Phylogenetic analysis of the aquatic Oligochaeta under the principle of parsimony

Phylogenetic relationships between five subfamilies of Tubificidae and ten other families of microdrile oligochaetes were estimated by a Wanger parsimony analysis using PAUP (Phylogenetic Analysis Using Parsimony, by D. L. Swofford). As the apomorph character state is ambiguous for some characters, different assumptions of directionality as well as deletions of some characters are tested in a number of analyses. A general pattern is evident from the study; (1) the majority of the aquatic families are members of a large monophyletic group (the order Tubificida in a somewhat restricted sense) defined by the shared possession of atria (generally with well developed external prostate glands), but the family Tubificidae is paraphyletic within this group; (2) the Enchytraeidae appear to form a second group (the Enchytraeida) together with the exclusively marine Capilloventridae and Randiellidae, all three families characterized by the anterior location of the spermathecae; (3) the Haplotaxidae are a plesiomorph family, which stands out as a branch of its own and constitutes the ancestral part of a group comprising also all the megadriles (the Haplotaxida). However, monophyly of the Haplotaxida is likely only if the haplotaxid octogonadial condition is assumed to be derived from the tetragonadial condition characterizing most microdriles, a situation not envisaged by previous authors. The implications of the parsimony method are briefly discussed.     

Incongruence between chloroplast and species phylogenies in Eucalyptus subgenus Monocalyptus (Myrtaceae)

Seventy-eight polymorphic cpDNA (chloroplast DNA) characters were found in 13 closely related taxa from Eucalyptus series Amygdalinae (subgenus Monocalyptus) and seven potential outgroup taxa. The strict consensus of six cladograms generated from cpDNA data confirmed monophyly of Monocalyptus. However, cpDNA phylogeny within Monocalyptus was incongruent with taxonomic classification, being more related to geography, even when accessions were from divergent series. Monocalyptus cpDNA formed two major clades. On the island of Tasmania cpDNA was restricted to a single clade, exhibited very little variation, and was phylogenetically related to cpDNA found in central and western Victoria. In contrast, cpDNA of mainland monocalypt taxa was more variable, even within the Amygdalinae. Four out of six Tasmanian Amygdalinae species were polymorphic. The difference between cpDNA of replicates was often greater than differences between species from different series. The low level of cpDNA variation and extensive morphological intergradation between the Tasmanian endemics suggest recent speciation. However, the transfer of cpDNA through hybridization between lineages is the most likely explanation for the observed sharing of cpDNA across series. This study highlights that the geographical pattern to cpDNA variation in Eucalyptus may be an important source of information on past plant distributions in Australia.     

Chloroplast DNA evidence for reticulate evolution in Eucalyptus (Myrtaceae)

Four highly differentiated chloroplast DNA (cpDNA) lineages were identified in the forest tree species Eucalyptus globulus Labill. (Myrtaceae) in Australia using restriction site polymorphisms from Southern analysis. The cpDNA variation did not conform with ssp. boundaries, yet there was a strong geographical pattern to the distribution of the lineages. One lineage (C) was geographically central and widespread, whereas the other three lineages were found in peripheral populations: Western (W), Northern (N) and Southern (S). Thirteen haplotypes were detected in E. globulus , seven of which belonged to clade C. At least three of the cpDNA lineages (C, N and S) were shared extensively with other species. On the east coast of the island of Tasmania, there was a major north–south difference in cpDNA in the virtually continuous distribution of E. globulus . Northern populations harboured haplotypes from clade C while southeastern populations harboured a single haplotype from clade S. This difference was also reflected in several co-occurring endemic species. It is argued that the extensive cpDNA differentiation within E. globulus is likely to originate from interspecific hybridization and 'chloroplast capture' from different species in different parts of its range. Superficially, this hybridization is not evident in taxonomic traits; however, large-scale common garden experiments have revealed a steep cline in quantitative genetic variation that coincides with the haplotype transition in Tasmania. Our cpDNA results provide the strongest evidence to date that hybridization has had a widespread impact on a eucalypt species and indicate that reticulate evolution may be occurring on an unappreciated scale in Eucalyptus .     

A cladistic analysis ofDipsacaceae (Dipsacales)

A cladistic study ofDipsacaceae (Asteridae, Dipsacales) was undertaken, based mainly on morphological and palynological characters, obtained by investigations of herbarium material and from the literature. Outgroups includedMorinaceae, Triplostegiaceae, and a subset ofValerianaceae. The consensus tree resulting from three equally parsimonious cladograms shows thatDipsacaceae are divided into two major clades, one withDipsacus andCephalaria, the other including the remaining genera. Within the latter clade,Knautia is the sister group of the rest of the taxa. This study is a reappraisal ofDipsacaceae phylogeny, and the results broadly match previous evidence.     

