfamilies. The lip configuration of the infective juveniles in the subfamilies Verutinae, Meloidoderinae, Cryphoderinae, Ataloderinae and Punctoderinae remains basically unchanged. The possible development of this character in the subfamily Heteroderinae is discussed and illustrated. The family Heteroderidae, its six subfamilies and 17 genera are defined or redefined, and for each of the genera the nominal species and their synonyms are listed. New synonyms introduced are: Heterodera rumicis and H. scleranthi of H. trifolii, H. ustinova of Bidera avenae and H. mali of Globodera chaubattia. Cactodera acnidae (Schuster & Brezina, 1979) n. comb. and Dolichodera andinus (Golden, Franco, Jatala & Astogaza, 1983) n. comb. are transferred from Heterodera and Thecavermiculatus respectively. Keys are provided for all taxa for which no suitable keys are available in the literature. Species inquirendae are listed. ac]19840606     

A numerical study of the Australian earthworm genera Cryptodrilus Fletcher and Trinephrus Beddard, with a new genus (Megascolecidae:Oligochaeta)

Species of Cryptodrilus and Trinephrus , South-east Australian genera previously suppressed as synonyms of Notoscolex , are redescribed. The affinities of the species of the Cryptodrilus-Trinephrus complex, both within the complex and with other groupings, are determined from considerations of general anatomy complemented by a taxonometric comparison of setal intervals in the species studied. It is concluded that Cryptodrilus must be re-established as an independent genus to include, in addition to the type-species (C. rusticus) , all species of Eastern Australia and Tasmania previously referred to Trinephrus. A key to all constituent species is provided and three species-groups are recognized. Cryptodrilus is unique in the Oligochaeta in having multiple excretory bladders in a segment, a condition hitherto unknown. Megascolides diaphanus Spencer, an avesiculate species from Victoria resembling Cryptodrilus in replication of nephridia, is shown to merit recognition of a new monotypic genus, Pseudocryptodrilus. Both Cryptodrilus and Pseudocryptodrilus are shown from the taxonometric study to have perionychine affinities, having close relationships with the vesiculate Plutellus sensu stricto and the avesiculate Simsia respectively. They show only low affinities with dichogastrine genera, represented by the type-species of Notoscolex and Megascolides. Pseudocryptodrilus is assignable to the Perionychini as previously defined but inclusion of Cryptodrilus would require emendment of the definition of the tribe to include total absence of holonephry, a step deferred pending further investigations. The validity of dividing Notoscolex into separate genera on the basis of the excretory system is confirmed. Association (in the Tribe Dichogastrini) of meronephric genera with tubular prostates (formerly placed in the family Octochaetidae) with those with racemose prostates (formerly the family Megascolecidae) receives strong support.     

Phylogenetic studies of Meloidae (Coleoptera), with emphasis on the evolution of phoresy

One of the most recent classifications of Meloidae is based on the assumption that phoretic first‐instar larvae evolved twice in the family, once in Meloinae and again in Nemognathinae. Within Meloinae, this scheme places all presumed phoretic taxa in Meloini regardless of other characteristics. This paper challenges this classification with a cladistic analysis of all meloid genera whose first‐instar larvae were available for study. It concludes that phoresy evolved several times in Meloinae alone and that Meloini, when defined to include all phoretic genera, is polyphyletic. Cladistic support also is presented for four subfamilies of Meloidae and for several of the traditional tribes recognized in recent classifications.