Phylogeny of <Emphasis Type="Italic">Gunnera</Emphasis>

 The phylogeny of the genus Gunnera is investigated for the first time. Twelve species representing the six currently recognised subgenera are analysed. Two chloroplast DNA regions, the rbcL gene and the rps16 intron, together provide 46 informative characters out of 2335. A combined analysis of both genes gives four most parsimonious trees, firmly establishing the east South American G. herteri as sister group to the rest of the genus. The African G. perpensa is sister group to two well-supported clades, one including the South American subgenera Misandra and Panke, the other the Australian/New Zealand/Malayan species of subgenera Milligania and Pseudogunnera. Thus, South America is a composite area for Gunnera, showing up at two different levels in the cladogram. Our analysis supports a close biogeographic relationship between Australia and New Zealand. The evolution of some morphological characters is discussed. Lastly, the unusual structure of some of the rbcL sequences is reported. Received July 6, 2000 Accepted October 24, 2000     

The biogeography of Gunnera L.: vicariance and dispersal

Aim The genus Gunnera is distributed in South America, Africa and the Australasian region, a few species reaching Hawaii and southern Mexico in the North. A cladogram was used to (1) discuss the biogeography of Gunnera and (2) subsequently compare this biogeographical pattern with the geological history of continents and the patterns reported for other Southern Hemisphere organisms. Location Africa, northern South America, southern South America, Tasmania, New Zealand, New Guinea/Malaya, Hawaii, North America, Antarctica. Methods A phylogenetic analysis of twenty‐six species of Gunnera combining morphological characters and new as well as published sequences of the ITS region, rbcL and the rps16 intron, was used to interpret the biogeographical patterns in Gunnera. Vicariance was applied in the first place and dispersal was only assumed as a second best explanation. Results The Uruguayan/Brazilian Gunnera herteri Osten (subgenus Ostenigunnera Mattfeld) is sister to the rest of the genus, followed sequentially upwards by the African G. perpensa L. (subgenus Gunnera), in turn sister to all other, American and Australasian, species. These are divided into two clades, one containing American/Hawaiian species, the other containing all Australasian species. Within the Australasian clade, G. macrophylla Blume (subgenus Pseudogunnera Schindler), occurring in New Guinea and Malaya, is sister to a clade including the species from New Zealand and Tasmania (subgenus Milligania Schindler). The southern South American subgenus Misandra Schindler is sister to a clade containing the remaining American, as well as the Hawaiian species (subgenus Panke Schindler). Within subgenus Panke, G. mexicana Brandegee, the only North American species in the genus, is sister to a clade wherein the Hawaiian species are basal to all south and central American taxa. Main conclusions According to the cladogram, South America appears in two places, suggesting an historical explanation for northern South America to be separate from southern South America. Following a well‐known biogeographical pattern of vicariance, Africa is the sister area to the combined southern South America/Australasian clade. Within the Australasian clade, New Zealand is more closely related to New Guinea/Malaya than to southern South America, a pattern found in other plant cladograms, contradictory to some of the patterns supported by animal clades and by the geological hypothesis, respectively. The position of the Tasmanian G. cordifolia, nested within the New Zealand clade indicates dispersal of this species to Tasmania. The position of G. mexicana, the only North American species, as sister to the remaining species of subgenus Panke together with the subsequent sister relation between Hawaii and southern South America, may reflect a North American origin of Panke and a recolonization of South America from the north. This is in agreement with the early North American fossil record of Gunnera and the apparent young age of the South American clade.     