Landscape and taxon age are associated with differing patterns of hybridization in two Eucalyptus (Myrtaceae) subgenera

Background and AimsHybridization is an important evolutionary process that can have a significant impact on natural plant populations. Eucalyptus species are well known for weak reproductive barriers and extensive hybridization within subgenera, but there is little knowledge of whether patterns of hybridization differ among subgenera. Here, we examine eucalypts of Western Australia’s Stirling Range to investigate how patterns of hybridization are associated with landscape and taxon age between the two largest Eucalyptus subgenera: Eucalyptus and Symphyomyrtus. In doing so, we tested a hypothesis of OCBIL (old, climatically buffered, infertile landscape) theory that predicts reduced hybridization on older landscapes.MethodsSingle nucleotide polymorphism markers were applied to confirm the hybrid status, parentage and genetic structure of five suspected hybrid combinations for subg. Eucalyptus and three combinations for subg. Symphyomyrtus.Key ResultsEvidence of hybridization was found in all combinations, and parental taxa were identified for most combinations. The older parental taxa assessed within subg. Eucalyptus, which are widespread on old landscapes, were identified as well-defined genetic entities and all hybrids were exclusively F₁ hybrids. In addition, many combinations showed evidence of clonality, suggesting that the large number of hybrids recorded from some combinations is the result of long-term clonal spread following a few hybridization events rather than frequent hybridization. In contrast, the species in subg. Symphyomyrtus, which typically occur on younger landscapes and are more recently evolved, showed less distinction among parental taxa, and where hybridization was detected, there were high levels of introgression.ConclusionsReduced hybridization in subg. Eucalyptus relative to extensive hybridization in subg. Symphyomyrtus affirmed the hypothesis of reduced hybridization on OCBILs and demonstrate that clade divergence times, landscape age and clonality are important drivers of differing patterns of speciation and hybridization in Eucalyptus.     

Biology of the Eucalyptus leaf beetle Paropsisterna selmani (Chrysomelidae: Paropsini): a new pest of Eucalyptus species (Myrtaceae) in Ireland

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A cladistic analysis of Hydrophis subgenus Chitulia (McDowell, 1972) (Serpentes, Hydrophiidae)

The Hydrophis subgenus Chitulia was analysed using the computer program Hennig86 (version 1.5). The character data set comprises 22 two-state characters, giving a minimum of 22 steps. Four trees with a length of 37 steps and a consistency index of 0.59 were found using the "ie*" option. The results indicate that the subgenus Chitulia is paraphyletic, and that the group has been based solely on plesiomorphic character states. The order of taxa input, the display of the root, and the effect of unknown character states using Hennig86 are commented on.     

A set of chloroplast microsatellite primers for Eucalyptus (Myrtaceae)

We present a set of 35 chloroplast microsatellite primers for Eucalyptus. Ten of the microsatellites displayed intraspecific polymorphism, and identified nine haplotypes among 16 Eucalyptus globulus individuals. Primer conservation was high, with a polymerase chain reaction (PCR) success rate of 98% when tested on four other Eucalyptus species and seven additional myrtaceous genera. Chloroplast microsatellites have applications in phylogeographic studies, fingerprinting and progeny analysis.     

A cladistic analysis of Asarum (Aristolochiaceae) and implications for the evolution of herkogamy

A cladistic analysis of Asarum was conducted to examine relationships among species within the genus and to test the monophyly of several groups of taxa that have often been treated as segregate genera. Thirty-two species were drawn from throughout the range of the genus, representing a broad sample of sections and all segregate genera. The data matrix included 37 characters derived from various aspects of vegetative and floral morphology. A strict consensus of all most parsimonious trees suggests that Asarum s.l. is monophyletic and consists of two main clades: an Asarum clade, which is characterized by connate styles and inferior ovaries, and an Asiasarum-Hexastylis-Heterotropa clade, which is characterized by ridges on the inner perianth surface, dorsal stigmas, and bifid style extensions. The latter is a large and morphologically diverse clade that includes the North American segregate Hexastylis and two Asiatic segregates. Examination of pollination mechanisms in the context of this phylogeny supports the conclusion that herkogamy, and thus obligate insect pollination, is derived from a plesiomorphic condition of autonomous self-pollination. Associated with herkogamy are characters such as glandular trichomes and other ornamentation of the surface of the calyx that probably represent increased specialization to attract insect pollinators. This study also indicates that chromosomal evolution has occurred via aneuploid decrease from an ancestral chromosome number of 2n = 26 to 2n = 24 in Heterotropa. The recognition of two subgenera, subgenus Asarum and subgenus Heterotropa, corresponding to the two clades in the cladistic analysis, is recommended.     

