Relationships among the major lineages of Dictyoptera: the effect of outgroup selection on dictyopteran tree topology

Abstract Dictyoptera, comprising Blattaria, Isoptera, and Mantodea, are diverse in appearance and life history, and are strongly supported as monophyletic. We downloaded COII, 16S, 18S, and 28S sequences of 39 dictyopteran species from GenBank. Ribosomal RNA sequences were aligned manually with reference to secondary structure. We included morphological data (maximum of 175 characters) for 12 of these taxa and for an additional 15 dictyopteran taxa (for which we had only morphological data). We had two datasets, a 59‐taxon dataset with five outgroup taxa, from Phasmatodea (2 taxa), Mantophasmatodea (1 taxon), Embioptera (1 taxon), and Grylloblattodea (1 taxon), and a 62‐taxon dataset with three additional outgroup taxa from Plecoptera (1 taxon), Dermaptera (1 taxon) and Orthoptera (1 taxon). We analysed the combined molecular?morphological dataset using the doublet and MK models in Mr Bayes , and using a parsimony heuristic search in paup . Within the monophyletic Mantodea, Mantoida is recovered as sister to the rest of Mantodea, followed by Chaeteessa; the monophyly of most of the more derived families as defined currently is not supported. We recovered novel phylogenetic hypotheses about the taxa within Blattodea (following Hennig, containing Isoptera). Unique to our study, one Bayesian analysis places Polyphagoidea as sister to all other Dictyoptera; other analyses and/or the addition of certain orthopteran sequences, however, place Polyphagoidea more deeply within Dictyoptera. Isoptera falls within the cockroaches, sister to the genus Cryptocercus. Separate parsimony analyses of independent gene fragments suggest that gene selection is an important factor in tree reconstruction. When we varied the ingroup taxa and/or outgroup taxa, the internal dictyopteran relationships differed in the position of several taxa of interest, including Cryptocercus, Polyphaga, Periplaneta and Supella. This provides further evidence that the choice of both outgroup and ingroup taxa greatly affects tree topology.     

The evolution of Synchaetidae (Rotifera: Monogononta) with a focus on Synchaeta: An integrative approach combining molecular and morphological data

This study represents the first phylogenetic reconstruction of Synchaetidae (Rotifera, Monogononta) and combines one morphological and two molecular data sets to derive the most comprehensive database for this taxon to date. Overall, 19 species were examined both morphologically (light- and scanning electron microscopy of habitus and trophi) and genetically (COI and 18S rRNA genes), thereby delivering new morphological information as well as the first molecular data for several species. Our results strongly support the monophyly of each of the three genera within Synchaetidae, with Synchaeta being the sister taxon of Polyarthra and Ploesoma. Resolution within each genus, however, was more poorly supported, possibly because of high levels of missing data for the more poorly known species that were also not re-discovered and revised in the present study. Nevertheless, an evolutionary reconstruction based on the total evidence topology indicates that the common ancestor of Synchaetidae was a rotatorivorous and pelagic freshwater rotifer exhibiting slightly enlarged lamellar trophi that were adapted to an enhanced pumping function. These features together with several, additional key character transformations (e.g., hatching independently from benthic or periphytic habitats) probably account for the success of Synchaetidae in pelagic environments and their being one of the most widely distributed and abundant rotifer taxa.     

Genetic diversity and photobiont associations in selected taxa of the Tephromela atra group (Lecanorales, lichenised Ascomycota)

Mycobiont and photobiont genetic diversity was investigated in four taxa of the Tephromela atra complex, which differ in ecology and substratum preference (from siliceous rocks, limestone to bark), and are differently interpreted by taxonomists. Phylogenetic analyses were performed using mycobiont nuclear ITS, beta tubulin and homologous polyketide synthase gene (PKS) sequences obtained from freshly collected material sampled from the Mediterranean region to the Southern Alps. The silicicolous samples from the Alps form a basal lineage of the entire complex, and despite the morphological similarity, they do not form a monophyletic group with the Mediterranean samples. No resolution was found among the calcicolous and the silicicolous taxa from Mediterranean habitats, which are traditionally segregated at variety or species level. The epiphytic taxon, although nested with the other ecotypes, splits in two well-supported lineages. Among the four taxa, Tephromela grumosa is the only morphologically, chemically and genetically distinct taxon. However, it is also nested in the large T. atra complex. Phylogenetic analysis of photobionts ITS sequences revealed that thalli from the Mediterranean region are associated with two distinct lineages of Trebouxia, but the lineages are not correlated with substrate or mycobiont phenotype. The thalli from the Alps are exclusively associated with T. simplex, suggesting a protracted isolation from the other lineages.     