Taxonomic implications of pollen exine morphology in infrageneric classification of Scutellaria (Lamiaceae)

Scutellaria L. in Lamiaceae subfamily Scutellarioideae is a subcosmopolitan genus with about 360 species. According to the latest subgeneric classification, two subgenera are recognized: S. subgen. Scutellaria and S. subgen. Apeltanthus. We studied pollen grains from 29 species of Scutellaria belonging to both subgenera and from both the Old and the New World to evaluate their taxonomic importance for the infrageneric classification of the genus. Pollen grains were acetolysed and studied by scanning electron microscope (SEM) and light microscope (LM). All examined pollen grains are isopolar, tricolpate, suboblate, oblate spheroidal, prolate spheroidal to subprolate. The exine is mainly bireticulate perforate, but with variations in lumina shape and size as well as perforation size and uniformity. Based on the exine sculpturing, two subtypes are recognized within the bireticulate pollen type. In subtype I the primary reticulum of the exine consists of angular muri; the lumen has an irregular polygonal shape and the perforations of the secondary reticulum are more or less uniform and of the same size. This pollen subtype was observed in species of subgenus Scutellaria, sectt. Scutellaria p.p., Anaspis, Perilomia and Salazaria. In subtype II the primary reticulum of the exine consists of a more or less curved murus; the lumen is rounded or obtusely angular and the secondary reticulum consists of one or more large central perforations surrounded by some smaller ones. This pollen subtype was observed in species of subgenus Scutellaria sectt. Scutellaria p.p., Salviifoliae and in all members of subgenus Apeltanthus. The results of this study reveal that the exine ornamentation is a diagnostic character useful for the infrageneric classification of Scutellaria.     

Systematics and Leaf Architecture of the Gunneraceae

Cladistic and phenetic analyses of leaf and other morphological characters ofGunnera strongly support monophyly of the genus, with the Saxifragaceae s.str. as the closest sister group. This morphologically based phylogeny provides a more coherent understanding of the evolutionary history ofGunnera than do recent phylogenetic hypotheses based on genetic data sets with Myrothamnaceae as the sister group. Simple, crenate, palinactinodromously veined leaves lacking freely ending veinlets and tricolpate, tectate-perforate pollen with a reticulate exine indicate a shared ancestry. Within the genusGunnera all six traditionally recognized subgenera are monophyletic, as supported by leaf architectural apomorphies. The monotypic subgenusOstenigunnera is the sister group to the other five subgenera, which can be divided into two principal lineages. One lineage includes the subgeneraMilligania andMisandra, characterized by a prostate stoloniferous habit with small, low-rank leaves and exclusively unisexual flowers, whereas the other lineage includes the subgeneraPerpensum, Pseudo Gunnera, andPanke, all of which possess at least some hermaphroditic flowers and larger, high-rank leaves. When the phylogeny of the subgenera is considered in light of biogeography and the fossil record, a number of cladogenetic events can be explained by continental vicariance in the Late Cretaceous. The AfricanPerpensum became distinct from the other large-leafed lineage with the separation of the African continent ca. 90 Ma. The two small-leafed lineages, the subgeneraMilligania andMisandra, split with the separation of New Zealand from Western Gondwana, about 80 Ma.Pseudo-Gunnera became isolated fromPanke prior to this time, whenPanke fossils occur in North America.Gunnera probably arose out of an early herbaceous radiation of tricolpate eudicots having close affinity to the basal Saxifragaceae, espethe genusChrysosplenium.     

Molecular phylogeny of the subgenus Holothuria (Selenkothuria) Deichmann, 1958 (Holothuroidea: Aspidochirotida)

The subgenus Selenkothuria comprises 12 species of tropical shallow water sea cucumbers that share morphological features, such as rods in the body wall and tube feet, modified tentacles for suspension feeding, and cryptic colours. The taxonomic status of this taxon has been controversial, but currently it is accepted as a subgenus of the genus Holothuria. Phylogenetic analyses of mitochondrial genes [cytochrome c oxidase subunit 1 (COI), 16S RNA] of ten species of Selenkothuria and related subgenera showed the polyphyly of this subgenus; monophyly was rejected by a likelihood ratio test. A geographical split divides the species of this subgenus into three different groups: one Indo‐West‐Pacific (IWP) group and two American groups. The IWP group is more closely related to Holothuria (Semperothuria) cinerascens and to other subgenera such as Roweothuria, Holothuria, and Vaneyothuria, whereas the two American groups are more closely related to each other and to some species of the subgenus Halodeima. These results suggest multiple parallel originations and diversification of ossicle morphology within the subgenus Selenkothuria. The current scheme of subgenera for the genus Holothuria is not supported, suggesting the need for a new classification. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 109–120.     

