An evaluation of biostratigraphic markers across multiple geological sections in the search for the GSSP of the base of the Serpukhovian Stage (Mississippian)

This paper aims to evaluate potential biostratigraphic markers for the Viséan–Serpukhovian boundary in sections of Europe and Asia, to help identify the base of the global Serpukhovian Stage, which is a high priority task for Carboniferous biostratigraphy. Sections in the Serpukhovian stratotype area in the Moscow Basin contain a gap at the base of the Tarusian Regional Substage (basal in the classical Serpukhovian), so the traditional boundary defined in these sections cannot be precisely correlated with other successions worldwide. The IUGS Task Group to establish a GSSP close to the traditional Viséan–Serpukhovian boundary focused on the search for a new boundary marker, primarily on the first appearance datum (FAD) of the conodont Lochriea ziegleri in the lineage Lochriea nodosa to Lochriea ziegleri, considering it to be a suitable biostratigraphic event. The FOD (first occurrence datum) of L. ziegleri has been recognized in many successions worldwide, although only in a few sections the supposed evolutionary lineage of L. ziegleri was inferred. There are serious impediments to the FAD of L. ziegleri being universally accepted as the boundary marker. This paper presents a review of the FOD levels of L. ziegleri documented so far from multiple sections along with other correlatable markers (foraminifers and ammonoids) that can serve as additional points of reference in sections where a conodont record is poor or absent. The reviewed sections are Naqing Section (South China), Verkhnyaya Kardailovka and Kugarchi sections (South Urals, Russia), Mariinsky Log and Ladeinaya Mountain Sections (western slope of the Middle Urals, Russia), Novogurovsky Section (Moscow Basin, Russia), Vegas de Sotres Section (Cantabrian Mountains, Spain), Lugasnaghta Section (County Leitrim, Ireland), Wenne River Bank Section (Germany), and Milivojevića Kamenjar Section (Družetić, NW Serbia). We also included a compilation of data from sections of North England and southern Scotland. In this paper, we will mainly focus on newly described sections, while the discussion of most previously described sections was summarized by Nikolaeva et al. (2001, 2002, 2005, 2009b) and other publications, so they are only briefly mentioned in this review. It should be added that there is no such a thing as a perfect GSSP section, as each section has certain disadvantages, either lithological, paleontological, or both, so it is important to hear and discuss all the different opinions to develop the optimum strategy for future research. In addition, we analyze published records from several sites in North England and southern Scotland. We discuss the first appearances of the ammonoid genera Cravenoceras, Edmooroceras, Lyrogoniatites, Dombarites, and Platygoniatites, the foraminifers Neoarchaediscus postrugosus, Hemidiscopsis muradymica, H. hemisphaerica, species of Janischewskina and Monotaxinoides, Eostaffella pseudostruvei group, Eostaffellina decurta, and Endothyranopsis plana. We publish here for the first time the useful accounts of foraminifers and conodonts from the Mariinsky Log Section and Ladeinaya Mountain Section (Middle Urals, Russia), and re-figure several important type specimens from Europe and the Urals.     

Molecular phylogenetics and the evolution of fruit and leaf morphology of Dichaea (Orchidaceae: Zygopetalinae)

Background and Aims

The orchid genus Dichaea, with over 100 species found throughout the neotropics, is easily recognized by distichous leaves on long stems without pseudobulbs and flowers with infrastigmatic ligules. The genus has previously been divided into four sections based primarily on presence of ovary bristles and a foliar abscission layer. The aim of this work is to use DNA sequence data to estimate phylogenetic relationships within Dichaea and map the distribution of major morphological characters that have been used to delimit subgenera/sections.

Methods

Sequence data for the nuclear ribosomal internal transcribed spacers and plastid matK, trnL intron, trnL-F spacer and ycf1 for 67 ingroup and seven outgroup operational taxonomic units were used to estimate phylogenetic relationships within Dichaea. Taxa from each of the four sections were sampled, with the greatest representation from section Dichaea, the most diverse and taxonomically puzzling group.

Key Results

Molecular data and morphology support monophyly of Dichaea. Results indicate that section Dichaeopsis is polyphyletic and based on symplesiomorphies, including deciduous leaves and smooth ovaries that are widespread in Zygopetalinae. There are at least three well-supported clades within section Dichaeopsis. Section Pseudodichaea is monophyletic and defined by setose ovaries and leaves with an abscission layer. Sections Dichaea and Dichaeastrum are monophyletic and defined by pendent habit and persistent leaves. Section Dichaeastrum, distinguished from section Dichaea primarily by a glabrous ovary, is potentially polyphyletic.