Analysis of hybridization between Eucalyptus pauciflora Sieber ex Spreng. and E. radiata Sieber ex DC. (Myrtaceae)

Eucalyptus is a large genus in which hybridization may produce populations which present taxonomic difficulties. A small group of plants, apparently morphologically intermediate between Eucalyptus pauciflora and E. radiata, was studied in an attempt to determine whether hybridization was occurring, and to investigate the use of different techniques of analysis (morphological, chemical and numerical). Taken as a whole, the data from adult and seedling morphology, flavonoid composition, and volatile oil composition of the plants studied prove that they are indeed hybrids between the two species of Eucalyptus; however, these data also indicate some interesting problems in the application of some of these techniques to this particular situation.     

A cladistic re-assessment of the Polycystis species complex (Polycystididae,Eukalyptorhynchia)

The taxonomy of the genus Polycystis Kölliker, 1845 sensu Schockaert &; Karling, 1975 is discussed. Character analysis reveals the monophyletic state of the three species groups these authors defined within it. As no synapomorphies for the three groups can be found, the genus is split in three genera: Polycystis Kölliker, 1845, Macrorhynchus Graff, 1882 and Paulodora Marcus, 1948. The systematic position of Macrorhynchus yaganus (Marcus, 1954) Evdonin, 1977 is also discussed, resulting in the erection of the new genus Marcusia gen. n.     

Circumscription of Malvaceae (Malvales) as determined by a preliminary cladistic analysis of morphological, anatomical, palynological, and chemical characters

We report a phylogenetic analysis of “core” Malvales (Tiliaceae, Sterculiaceae, Bombacaceae, and Malvaceae) based on morphological, anatomical, palynological, and chemical features. The results of the analyses lead to the conclusion that Tiliaceae, Sterculiaceae, and Bombacaceae, as variously delimited, are paraphyletic; only the Malvaceae are likely monophyletic. The genera of “core” Malvales form a well-defined clade. Genera of “Tiliaceae” constitute the basal complex within “core” Malvales. The “Sterculiaceae” (most genera)+ “Bombacaceae” + Malvaceae form a clade on the basis of a monadelphous androecium; “Bombacaceae”+ Malvaceae also form a clade, which is diagnosable on the basis of monoloculate anthers. It is clear that the traditional classification, with its arbitrarily delimited evolutionary grades, is unsatisfactory, especially if one seeks to reflect phylogeny accurately. Thus, Malvaceae is redefined to refer to the most recent common ancestor of plants previously considered to be “Tiliaceae,” “Sterculiaceae,” “Bombacaceae,” and Malvaceae, and all of the descendants of that ancestor. This broadly circumscribed Malvaceae can be diagnosed by several presumed synapomorphies, but we draw special attention to the unusual floral nectaries that are composed of densely packed, multicellular, glandular hairs on the sepals (or less commonly on the petals or androgynophore).     

Suprageneric relationships of galliform birds (Aves, Galliformes): a cladistic analysis of morphological characters

Of the basal clades of extant birds (Neornithes) the 'landfowl' or galliforms (Aves, Galliformes) are the most speciose. Cladistic analysis of more than 100 morphological characters coded at the generic level for most putative galliform genera confirms that the megapodes ('mound builders'; Megapodiidae) are the most basal clade within the order. They are followed successively by the curassows, guans and chachalacas (Cracidae), which comprise the sister-group to all other extant Galliformes (i.e. Phasianoidea). Within this large 'phasianoid' clade, analyses suggest that the guineafowl (Numididae) are the most basal taxon, although monophyly of this 'family' is not strictly supported on the basis of the morphological characters employed. An additional major clade within the phasianoid Galliformes is recovered by this analysis, comprising the traditional groupings of New World quails (Odontophoridae) and Old World quails ('Perdicini'), yet only monophyly of the former is supported unambiguously by morphological characters. Relationships within the remainder of the phasianoid taxa, including the grouse (Tetraonidae), turkeys (i.e. Meleagris / Agriocharus spp.) as well as other 'pavonine' galliforms (i.e. peafowl; Pavo , Afropavo , Rheinardia , Argusianus and Polyplectron spp.) remain largely unresolved on the basis of morphological characters, yet monophyly of the major subdivisions is supported here. Although there are a number of important differences, especially with regard to relationships within the nonquail phasianoids, the results of this morphological phylogenetic (cladistic) analysis are broadly congruent both with traditional classifications and existing molecular hypotheses of galliform phylogenetic relationships.     