Tumbleweed (Salsola, section Kali) species and speciation in California

Tumbleweeds (Salsola species, section Kali) are road side and rangeland pest plants throughout the 48 contiguous states in the US. Three described tumbleweed species and two undescribed Salsola taxa occur in California. The known species are Russian thistle, Salsola tragus, introduced from Eurasia in the 1800s, Russian barbwire thistle, S. paulsenii, which grows in the desert regions of California, and is also native to Eurasia, and the recently identified S. kali subspecies austroafricana, possibly native to South Africa. Our goals were to investigate karyology, genome size, and molecular genetic affinities of the described species and the other taxa within their ranges in California using recently developed microsatellite loci, dominant nuclear DNA markers (RAPD and ISSR), and DNA sequence data. Chromosome counts and genome size assessments made with flow cytometry were compared. These analyses indicated that one undescribed taxon is a new allopolyploid hybrid between S. tragus and S. kali subspecies austroafricana, and the other undescribed taxon appears to be a complex hybrid involving all three described species. The invasion potentials for the hybrid taxa are unknown. Tumbleweeds are the focus of biological controls efforts but the identification of suitable agents for the hybrid taxa may be problematic because of the large amount of genetic variability encompassed within this evolving Salsola complex.     

TAXONOMY AND ULTRASTRUCTURE OF GOMPHONEIS MESTA SP. NOV. (BACILLARIOPHYTA), A NEW EPILITHIC DIATOM FROM THE MESTA RIVER,BULGARIA1

A new species from the diatom genus Gomphoneis, G. mesta, was described following morphological and ultra-structural analyses of its frustules by light and scanning electron microscopy. The valve striation consisting of double rows of areolae and the presence of longitudinal lines dictate the systematic position of this taxon in the genus Gomphoneis. The set of features that distinguishes G. mesta from allied taxa is small valves, 26.5–81.6 μm in length and 7–12.5 μm in width, lanceolate to clavate shape, with a broadly lanceolate axial area, and longitudinal lines positioned next to the valve margins, thus contributing to a general thickening of the valve outline. When compared to other members of the genus, G. mesta most resembles G. magna Kociolek & Stoermer. Gomphoneis mesta dominates the epilithic community, growing in the Cherna Mesta, a pristine high-mountainous tributary of the Mesta River in Bulgaria. The species has been also found as small populations in all other benthic habitats of the Cherna Mesta and in a few locations along the Mesta River.     

A phylogenetic analysis of Pisum based on morphological characters, and allozyme and RAPD markers

Cladistic analyses of 17 wild and cultivated pea taxa were performed using morphological characters, and allozyme and RAPD (random amplified polymorphic DNA) markers. Both branch-and-bound and bootstrap searches produced cladograms that confirmed the close relationships among the wild species and cultivars of Pisum proposed by a variety of systematic studies. Intraspecific rankings were supported for northern P. humile, southern P. humile, P. elatius and P. sativum, which together comprise a single-species complex. P. fulvum, while clearly the most divergent of the pea taxa, could also be assigned to the same species complex without violating the hierarchial logic of the cladogram. Its inclusion or exclusion depends on whether the level of interfertility it displays with other pea taxa or its overall morphological and chromosomal distinction are emphasized. As suggested by previous studies, northern P. humile was the most likely sister taxon to cultivated P. sativum; although, rigorous phylogenetic evaluation revealed a close genealogical affinity among P. elatius, northern P. humile and P. sativum. Despite their limited number, the 16 morphological characters and allozyme markers used precisely organized the pea taxa into established taxonomic groupings, perhaps in part reflecting the role morphology has played historically in pea classification. The RAPD data also generally supported these same groupings and provided additional information regarding the relationships among the taxa. Given that RAPDs are relatively quick and easy to use, are refractory to many environmental influences, can be generated in large numbers, and can complement traditional characters that may be limited in availability, they provide a valuable new resource for phylogenetic studies.     