Conclusions

The leaf abscission layer was lost once, occurring only in the derived sections Dichaea and Dichaeastrum. The setose fruit is a more homoplasious character with several losses and gains within the genus. We propose an informal division of the genus based upon five well-supported clades.Key words: Dichaea, matK, nrITS, Orchidaceae, trnL intron, trnL-F spacer, ycf1, Zygopetalinae     

The systematic value of nuclear DNA content for all species of Narcissus L. (Amaryllidaceae)

The taxonomy of all species of Narcissus (Amaryllidaceae), an important horticultural crop, has not been investigated recently. As a new approach, genome size was determined by flow cytometry with propidium iodide from 375 accessions. The somatic nuclear DNA contents (2C) were shown to range from 14 to 38 pg for the diploids. Narcissus assoanus and N. gaditanus are, based on their nuclear DNA content, removed from section Apodanthi and placed in a new section Juncifolii. The different ploidy levels and species involved were entangled for N . “fernandesii” s.l. and a new allotetraploid form is named here. Section Pseudonarcissus was much more heterogeneous in nuclear DNA content than expected. Sixty-five accessions of N. pseudonarcissus possessed, with 23.7 pg, similar amounts of DNA. However, several species from this section were clearly distinctive in nuclear DNA content. It runs from the diploid N. primigenius with 21.7 pg to the also diploid N. nevadensis with 38.2 pg. Also N. abscissus and N. moleroi are with about 26 pg clearly different from N. pseudonarcissus. For the first time, in 11 accessions, hexaploidy was found in N. pseudonarcissus ssp. bicolor. A new section Nevadensis with 30–39 pg of nuclear DNA was split off from the section Pseudonarcissus with now 21–27 pg. A nonoploid N. dubius with 96.3 pg has by far the highest amount of nuclear DNA and can be calculated to have the highest ploidy ever reported in Narcisssus. The total number of Narcissus species was determined as 36, nine more than in Flora Europaea and they were divided up in two subgenera and 11 sections. Flow cytometry is shown to produce easily obtainable and original systematic data that lead to new insights. Genome size or C-value turns out to be one of the most salient features to define the status of the species in the genus Narcissus.     

Aspergillus yunnanensis,a new and rare species in the Aspergillus section Jani

Aspergillus is a monophyletic genus comprising the subgenera Aspergillus, Circumdati, Cremei, Fumigati, Nidulantes and Polypaecilum. The subgenus Circumdati contains many economically important species and mycotoxin producers. Section Jani was recently introduced with morphological and molecular support. In the present study, two strains isolated from farmland soil were assigned in section Jani based on multi-locus phylogenetic analyses but showed low similarity with existing species. Further morphological observation found they had wider vesicles and conidia connections which were different from the known species. Based on phylogenetic and morphological data, Aspergillus yunnanensis was introduced as the third species in section Jani. Members in section Jani are rarely distributed, this is the first report of this section in China.     

An expanded phylogeny of Cuphea (Lythraceae) and a North American monophyly

A phylogenetic analysis of the New World genus Cuphea was conducted employing sequences from the nuclear rDNA internal transcribed spacer (ITS) and chloroplast trnL-trnF spacer and rpl16 intron. The analysis expands the number of Cuphea species from 53 in an earlier ITS study to 70 and adds two chloroplast data sets in order to generate a more complete and robust phylogeny and to test a previous result that suggested the presence of a large North American clade. Results reaffirm the monophyly of Cuphea with Pleurophora as the sister genus and recover a basal divergence event that mirrors the two subgenera of the current classification. Phylogenies of the two chloroplast regions are largely unresolved beyond the initial dichotomy and some resolution at the terminal and subterminal nodes. Based on the ITS phylogeny, five major clades are recognized. Subgenus Cuphea (Clade 1), defined morphologically by the synapomorphic loss of bracteoles, is sister to the much larger subg. Bracteolatae (Clades 2–5). Clades 2–4, comprising the South American and Caribbean species, grade successively to Clade 5, an exclusively North American lineage of 29 species. Among the 12 sections included in the study, only section Trispermum, a subclade of Clade 4, is monophyletic. Section Pseudocircaea is nested within Clade 3, which is largely equivalent to section Euandra. The North American endemic clade includes four sections, of which none are recovered as monophyletic in this study.     

Systematics of Fraxinus (Oleaceae) and evolution of dioecy

Phylogenetic relationships among 40 of the 43 recognized species of Fraxinus L. (Oleaceae) were estimated on the basis of 106 nuclear ribosomal ITS sequences. ITS trees resulting from maximum likelihood (ML), maximum parsimony (MP) and Bayesian inference (BI) are congruent and identify six distinct lineages. These clades allow establishing sections with high molecular and morphological support. The basal resolution generally has low ML bootstrap and MP jackknife support, but the Bayesian posterior probabilities are high for certain relationships. An independent data set of combined sequences from the chloroplast rps16 and trnL-F regions contains few informative sites but corroborate most of the relationships in the ITS tree. The molecular phylogeny is discussed in the light of morphological and other data and a revised infrageneric classification with six sections are presented. The subgenera and subsections are abandoned and the section Pauciflorae is a new combination. Fraxinus quadrangulata and Fraxinus anomala are united with Fraxinus dipetala in the section Dipetalae and Fraxinus platypoda is transferred to the section Fraxinus. Fraxinus chiisanensis, Fraxinus spaethiana and Fraxinus cuspidata are treated as incertae sedis. A sectional key is given, together with a systematic list of the 43 recognized species, with common synonyms and distribution. Breeding system and other traits mapped on the phylogeny show that dioecy has three separate origins, and in each case followed after the transition from insect to wind pollination. In one instance dioecy evolved from hermaphroditism via androdioecy and twice via polygamy.     

