CLADISTIC ANALYSIS OF PATTERNS OF ENDOTHECIAL THICKENINGS IN THE POALES/RESTIONALES

The three-dimensional structure of the endothecial thickenings in the anthers was investigated in 87 species from 70 genera, chosen to provide representative coverage of the families Cyperaceae, Restionaceae, Anarthriaceae, Ecdeiocoleaceae, Centrolepidaceae, Joinvilleaceae, Flagellariaceae, Poaceae, Xyridaceae, and Eriocaulaceae. There is complex variation in the patterns of endothecial thickening: the Eriocaulaceae, Flagellariaceae, and most Poaceae have thickenings with a complete baseplate; the Cyperaceae and most of the Restionaceae are characterized by helical thickenings; some genera in the Bambusoideae have annular thickenings; and U-shaped thickenings occur in the Xyridaceae and Eriocaulaceae and in some Poaceae and Restionaceae. Joinvillea and Ecdeiocolea have unique thickening types. Endothecial characters were subjected to cladistic analysis. Including endothecial characters in an existing cladogram of the group indicates that there is no single, well-corroborated cladogram available for the Poales/Restionales.     

VESSELS IN JUNCALES: II. CENTROLEPIDACEAE AND RESTIONACEAE

The occurrence and specialization of vessels in 6 genera and 14 species of Centrolepidaceae and 25 and 72, respectively, of Restionaceae are described. Those in Anarthriaceae (1 and 3) and Ecdeiocoleaceae (1 and 1), newly separated from Restionaceae, were also examined. Vessels in Ecdeiocolea are highly specialized; those in Centrolepidaceae are relatively primitive. Vessels in Restionaceae and Anarthriaceae are intermediate. Vessels in Cyperaceae and Gramineae are compared with those in Juncales in a review of vessels in Glumiflorae. Thurniaceae has most primitive vessels on the average, Juncales has more primitive vessels in leaves and inflorescence axes than does Cyperales; the opposite is true for roots, rhizomes and stems. Vessels in Graminales on the average are much more specialized than those in Cyperales and Juncales.     

Gondwanan evolution of the grass alliance of families (Poales)

Phylogenetic interrelationships among all 18 families of Poales were assessed by cladistic analysis of chloroplast DNA rbcL and atpB sequences from 65 species. There are two well-supported main clades; the graminoid clade with Poaceae (grasses), Anarthriaceae, Centrolepidaceae, Ecdeiocoleaceae, Flagellariaceae, Joinvilleaceae, and Restionaceae; and the cyperoid clade with Cyperaceae, Juncaceae, and Thurniaceae. A sister group relationship between Poaceae and Ecdeiocoleaceae is identified with strong support. The sister group of this pair is Joinvilleaceae. These relationships help in elucidating the evolution of grasses and the grass spikelet. Dating of the tree was done by nonparametric rate smoothing of rbcL molecular evolution. Most Poales families date back to the Cretaceous >65 million years ago (mya). Dispersal-vicariance analysis indicates that the Poales originated in South America, the cyperoid clade in West Gondwana (South America or Africa), and the graminoid clade in East Gondwana (Australia). The Trans-Antarctic connection between South America and Australia, and its breakup about 35 mya, probably influenced the evolution of the Poales and the graminoid clade in particular, leading to vicariance between the continents, but the separation of Africa from the other Gondwanan areas, completed about 105 mya, is too old for such a relation.     