Foliar micro‐morphology of Carex sect. Phacocystis in northern Europe

A micro‐morphological analysis of leaf epidermis (adaxial and abaxial sides) of fifteen taxa of Carex section Phacocystis was carried out using light microscopy and scanning electron microscopy (SEM). Three taxa were epistomatic (C. nigra var. nigra, C.nigra var. juncea and C. subspathacea), seven taxa were hypostomatic (C. acuta, C. bigelowii ssp. bigelowii, C. bigelowii ssp. rigida, C. cespitosa, C. elata, C. lyngbyei and C. paleacea) and five amphistomatic (C. aquatilis, C.×halophila, C. rufina, C. stans and C. trinervis). Epidermal modifications such as prickles were present in many species. The micro‐morphological leaf characters of the investigated species were found to be important for distinguishing individual taxa but not for subsectional classification.     

Morphology, Biometry, and Taxonomy of Freshwater and Marine Interstitial Cyphoderia (Cercozoa: Euglyphida)

ABSTRACT. Good taxonomy is essential for ecological, biogeographical, and evolutionary studies of any group of organisms. Therefore, we performed detailed light- and scanning electron microscopy investigations on the shell ultrastructure and biometric analyses of the morphometric variability of five freshwater and marine interstitial testate amoebae of the genus Cyphoderia ( C. trochus var. amphoralis, C. ampulla, C. margaritacea var. major, C. compressa , and C. littoralis ), isolated from different populations in Bulgaria and Switzerland. Our aims were (1) to clarify the morphological characteristics of these taxa, and (2) to compare the morphology of a given taxon ( C. ampulla ) among different locations in Bulgaria and Switzerland as a first step towards an assessment of the geographical variation within a supposedly cosmopolitan taxon. Four of the studied taxa are characterized by a well-expressed main-size class and by a small size range of all the characters and can be defined as size-monomorphic species. Based on these results, the following systematic changes are proposed: C. major ( Penard, 1891 ) n. comb. (Syn.: C. margaritacea var. major ( Penard, 1891 ) and C. amphoralis ( Wailes & Penard, 1911 ) n. comb. (Syn.: C. trochus var. amphoralis ( Wailes & Penard, 1911 )). However, we also show significant morphological variability between the Swiss and Bulgarian populations of C. ampulla , suggesting the possible existence of more than one taxon within this species. Further studies are required to assess (1) if these two morphologically different taxa represent individual species, (2) if so, if more species exist, and if this diversity is due to limited distribution ranges (endemism) or if several closely related taxa occur together in different geographical areas.     

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.     

A phylogenetic hypothesis for Asilidae based on a total evidence analysis of morphological and DNA sequence data (Insecta: Diptera: Brachycera: Asiloidea)

An hypothesis of phylogenetic relationships of Asilidae and its constituent taxa is presented, combining morphological and DNA sequence data in a total evidence framework. It is based on 77 robber fly species, 11 Asiloidea outgroup species, 211 morphological characters of the adult fly, and approximately 7300 bp of nuclear DNA from five genes (18S and 28S rDNA, AATS, CAD, and EF-1α protein-encoding DNA). The equally weighted, simultaneous parsimony analysis under dynamic homology in POY resulted in a single most parsimonious cladogram with a cost of 27,582 (iterative pass optimization; 27,703 under regular direct optimization). Six of the 12 included subfamily taxa are recovered as monophyletic. Trigonomiminae, previously always considered as monophyletic based on morphology, is shown to be non-monophyletic. Two of the three Trigonomiminae genera, Holcocephala Jaennicke, 1867 and Rhipidocephala Hermann, 1926, group unexpectedly as the sister taxon to all other Asilidae. Laphriinae, previously seen in the latter position, is the sister group of the remaining Asilidae. Five other subfamily taxa, i.e. Brachyrhopalinae, Dasypogoninae, Stenopogoninae, Tillobromatinae, and Willistonininae, are also shown to be non-monophyletic. The phylogenetic relationships among the higher-level taxa are partly at odds with findings of a recently published morphological study based on more extensive taxon sampling. The total evidence hypothesis is considered as the most informative one, but the respective topologies from the total-evidence, morphology-only, and molecular-only analyses are compared and contrasted in order to discuss the signals from morphological versus molecular data, and to analyze whether the molecular data outcompete the fewer morphological characters. A clade Apioceridae+Mydidae is corroborated as the sister taxon to Asilidae.     