Microsporogenesis, microgametogenesis and pollen morphology of Passiflora spp. (Passifloraceae)

Microsporogenesis, microgametogenesis and pollen morphology of six species of genus Passiflora L. belonging to three subgenera ( Passiflora , Dysosmia , Decaloba ) were studied with light and scanning microscopy; P. caerulea was also examined with transmission microscopy. The tapetum is secretory, microspore tetrads are tetrahedral and pollen grains are two-celled when shed. Small Ubisch bodies are attached to a peritapetal membrane; they are a product of tapetal activity and the rough endoplasmic reticulum (ERr) appears to be involved in their origin. The pollen grains of all the species are subspheroidal, zonocolpate, geminicolpate. Each pair of colpi anastomoses at the poles. The exine is semitectate, reticulate, heterobrochate. The muri are simplibaculate, wavy. The lumina have clavate bacula of varying height. The colpus structure is similar to that of the lumina but generally with fewer and smaller bacula. Lumina size and amount of bacula inside the lumina vary between subgenera. The grains from subgenera Passiflora and Dysosmia differ from those of Decaloba in their size and number of colpi. The pollen and microsporangium morphology of the species of subgenera Passiflora and Dysosmia are more similar than those of subgenus Decaloba . The results are discussed in relation to the current taxonomic classification.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society of London , 2002, 139 , 383–394.     

Pollen morphology of Chamaebuxus (DC.) Schb., Chodatia Paiva and Rhinotropis (Blake) Paiva (Polygala L., Polygalaceae)

Polygala L. is a large and highly diverse genus with complex taxonomy, but pollen morphological information for this taxon is scarce. In the present study, pollen characters have been used to assess the taxonomic delimitation and phylogenetic relationships of three newly established subgenera of Polygala: Chamaebuxus, Chodatia and Rhinotropis (sensu Paiva). The pollen morphology of 22 species has been examined using light microscopy and scanning electron microscopy of acetolysed material. The pollen of 15 of the species is examined for the first time. The pollen grains are isopolar, radially symmetrical, tectate and, typically, polyzonocolporate with numerous colpi running parallel to the polar axis, and an endocingulum around the equator. Two pollen types can be distinguished: Type I, which includes species belonging to Rhinotropis, and Type II, which includes species from Chamaebuxus and Chodatia. The two pollen types are described and the pollen of the three studied subgenera is illustrated. Despite the low infrageneric morphological diversity observed within the genus Polygala, quantitative characters of pollen grains support the current classification of the subgenera Chamaebuxus, Chodatia and Rhinotropis, and reveal a closer relationship between the first two taxa. Pollen characters are shown to be a useful and informative tool for assessing taxonomic position and phylogenetic relationships within Polygalaceae, especially at higher taxonomic levels.     

Molecular phylogenetics of the burrowing crayfish genus Fallicambarus (Decapoda: Cambaridae)

Ainscough, B.J., Breinholt, J.W., Robison, H.W. & Crandall, K.A. (2013). Molecular phylogenetics of the burrowing crayfish genus Fallicambarus (Decapoda: Cambaridae). —Zoologica Scripta, 42, 306–316. The crayfish genus Fallicambarus contains 19 species of primary burrowing freshwater crayfish divided into two distinct subgenera. We test current hypotheses of the phylogenetic relationships among species within the genus as well as the monophyly of the genus. Our study samples all 19 species for five gene regions (both nuclear and mitochondrial) to estimate a robust phylogenetic hypothesis for the genus. We show that the genus is not a monophyletic group. The subgenus Creaserinus does fall out as a monophyletic group, but distinct from the subgenus Fallicambarus. The subgenus Fallicambarus appears to be monophyletic with the exception of the species Procambarus (Tenuicambarus) tenuis, which falls in the midst of this subgenus suggesting that it might be better classified as a Fallicambarus species. We also show that the species Fallicambarus fodiens is a species complex with distinct evolutionary lineages that are regionalized to different geographic areas.     