Restionaceae seedlings: Morphology,anatomy and systematic implications

Restionaceae differ from most monocot families in having both epigeal and hypogeal germination. The green cotyledons associated with epigeal germination have a central vascular strand as found in most epigeal monocotyledons. In some genera the cotyledon may have a hairpin‐like structure, also described for Anthericaceae. The cotyledon of the hypogeal seedlings is short, without green pigment and largely remains embedded in the seed coat. Hypogeal germination is correlated with large, woody, indehiscent, frequently myrmecochorous nuts, while epigeal germination is found in species with smaller indehiscent nutlets or seeds, dispersed in a variety of ways. The primitive condition is most likely epigeal germination. In hypogeal seedlings of some African and Australian taxa an epicotyledonary rhizome is found between the primary root and the first leaves. Seedlings of African Restionaceae frequently have elongated culm internodes, whereas in the Australian species studied, internodes are very short, resulting in a cluster of seedling leaves. The leaf blades, which in most species are only found on the seedlings, are very simple anatomically. However, they appear to be unifacial, similar to the leaf blades of Anarthria (Anarthriaceae). The anatomical specialisations in the blades mirror those recorded for the culm anatomy. These observations are consistent with the hypothesis that Centrolepidaceae may be neotonous Restionaceae. They also corroborate the morphology of the African Restionaceae, and the presently accepted phylogeny of the African genera of Restionaceae.     

Morphology and development of the gynoecium in Centrolepidaceae: The most remarkable range of variation in Poales

This paper explores the relative impacts of reduction and polymerization on the evolution of reproductive structures in the small but morphologically diverse family Centrolepidaceae. Centrolepidaceae are closely related to Restionaceae and belong to the large order Poales, which also includes the grasses. In the largest genus of Centrolepidaceae, Centrolepis, the reproductive structures are viewed either as highly unusual aggregations of reduced flowers (the pseudanthial interpretation) or as unique flowers evolved through extreme reduction in the androecium, usually accompanied by a drastic increase in carpel number and elaboration of the entire gynoecium. Comparative data are here presented on gynoecia of all three genera of Centrolepidaceae; these data strongly support the latter (euanthial) interpretation. The combined phenomenon of carpel multiplication and decrease in stamen number is unexpected in a predominantly wind-pollinated lineage. Applying a pseudanthial interpretation would create a considerable morphological gap with reproductive structures of other Poales, whereas accepting a euanthial concept allows an almost continuous morphological series with related taxa.     

MEGAGAMETOPHYTE AND NUCELLUS IN RESTIONACEAE AND FLAGELLARIACEAE

Restionaceae are characterized by Polygonum-type development, a well-developed hypostase, the presence of starch grains in the mature female gametophyte, and a thick nucellus that is uniseriate at the micropylar end. Flagellaria indica differs in these characters. In general, proliferation of antipodals occurs in South African taxa of Restionaceae but not in Australasian taxa. The presence of multiplied antipodals is the usual condition in the allied family Poaceae, but the sporadic occurrence of this character in Restionaceae cannot be used to support the view that the two families are sister groups.     

The systematic position of Meruidae (Coleoptera, Adephaga) and the phylogeny of the smaller aquatic adephagan beetle families

A phylogenetic analysis of Adephaga is presented. It is based on 148 morphological characters of adults and larvae and focussed on a placement of the recently described Meruidae, and the genus‐level phylogeny of the smaller aquatic families Gyrinidae, Haliplidae and Noteridae. We found a sister group relationship between Gyrinidae and the remaining adephagan families, as was found in previous studies using morphology. Haliplidae are either the sister group of Dytiscoidea or the sister group of a clade comprising Geadephaga and the dytiscoid families. Trachypachidae was placed as the sister group of the rhysodid‐carabid clade or of Dytiscoidea. The monophyly of Dytiscoidea including Meru is well supported. Autapomorphies are the extensive metathoracic intercoxal septum, the origin of the metafurca from this structure, the loss of Mm. furcacoxalis anterior and posterior, and possibly the presence of an elongated subcubital setal binding patch. Meruidae was placed as sister group of the Noteridae. Synapomorphies are the absence of the transverse ridge of the metaventrite, the fusion of abdominal segments III and IV, the shape of the strongly asymmetric parameres, and the enlargement of antennomeres 5, 7 and 9. The Meru‐noterid clade is the sister group of the remaining Dytiscoidea. The exact position of Aspidytes within this clade remains ambiguous: it is either the sister group of Amphizoidae or the sister group of a clade comprising this family and Hygrobiidae + Dytiscidae. The sister group relationship between Spanglerogyrinae and Gyrininae was strongly supported. The two included genera of Gyrinini form a clade, and Enhydrini are the sister group of a monophylum comprising the remaining Enhydrini and Orectochilini. A branching pattern (Peltodytes + (Brychius + Haliplus)) within Haliplidae was confirmed. Algophilus, Apteraliplus and the Haliplus‐subgenus Liaphlus form a clade. The generic status of the two former taxa is unjustified. The Phreatodytinae are the sister group of Noterinae, and Notomicrus (+ Speonoterus), Hydrocoptus, and Pronoterus branch off successively within this subfamily. The search for the larvae of Meru and a combined analysis of morphological and molecular data should have high priority. © The Willi Hennig Society 2006.     