DEFINING TAXON BOUNDARIES IN MEMBERS OF THE MORPHOLOGICALLY AND GENETICALLY PLASTIC GENES CAULERPA (CAULERPALES,CHLOROPHYTA)1

Numerous attempts to capture the morphological variability of the genus Caulerpa have resulted in an unstable classification of numerous varieties and formae. In the present study we attempted to test taxon boundaries by investigating morphological and genetic variation within and between seven taxa of Caulerpa, supposedly belonging to four species, sampled at different sites in a Philippine reef system. Using both field and culture observations, we described the relation between the variability of a set of morphological characters and ecological parameters, such as wave exposure, light intensity, and substrate type. Statistical analyses showed that the limits between two (out of three) ecads of the C. racemosa (Forsskål) J. Agardh complex were obscured by the presence of morphological plasticity. Other studied taxa of Caulerpa (i.e. C. cupressoides [Vahl] C. Agardh, C. serrulata [Forsskål] J. Agardh, and two formae of C. sertularioides [S. Gmelin] Howe) could be grouped based on morphology despite the presence of morphological plasticity. Our results indicated a strong association between light intensity and several quantitative morphological variables. Genetic diversity of these taxa was assessed by partial sequencing chloroplast rbcL and tufA genes and the ycf10‐chlB chloroplast spacer. In all phylogenetic analyses, C. serrulata, C. cupressoides, C. sertularioides, and the three ecads of C. racemosa emerged as distinct genetic units. Despite the presence of morphological plasticity and morphological convergence, a subset of morphological characters traditionally used in taxonomic delimitation still had sufficient discriminative power to recognize the terminal phylogenetic clades.     

The marine algae of the Hebridean Island of Colonsay

Four specimens attributable to the freshwater red algal taxon, Ptilothamnion richardsii, were examined from locations in Australia and French Guiana, and partial rbcL gene sequence data were obtained. Phylogenetic analyses showed these specimens to be allied to taxa in the Batrachospermales. The two specimens from Australia were determined to be Batrachospermum antipodites, differing from that taxon by only 15?bp (1.2%). The two specimens from French Guiana were within the Batrachospermum section Contorta clade, but differed considerably (≥8%) from the taxa for which there are presently rbcL gene data. The sequence data clearly show these specimens to be chantransia stages of Batrachospermum taxa. Recently, other described freshwater Audouinella taxa have also been shown to be chantransia stages. Like previous authors, we argue that it is useful to continue to recognize entities with particular morphological characters and propose the form taxa Chantransia richardsii, C. macrospora and C. pygmaea. A key to these is provided.     

A new viviparous acoel Childia vivipara sp. nov. with observations on the developing embryos, sperm ultrastructure, body wall and stylet musculatures

A free‐living viviparous acoel, Childia vivipara sp. nov., from the Gullmar fjord of the Swedish coast is described. The new species is assigned to the taxon Childia based on histological, ultrastructural and molecular sequence similarities. All available molecular markers (18S rRNA, 28S rRNA and histone H3) and several morphological characters, obtained using transmission electron microscopy and confocal scanning laser microscopy of whole mount specimen stained with TRITC‐labelled phalloidin, support the placement of C. vivipara in the taxon Childia. Childia vivipara and other Childia species share the following morphological synapomorphies: well‐developed copulatory organs built of tightly packed stylet needles, proximal part of the stylet inserted into the seminal vesicle, reversed body‐wall musculature, absence of ventral diagonal muscles, presence of dorsal diagonal muscles, and presence of ventral straight longitudinal muscles between frontal pore and mouth, 9 + 1 sperm axoneme structure, six distal sperm cytoplasmic microtubules, and extensive overlap of axonemes and nucleus. The new species can be easily distinguished from other Childia species by its viviparous mode of reproduction and single curved stylet. Observations on late embryonic development based on the oldest developing embryos are discussed.     