A phylogenetic study of the free-living amoeba

Immunological distances were determined for four strains of the free-living amoeba classified as Amoeba proteus, two strains classified as Polychaos dubia and a single strain classified as Chaos carolinensis. The data show that the _ShP strain does not belong to the proteus group; that A. proteus is more closely related to C. carolinensis and is derived from Chaos as is the _ShP strain; that P. dubia and C. carolinensis are the more distantly related species and appear to be the first of the above to have diverged from a common ancestor; and that the amoebae have a long evolutionary history. The accuracy of the phylogenetic tree and the distance Wagner network was discussed. Since the amoebae may be polyphyletic in origin, the latter was assumed to be a more accurate representation of the immunological distance data.     

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.     

Larvicidal potentials of leaf and bark extracts of Eucalyptus camaldulensis (Schlect) and Eucalyptus citriodora (Hook) on Culex quinquefasciatus (Say) larvae

The larvicidal activity of leaf and bark extracts of Eucalyptus camaldulensis and Eucalyptus citriodora on juveniles of Culex quinquefasciatus was determined. Larvae were successfully bred in the laboratory to obtain a “pure” colony. One hundred and sixty‐two batches of 25 larvae each were treated with 3.90, 15.63, 62.50, 250 and 1000 mg/L of plant extract in three replicates, for 24 h to record mortality and for up to 96 h to record feeding and fecundity. There was an increase in percentage mortality with increase in concentration of the petroleum ether plant extracts. There was a statistically significant difference (P < 0.05) between extracts used in the bioassay and mortality of larvae, but there was no significant difference (P > 0.05) between the two plant species. Statistical analysis of average mortality figures using Probit analysis for the leaf extracts of E. citriodora and E. camaldulensis, respectively, had LC₅₀ of 245.47 (26.89 ± 10.13 [SE]) and 316.23 (22.89 ± 8.95) for the crude extracts; 97.72 (44.95 ± 12.89) and 223.87 (29.17 ± 9.56) for the petroleum ether extracts; and 162.18 (30.7 ± 11.16) and 257.04 (26.47 ± 9.28) for the methanol extracts. The bark extracts of E. citriodora and E. camaldulensis, respectively, had LC₅₀ of 251.19 (27.78 ± 8.39) and 398.11 (20.7 ± 7.17) for methanol; and 302.0 (21.56 ± 8.11) and 630.96 (13.56 ± 6.4) for hexane. Both plant species showed anti‐feeding properties against larvae of Culex quinquefasciatus.     

Phylogeny and evolutionary history of the Aplodontoidea (Mammalia: Rodentia)

Athough over a hundred species of fossil aplodontoids have been described since the extant species, Aplodontia rufa (the mountain beaver), was first described by Rafinesque in 1817, a thorough survey of the relationships among all the species in this clade has not been undertaken since McGrew's study in 1941. Here, a complete phylogenetic analysis of all published species of aplodontoids is used to reconstruct the evolutionary relationships within the clade, and to present an updated classification of the Aplodontoidea. Several of the traditionally recognized subfamilies are found to be paraphyletic, namely the Prosciurinae, the Allomyinae, and the Meniscomyinae. Others, however, including the Aplodontinae and the Mylagaulidae, appear to be monophyletic. These latter two taxa, which include all of the hypsodont members of the aplodontoid clade, seem to be sister taxa. The history of the aplodontoid clade shows several episodes of rapid diversification in the Early Oligocene, the Late Oligocene, and the Early to Middle Miocene. The Ansomyinae and Aplodontinae show comparatively low speciation rates. The patterns of change in morphology and evolutionary rates suggest a need for a more detailed study of the causes of diversification, extinction, and ecological change in this lineage.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 769–838.     

Guticular features of the Central Australian bloodwoods Eucalyptus, section Corymbosae (Myrtaceae)

The morphological features of the cuticles of the leaves, nectaries and stamen filaments of eleven newly published species of Eucalyptus from Central Australia are described and illustrated. The species form four groupS. Within each group, the gross morphology of the species is sufficiently uniform to present difficulty in identification of specimenS. However, the species differ from each other in both qualitative and quantitative cuticular features. The quantitative features include stomatal size and the frequency spectra of subsidiary cells. The abaxial and adaxial frequency spectra of a given species are usually different from each other, but the spectra of different specimens of the same species are closely similar and differ from those of other species of the same group. The general features of the nectary are described. The species differ in nectary pore size, whether the pores are complanate or sunk, etc. Other microscopic features of the nectary cuticles are also species-specific. Different species are shown to have different patterns of the cuticular ornamentation of the stamen filaments.
The importance of cuticular features in identification and definitions of the term 'subsidiary cell' are discussed. Cuticular ornamentation differs in leaves, nectaries and stamen filaments, yet the nature of the cuticle remains the same. Species with relatively large stomata may have small nectary pores, and vice versa. The genetic control of the cuticular features is different in and independent for each tissue, in the leaf even for the abaxial and adaxial surfaces. The adaptive value of cuticular features, especially of the leaves, and the radiative evolution of the Central Australian bloodwoods are discussed.