A revised evolutionary history of Poales: origins and diversification

Poales represents more than one‐third of all monocotyledons (c. 20 000 species in 16 families) and constitutes a microcosm of the angiosperms. The extreme variation in species richness among the families of Poales is still not understood: Poaceae includes ～10 000 species, whereas six families have fewer than ten species. Here, using the largest phylogenetic analysis of Poales to date, molecular dating, ancestral reconstructions and diversification analyses, we develop a macro‐evolutionary and macro‐ecological approach to seek correlates for changing diversification patterns. We show that the poalean families diverged in the Late Cretaceous, a time of high levels of CO₂ and high rainfall. Our habitat reconstructions indicate that Poales inhabited open and dry habitats in this environment. We also demonstrate that lineages with CO₂‐concentrating mechanisms inhabiting dry and open environments exhibited higher diversification rates than C₃, shade and wet lineages. CO₂‐concentrating mechanisms counteract the effects of low atmospheric CO₂ and reduce phototranspiration. It is believed that the parallel evolution of C₄ and CAM (Crassulacean acid metabolism) photosynthesis in Poaceae, Cyperaceae and Bromeliaceae is an adaptation to changes in atmospheric CO₂ concentrations. Combinations of extrinsic and intrinsic factors might have played a role in shifts in diversification rates and may explain the variation in species richness in Poales. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 4–16.     

Ovule,fruit and seed development in Abolboda (Xyridaceae,Poales): implications for taxonomy and phylogeny

Xyridaceae belongs to the xyrid clade of Poales, but the phylogenetic position of the xyrid families is only weakly supported. Xyridaceae is divided into two subfamilies and five genera, the relationships of which remain unclear. The development of the ovule, fruit and seed of Abolboda spp. was studied to identify characteristics of taxonomic and phylogenetic value. All of the studied species share anatropous, tenuinucellate and bitegmic ovules with a micropyle formed by the inner and outer integuments, megagametophyte development of the Polygonum type, seeds with a tanniferous hypostase, a helobial and starchy endosperm and an undifferentiated embryo, seed coat derived from both integuments with a tanniferous tegmen and a micropylar operculum, and fruits with a parenchymatous endocarp and mesocarp and a sclerenchymatous exocarp. Most of the ovule and seed characteristics described for Abolboda are also present in Xyris and may represent a pattern for the family. Abolboda is distinguished by the ovule type, endosperm formation and the number of layers in the seed coat, in agreement with its classification in Abolbodoideae. The following characteristics link Xyridaceae to Eriocaulaceae and Mayacaceae, supporting the xyrid clade: tenuinucellate, bitegmic ovules; seeds with a tanniferous hypostase, a starchy endosperm and an undifferentiated embryo; and a seed coat with a tanniferous tegmen. A micropylar operculum in the seeds of Abolboda is described for the first time here and may represent a synapomorphy for the xyrids. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 144–154.     

Phylogenetic relationships among Poaceae and related families as inferred from morphology, inversions in the plastid genome, and sequence data from the mitochondrial and plastid genomes