Ornamentation of achene silica walls and its contribution to the systematics of <Emphasis Type="Italic">Eleocharis</Emphasis> (Cyperaceae)

Micromorphology of the achene surface of 26 Brazilian species of Eleocharis was studied by scanning electron microscopy in order to evaluate its usefulness in the taxonomy of the genus. The results point out two patterns of cell organization according to silica structures of achenes. The first corresponds to a group of species (group A) that have small to medium cells arranged vertically. The second is found in those species (group B) with medium to large cells arranged horizontally. These data were useful in separating species of Eleocharis subgenus Scirpidium and E. subgenus Limnochloa (group B) from E. subgenus Eleocharis (group A). However, group A shows considerable variation in silica wall arrangement. Eleocharis squamigera, previously considered as part of E. subgenus Eleocharis, shows features rather similar to those of Scirpidium, confirming recent phylogenies. The subgenus Limnochloa was clearly distinguished from others by achenes with large cells (over 55 μm width), presence of crenate or repand anticlinal walls, and some orifices near the wall in some species. The silica wall ornamentation seems to be a useful morphological tool for studying relationships between subgenera and distinguishes Limnochloa from the other subgenera.     

Physiological,Biochemical, and Molecular Characters for the Taxonomy of the Subgenera of Scenedesmus (Chlorococcales,Chlorophyta)

Biochemical and physiological properties of 16 Scenedesmus species representing the three subgenera Scenedesmus, Acutodesmus, and Desmodesmus are not suitable for species differentiation. All Scenedesmus species studied thus far produce secondary carotenoids, e.g. astaxanthin and canthaxanthin, under nitrogen-deficient conditions. In addition, with the exception of one strain, hydrogenase activity under anaerobic conditions is generally present. Sequence analyses of ribosomal 18S RNAs indicate that the subgenus Desmodesmus is phylogenetically well separated from the other subgenera, whereas the separation of Scenedesmus and Acutodesmus appears less convincing and is dismissed in favour of a single subgenus, Scenedesmus. Three taxa formerly assigned to the genus Chlorella are shown to be unicellular species of the genus Scenedesmus. “Chlorella” fusca var. vacuolata and “C.” fusca var. rubescens, which is closely related to S. obliquus, belong to the Scenedesmus/‘Acutodesmus group. “C.” fusca var. fusca is closely related to S. communis and thus belongs to the subgenus Desmodesmus. Inclusion of Kermatia pupukensis into the genus Scenedesmus is also strongly supported by the RNA data which furthermore indicate a relationship with the subgenus Desmodesmus.     

The palynology of the genus Rondeletia L. (Rubiaceae-Cinchonoideae-Rondeletieae)

Pollen of 20 species of Rondeletia, covering three subgenera and seven sections or “species groups” of the genus, was studied using light and scanning microscopy. Rondeletia, characterized by binucleate mature pollen grains like numerous other Rubiaceae, was found to be eurypalynous. The pollen is isopolar and small-sized (average equatorial diameters 11.4–22.8 μm) and 3-colporate (rarely 4-colporate). In equatorial view, the pollen grains are mostly ± spheroidal (rarely suboblate); in polar view (ambitus), they are mainly circular or semiangular (intersemiangular in one species only). Three different exine types are distinguishable: exine with 1) tectum minutely perforate, perforations usually isodiametric, 2) tectum with comparatively large, not isodiametric perforations or 3) coarsely reticulate; supratectal sculpturing elements are lacking; the width of the perforations and lumina is either equal or different at the apo- and mesocolpia; in a few species rod-like luminal processes are present. Palynological features of most of the investigated taxa tend to overlap so that palynology is not very helpful in supporting the subgeneric division of Rondeletia. It was, however, noted that pollen morphology, to some extent, supports the sections of subgen. Arachnothrix. Palynologically, the subgen. Rogiera was found to be heterogeneous in that each of the three investigated species is characterized by a different exine type; one of these, R. strigosa, differs from all other Rondeletia species investigated in having a coarsely reticulate exine. R. odorata (subgen. Rondeletia) differs from all other investigated species in its intersemiangular pollen shape.     