A comparison of the pollen wall ultrastructure of aureoid and non-aureoidSenecio species (Asteraceae) in North America

The structure of the pollen wall as revealed by transmission electron microscopy is presented for 29 speciesSenecio including 19 aureoid and ten non-aureoid species. The previous report of a helianthoid wall structure (exine with internal foramina) is confirmed for all the aureoid species examined while all of the non-aureoid taxa showed a senecioid pattern. The data provide support for the treatment of the aureoid complex as a distinct taxon.     

Genetic and Morphological Analysis of Some European Species of the “coriaceus group” of Carabodes (Acari, Oribatida, Carabodidae) and Description of C. tyrrhenicus sp. nov.

The oribatid mite Carabodes reticulatus Berlese, 1913 is redescribed from typical and topotypical material. A new species, C. tyrrhenicus, widespread in Sardinia and other Tyrrhenian islands, is also described on the basis of morphological evidence. Scanning electron microscopy was used to study intra- and inter-specific variation, in order to determine diagnostic characters. The morphology-based taxonomy was tested through allozyme analysis of the two species and of two closely related species, namely C. coriaceus C. L. Koch, 1836 and C. arduinii Valle, 1955. Allozyme data are largely congruent with morphological evidence, indicating that the four entities represent well-differentiated evolutionary lineages. Molecular results show large inter-specific genetic differentiation, suggesting that these taxa arose from ancient cladogenetic events.     

The curse of taxonomic uncertainty in biogeographical studies of free-living terrestrial protists: a case study of testate amoebae from Amsterdam Island

Aim A current debate in microbial biogeography contrasts two views concerning the distribution of free‐living microorganisms. The first view assumes a ubiquitous distribution, while the second assumes that at least some species have limited geographical distributions. We tested for limited geographical distributions by identifying testate amoebae morphospecies from an extremely remote oceanic island where the potential for endemism is high. Location Amsterdam Island, Indian Ocean. Methods Sixty moss and water samples collected from the top of the volcano to the lowland were investigated for their testate amoeba content. Due to taxonomic uncertainties among the Argynnia (Nebela) dentistoma species complex (including A. antarctica), we also performed light and scanning electron microscopy investigations on the shell ultrastructure and biometric analyses on several specimens of this taxon. Results We identified a total of 43 testate amoeba taxa belonging to 15 genera. Only four testate amoeba taxa had previously been recorded on this island. Testate amoeba communities of Amsterdam Island are dominated by cosmopolitan ubiquitous euglyphid taxa such as Trinema lineare, Assulina muscorum and Corythion dubium. The length and width ranges for Argynnia dentistoma on Amsterdam Island overlap with other records of this species and of A. antarctica, suggesting that A. antarctica is not a distinct taxon. Main conclusions Although Amsterdam Island is among the most remote islands in the world, an extensive inventory of testate amoeba morphospecies provided no clear evidence for endemism. On the one hand, our detailed morphometric analysis of the A. dentistoma complex revealed that A. antarctica, a morphospecies previously suggested to display endemism, cannot be confidently distinguished from the cosmopolitan morphospecies A. dentistoma. On the other hand, five morphotaxa could not be identified with certainty and might represent new species, potentially with limited distribution. These examples illustrate how taxonomic uncertainties undermine biogeographical studies of testate amoebae. In order to allow better interpretation of morphology‐based testate amoeba distribution data, an assessment of genetic diversity among and within morphotaxa in relation to geographical distance for some common testate amoebae should be given high priority.