A phylogenetic analysis of the Poales was conducted to assess relationships among Poaceae and allied families. The analysis included 40 taxa, representing all families of the Poales as circumscribed by the Angiosperm Phylogeny Group (APG), plus five of the six unplaced Commelinid families in the APG system. The data matrix included 98 informative characters representing variation in morphology and chloroplast genome structure (including three inversions in the chloroplast genome), and 563 informative characters derived from rbcL and atpA nucleotide sequences. Ecdeiocolea has the 6-kilobase (kb) chloroplast genome inversion previously reported in Joinvillea and Poaceae, and like Joinvillea it lacks the trnT inversion that occurs in grasses. Analysis of the morphological data places Poaceae in an unresolved relationship relative to several other taxa, including Joinvillea and Ecdeiocolea, while analysis of the molecular and combined data resolves Ecdeiocolea as sister of Poaceae, with Joinvillea the sister of this group. Although the 6-kb and trnT inversions are non-homoplasious in the phylogenies obtained in this study, the 28-kb inversion is optimized as having originated twice (once in Restionaceae and another time in the most recent common ancestor of Ecdeiocolea, Joinvillea, and the grasses); an alternative interpretation is that it arose once and was later lost in Anarthria. Ecdeiocolea shares with Poaceae the presence of operculate, annulate pollen that lacks scrobiculi, and a dry, indehiscent fruit.     

Illuminating the evolutionary history of liverworts (Marchantiophyta)—towards a natural classification

The phylogenetic relationships of liverworts were reconstructed using the sequence data of four genome regions including rbcL, rps4 and trnL‐F of the chloroplast and 26S large subunit ribosomal rRNA gene of the nucleus, and 90 characters of morphological, ultrastructural and developmental aspects. The taxa sampled consisted of 159 species including 135 liverworts (108 genera, 54 families and 29 suborders), 13 mosses, two hornworts, seven vascular plants and two charophyte algae. Analyses based on maximum parsimony using both direct optimization (POY) and static alignment (NONA), as well as Bayesian inference (MrBayes) were done. All the data sets were analyzed simultaneously. Our study confirms that liverworts compose a monophyletic group which consists of three classes. The class Treubiopsida including both Treubia and Haplomitrium is resolved as the earliest diverging liverwort lineage. Blasia and the complex thalloids are assigned to the Marchantiopsida, under which Blasiidae and Marchantiidae are divided. Marchantiidae include Sphaerocarpales and Marchantiales. The simple thalloid and leafy liverworts form the Jungermanniopsida, which is further divided to subclasses Pelliidae subclassis nov., Metzgeriidae and Jungermanniidae. Metzgeriidae here is defined to include only Metzgeriaceae, Aneuraceae and Vandiemeniaceae, and is the sister group to the leafy liverworts. The leafy liverworts Jungermanniidae include the orders Pleuroziales, Porellales and Jungermanniales. It is assumed that the Porellales and the Jungermanniales have split early, at least in the Jurassic period. In the Porellales, the diversification rate may have remained relatively constant for long periods of time but speeding up only recently within some of the families, associated with an explosive radiation of angiosperms. The Jungermanniales are most probably a recently diversified group which has attained the greatest profusion of structure and the most remarkable diversity of leaf development and protective devices for maturing sporophytes. A detailed classification scheme for liverworts is presented. © The Willi Hennig Society 2006.     

An integrative approach to characterize Malagasy bats of the subfamily Vespertilioninae Gray, 1821, with the description of a new species of Hypsugo

Although important advances have been made in recent years in the taxonomy of different families and subfamilies of Malagasy bats, those belonging to the Vespertilioninae remain partially unresolved. Herein using a mitochondrial marker (cytochrome b) as the point of departure for 76 specimens of Malagasy vespers and appropriate African taxa, we diagnose the six taxa of this subfamily on the island by overlaying different morphological and bioacoustic characters on the clade structure of sequenced animals. The species include: endemic Neoromicia matroka, which is sister to African N. capensis; endemics N. malagasyensis and N. robertsi, which form sister species; a member of the genus Hypsugo, which is sister to African H. anchietae and described herein as new to science; Pipistrellus hesperidus for which Madagascar animals are genetically close but distinct from African populations of the same species; and endemic P. raceyi, which shows segregation of eastern (mesic) and western (dry) populations and its sister species relationships are unresolved. While the external and craniodental measurements, as well as bioacoustic variables, allow only partial differentiation of these six species of Vespertilioninae, molecular characters provide definitive separation of the taxa, as do male bacular morphology. © 2015 The Linnean Society of London     

A COMPARATIVE STUDY OF OVULE AND MEGAGAMETOPHYTE DEVELOPMENT IN FIELD-GROWN AND GREENHOUSE-GROWN PLANTS OF GLYCINE MAX AND PHASEOLUS AUREUS (PAPILIONACEAE)