Marker chromosomes commonly observed in the genus Glycine

Summary Soybean [Glycine max (L.) Merr.] chromosomes were analyzed using the chromosome image analyzing system, CHIAS, and seven groups, including subgroups, were identified based on morphological characteristics. Two pairs of chromosomes were conspicuous in their morphological traits. One pair of chromosomes, which had the largest arm ratio among all the chromosomes, was commonly observed in the species in all three subgenera of the genus Glycine. These chromosomes also displayed a unique pattern after N-banding and were detected as marker chromosomes. G. soja, which is considered to be the ancestor of G. max, has two types of marker chromosomes. The lines that carry the same type as G. max may be the ancestors of G. max among the lines of G. soja. The morphological differences of the marker chromosomes within the species in the subgenus Soja are discussed in relation to the domestication process of soybean.     

REVISION OF THE GENUS OPACOPTERA GOZMANY* (LEPIDOPTERA: LECITHOCERIDAE)

Abstract The characters of the genus Opacoptera Gozmány are revised and 3 species belonging to 2 subgenera are treated. A new subgenus, Fulvitalia, and a new species, O. (O.) ecbasta, are described and figured. Key to the species is provided.     

Mise à jour taxonomique et répartition des puces du genre Ctenophthalmus Kolenati 1856 en région paléarctique occidentale (Insecta : Siphonaptera : Ctenophthalmidae)

Summary. Taxonomic update and geographic distribution of fleas of the genus Ctenophthalmus Kolenati 1856 in the Western Palearctic Region (Insecta: Siphonaptera: Ctenophthalmidae). Among fleas (Siphonaptera), the genus Ctenophthalmus is the one that comprises the largest number of taxa and is also characterized by a large geographical range. Here, we present a taxonomic revision of the Western Paleartic subgenera, groups, species and subspecies. We recognized a total of 143 taxa (57 species and 86 subspecies). These taxa are clustered into 23 groups of species, which fall into seven of the 16 subgenera of the genus Ctenophthalmus. According to Hopkins & Rothschild (1966), the subgenus Ctenophthalmus would only include the agyrtes group, itself divided into subgroups. We decided to raise these subgroups to group status to clarify taxonomic relationships within the subgenus Ctenophthalmus. Within this subgenus, the arvernus group is renamed baeticus, the fransmiti group is confirmed, and the egregius group is created. For each taxon, we provided information on geographical distribution, mammalian hosts, and host specificity.     

A review of the tribe Auletini (Coleoptera,Rhynchitidae) from the Russian fauna: 1. Subtribe Auletobiina

The genus Auletobius in the Russian fauna is revised. Five species (A. egorovi, A. irkutensis, A. puberulus, A. sanguisorbae, and A. submaculatus) belonging to two subgenera are revealed. The distribution of these species in Russia is given. The data on the trophic associations of the species are summarized. Keys to the subtribes of the tribe Auletini, subgenera of the genus Auletobius, and species of the subgenus Auletobius s. str. are given. All the taxa are redescribed.     

Molecular phylogenetics and biogeography provide insights into the subgeneric classification of Wiedemannia Zetterstedt (Diptera: Empididae: Clinocerinae)