A study of ovule and megagametophyte development in field- and greenhouse-grown plants of Glycine max (L.) Merrill and Phaseolus aureus Roxb. reveals several consistent features for both species. These features include: a multiple archesporium, enlargement of a primary sporogenous cell directly into a megasporocyte, production of unequal dyad cells, a functional chalazal megaspore, Polygonum-type development, and a hypostase. A filiform apparatus was not observed in either species. Several marked differences in development also occur. Phaseolus usually produces one sporogenous cell per ovule; Glycine produces 2–3 sporogenous cells per ovule. Meiosis II is synchronous in Phaseolus but nonsynchronous in Glycine. Linear tetrads are produced in Phaseolus, whereas linear and T-shaped tetrads are found in Glycine. Starch grains accumulate in the mature megagametophyte of Glycine but are absent at that stage in Phaseolus. The usefulness of the modified clearing fluid, benzyl benzoate-4½, for the study of ovule and megagametophyte development in Glycine max and Phaseolus aureus is here demonstrated. In addition, the study indicates for both species that megagametophyte development in plants grown under field conditions is markedly similar to development in plants grown in the more uniform conditions of the greenhouse. Accordingly, these findings suggest generally that embryological data collected from plants grown under greenhouse conditions will reflect those from plants found in nature.     

In vitro propagation of Western Australian Rushes (Restionaceae and related families) by embryo culture. Part 2. Micropropagation

Micropropagation of 21 species of Restionaceae and the closely relatedmonotypic families Anarthriaceae and Ecdeiocoleaceae is discussed. Multiplication rates ranged from 2-fold to 14-fold each 4–6 week subculture passage, with most species averaging 3–5-fold. The majority of taxa preferred half-strength Murashige and Skoog basal media with 1 M benzyladenine, with certain species requiring other specific treatments (e.g. Woody Plant medium). Approximately half of the species produced roots successfully (i.e. >50%) in vitro on half-strength MS with no growth regulators (or no auxins), or initiated roots after transfer to soil; other species required a longer (6–7 week) incubation on quarter-strength MS medium for initiation to occur. This paper describes the first successful micropropagation of these taxa with application for horticultural development of this important southern-hemisphere family.Abbreviations MS Murashige & Skoog (1962) basal medium - 1/4 MS quarter-strength MS (minerals only) - 1/2 MS half-strength MS (minerals only) - BA benzyladenine - WPM woody plant medium (Lloyd and McCown 1980) - GA₃ gibberellic acid - TDZ thidiazuron - IBA indolebutyric acid     

Phylogeny of scolopendromorph centipedes (Chilopoda): morphological analysis featuring characters from the peristomatic area

Alternative schemes for the higher‐level systematics of the centipede order Scolopendromorpha have been established from characters of trunk segmentation, including the segmental position of spiracles, and the presence or absence of eyes. A comparative survey of the preoral chamber by light and scanning electron microscopy contributes 16 new characters of the epipharynx and hypopharynx, sampled from 26 species that represent 20 genera. These new data together with 49 additional morphological characters permit cladistic analysis of the major scolopendromorph groups. The shortest cladograms resolve blind Scolopendromorpha as a basal grade within which a clade now classified as Plutoniuminae + Scolopocryptopidae (supported by unreversed characters from the preoral chamber) is sister to the remaining scolopendromorphs. A unique row of bullet‐shaped sensilla between the labral and clypeal parts of the epipharynx provides a new autapomorphy of the Scolopendromorpha. Either 21 or 23 trunk segments optimize at the base of the Scolopendromorpha but in either case homoplasy is forced on the cladogram. New characters from the epipharynx give additional support for the monophyly of several traditional groupings, including Cryptopinae, Scolopendridae, Otostigmini, and Scolopendrini, and a basal resolution of Asanadini within Scolopendridae. Of the two competing hypotheses for the position of the enigmatic Mediterranean Plutonium zwierleini—being either sister to the cryptopid Theatops or sister to all other Scolopendromorpha—the former hypothesis is strongly supported; spiracles on all trunk segments in Plutonium are homoplastic with the state in Geophilomorpha. Observations on feeding behaviour are needed to illuminate convergence in characters of the epipharynx and mandible in Edentistoma (Otostigminae) and Campylostigmus (Scolopendrini). © The Willi Hennig Society 2008.     

Limb myology and phylogenetic relationships in the superfamily Dipodoidea (birch mice, jumping mice, and jerboas)

Limb muscles were dissected in seven genera, representing all six superfamilies, of dipodoid rodents and myoloic characters were used to construct a phylogenetic hpothesis of relationships within this cfade. Mologic differences among genera suported tie monophyly of the superfamily Dipodoidea reLtive to the outrou taxon and reveaEd thatSicista is the sister group to all other zapodid and dipodid enera. Tkis picement of Sicista differs markedly from its position in previous classifications where it has been regarded merely as a primitive zapodid genus. The phlograrn based on rnyologic characters also indicated that Cardiocranius is not a rimitive dipodid genus; it is the sister group to the subfamily Dipodinae. Although myologic differences among taxa were not sufficient to warrant the continued separation of zaodids and dipodids into two families, a new classification that places Sicista in its own family, ficistidae, and places the remaining zaodids and dipodids in the family Dipodidae, is proposed. Differences in karyology, genitaP morholoy, and postcranial osteological characters among dipodoid rodents are discussed in light or this pjylogeny.     

Identifying the sister group to the bees: a molecular phylogeny of Aculeata with an emphasis on the superfamily Apoidea

Debevec, AH., Cardinal, S & Danforth, BN. Identifying the sister group to the bees: a molecular phylogeny of Aculeata with an emphasis on the superfamily Apoidea. —Zoologica Scripta, 41, 527–535. The hymenopteran superfamily Apoidea includes the bees (Anthophila) as well as four predatory wasp families (Heterogynaidae, Ampulicidae, Sphecidae and Crabronidae) collectively referred to as the “sphecoid” or “apoid” wasps. The most widely cited studies suggest that bees are sister to the wasp family Crabronidae, but alternative hypotheses have been proposed based on both morphological and molecular data. We combined DNA sequence data from previously published studies and newly generated data for four nuclear genes (28S, long‐wavelength rhodopsin, elongation factor‐1α and wingless) to identify the likely sister group to the bees. Analysis of our four‐gene data set by maximum likelihood and Bayesian methods indicates that bees most likely arise from within a paraphyletic Crabronidae. Possible sister groups to the bees include Philanthinae, Pemphredoninae or Philanthinae + Pemphredoninae. We used Bayesian methods to explore the robustness of our results. Bayes Factor tests strongly rejected the hypotheses of crabronid monophyly as well as placement of Heterogynaidae within Crabronidae. Our results were also stable to alternative rootings of the bees. These findings provide additional support for the hypothesis that bees arise from within Crabronidae, rather than being sister to Crabronidae, thus altering our understanding of bee ancestry and evolutionary history.     

Toward a Better Understanding of the Phylogeny of the Asplanchnidae (Rotifera)

We investigated the phylogenetic relationships of Family Asplanchnidae using both morphological and molecular data. The morphological database, comprising 23 characters from 19 taxa (15 Asplanchnidae and 4 outgroups), was compiled from a survey of the literature and our own observations; the molecular data (ITS and V4 region nuclear regions and mitochondrial cox1) was sequenced from specimens that we collected. Our analysis of the morphological data set (maximum parsimony) yielded 12 most-parsimonious trees with a tree length of 27 steps. From this analysis we conclude (1) Asplanchnidae is a monophyletic group as are the three genera comprising it, (2) there is no compelling support for the argument that Asplanchna should be separated into two discrete genera, and (3) there is some support for the proposal that Asplanchnidae and Synchaetidae are sister groups. Our analysis of the molecular data set supports the first two of these conclusions while the sister group of the family varied depending on the gene region analyzed and families and genera included. Current understanding of the phylogeny of Asplanchnidae is hampered by the need for additional informative morphological characters and a lack of molecular data for the genus Harringia and several other members of the Asplanchnidae.