The subgenera of Wiedemannia are poorly defined and, as such, most recently described species are not assigned to a subgenus or have been assigned to a subgenus without explanation. In this study we perform a molecular phylogenetic analysis to elucidate relationships within the genus Wiedemannia. We sequenced two mitochondrial (cytochrome oxidase c subunit I and cytochrome β) and two nuclear (carbomoylphosphate synthase domain of rudimentary and elongation factor‐1α) gene fragments to reconstruct phylogenetic relationships among the subgenera Chamaedipsia, Eucelidia, Philolutra, Pseudowiedemannia, Roederella and Wiedemannia (s.s.) using both Bayesian inference and maximum likelihood approaches. The genus was found to be monophyletic, but most of the subgenera were not. We propose eliminating the present subgeneric division altogether. Molecular dating using a log‐normal clock model and calibration with fossil species indicated that Wiedemannia diversified about 48 Ma, while there was still land connectivity between Europe and Asia with North America. Wiedemannia has a near‐worldwide distribution apart from the Australasian and Neotropical regions and Antarctica, with greatest species richness in the western Palaearctic, especially the Mediterranean region. Molecular phylogenetics support more recent morphological studies. The subgenera of Wiedemannia are invalid and rejected. Biogeographical data suggest potential hotspots, and the current distribution is discussed.     

Phylogenetic relationships of the chloroplast genomes in the genus Glycine inferred from four intergenic spacer sequences

The nucleotide sequences of four intergenic spacer regions of chloroplast DNA, atpB-rbcL, trnS-trnG, rps11-rpl36, and rps3-rpl16, were analyzed in the genus Glycine. Phylogenetic analysis based on the sequence data using Neonotonia wightii as the outgroup generated trees supporting the classification of two subgenera, Soja and Glycine, and three plastome groups in the subgenus Glycine. The results were consistent with the presence of diversified chloroplast genomes within tetraploid plants of G. tabacina and G. tomentella, as well as with a close relationship between G. tomentella and G. dolichocarpa that had been suggested based on morphological analyses. Little sequence variation was found in the subgenus Soja, suggesting that G. soja rapidly expanded its distribution in East Asia. The analysis also showed that the differentiation into three plastome groups in the subgenus Glycine occurred in the early stages of its evolution, after the two subgenera diverged.     

Pollen morphology of selected Central European species from subgenera Vignea and Carex (Carex,Cyperaceae) and its relation to taxonomy

This study analysed the taxonomic utility of selected features of pollen grains of 45 Central European species of subgenera Vignea and Carex of the genus Carex. Seven quantitative features were analysed: [length of polar axis (P), length of equatorial axis (E), exine thickness on the pole (Exp), exine thickness on the equator plane (Exe), P/E ratio and the relative thickness of exine (Exp/P and Exe/E ratio)] and outline, shape and exine sculpture. Among the biometrical features, the most important for Carex taxonomy (particularly at the level of subgenus) is the thickness of exine (Exp). Marked differences in the morphological structure of inflorescences at the subgenus level are reflected in the differences of pollen features. Results of our studies clearly show an explicit (and not previously described in the palynological literature) taxonomic division of the genus Carex on the basis of pollen biometrical features into two units corresponding exactly to the subgenera (Vignea and Carex). However, the features of pollen grains analysed did not contribute any essential taxonomic criteria at levels lower than subgenus (e.g. sections). An exception was for C. disticha, where it was observed that the poroid areas occurred in pairs, a new feature for sedges. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 422–439.     

Pollen morphology and systematics of palaeotropical Phyllanthus and related genera of subtribe Phyllanthinae (Euphorbiaceae)

The observations of pollen from 27 species of subtribe Phyllanthinae using scanning electron microscopy reveal considerable morphological diversity in palaeotropical Phyllanthus and the related palaeotropical genera Breynia, Glochidion, and Sauropus. The tribe appears to be monophyletic, but the pollen morphology does not support the monophyly of Phyllanthus or Sauropus. Within Phyllanthus, the pollen characters suggest a close relationship between the subgenera Emblica and Phyllanthodendron. They also reveal a surprising morphological congruence between the pollen grains of section Ceramanthus (subgenus Isocladus) and those of subgenus Eriococcus, although it is not clear whether this similarity is homoplastic. The presence of diploporate colpi is a synapomorphy uniting Breynia and Sauropus, but may be homoplastic in Phyllanthus. The affinities suggested by the morphological features of the pollen in the Phyllanthinae are concordant with recent molecular phylogenies. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 591–608.