Biosystematic study in the genus Cousinia Cass. (Asteraceae), section Cousinia

The genus Cousinia Cass. (Family Asteraceae) with about 600–700 species is distributed in central and western Asia. Iran has more than 200 Cousinia species that grow in mountainous areas. Cousinia is not monophyletic and with the genus Arctium, comprise the Arctium-Cousinia complex. The Cousinia species are distributed in 70 sections. The species relationship in the sect. Cousinia is not known, moreover, in few cases, inter-specific hybrids and intermediate forms are recognized. Therefore, we undertook a detailed morphological, micro-morphological and molecular (Cp-DNA and ISSRs) study of the section Cousinia with the following objectives: 1- to delimitate the species, 2- to identify potential intermediate forms, 3- to investigate the species inter-relationship. The present study revealed that combination of morphological, micro-morphological and ISSR data can delimit the species. We did not encounter any intermediate forms throughout the studied area. AMOVA and STRUCTURE analysis revealed that the species of this section are genetically differentiated but have some degree of shared common alleles. Reticulation analysis by T-REX also showed some degree of gene flow for Cp-DNA. This low degree of genetic and morphological variability in Cousinia, can help the species to cope with environmental variability they face.     

Phylogenetic comparison among the heterocystous cyanobacteria based on a polyphasic approach

Phylogenetic comparison has been done among the selected heterocystous cyanobacteria belonging to the sections IV and V. The hierarchical cluster analysis based on antibiotics sensitivity showed a distant relationship between the members of Nostocales and Stigonematales. Thus, multiple antibiotic resistance pattern used as marker provide easy, fast, and reliable method for strain discrimination and genetic variability. However, morphological, physiological (both based on principal component analysis) and biochemical analysis grouped true branching cyanobacteria along with the members of section IV. Molecular analysis based on 16S rRNA gene sequences revealed that Hapalosiphon welwitschii and Westiellopsis sp. were grouped in cluster I whereas Scytonema bohnerii, a false branching genera showed a close proximity with Calothrix brevissima in cluster II. Cluster III of clade 2 included Nostoc calcicola and Anabaena oryzae which proved the heterogeneity at the generic level. Cluster IV the largest group of clade 2 based on 16S rRNA gene sequences includes six strains of the genera Nostoc, Anabaena, and Cylindrospermum showing ambiguous evolutionary relationship. In cluster IV, Anabaena sp. and Anabaena doliolum were phylogenetically linked by sharing 99% sequence similarity. Probably, they were of the same genetic makeup but appear differently under the diverse physiological conditions. Section IV showed polyphyletic origin whereas section V showed monophyletic origin. Results suggested that either morphological or physiological or biochemical or molecular attribute is not sufficient to provide true diversity and phylogeny of the cyanobacteria at the generic level and thus, a polyphasic approach would be more appropriate and reliable.     

The Guadalupian–Lopingian boundary (Permian) in a pelagic sequence from Panthalassa recognized by integrated conodont and radiolarian biostratigraphy

Guadalupian–Lopingian sedimentary rocks are widely distributed in accretionary complexes in Japan, but the Guadalupian–Lopingian boundary (G–LB) is not well documented from these pelagic sediments. To identify the G–LB and to better correlate an extinction event that occurred around the Guadalupian–Lopingian boundary, we examined the conodont biostratigraphy from a Permian pelagic chert sequence in the Gujo-hachiman section, Gifu, southwest Japan. Age-diagnostic conodonts, including Clarkina postbitteri postbitteri, were found in this section. The biostratigraphic occurrences of these age-diagnostic conodonts can pinpoint the “G–L transitional zone” in the Gujo-hachiman section by comparison with well-studied sections from south China, including the GSSP section. The transitional zone was recognized by the first occurrence horizons of both Clarkina postbitteri hongshuiensis and C. p. postbitteri. The G–LB has been placed at or above the first occurrence horizon of the radiolarians Albaillella yamakitai or Albaillella cavitata in previous studies from China and Japan. We detected the first occurrence horizon of A. yamakitai below the base of the “G–L transitional zone,” in the Upper Capitanian. The conodont biostratigraphy is consistent with the radiolarian biostratigraphy in this section, which can be correlated to relevant sections in China.     

Analysis of genetic diversity and sectional relationships in Musa using AFLP markers

The AFLP technique was used to assess the genetic diversity and sectional relationships in 39 accessions representing the four main sections of the genus Musa. Eight AFLP + 3 primer pairs produced 260 polymorphic bands that were used in cluster and PCO analysis. A wide range of variability was observed among the species within the sections of the genus Musa. AFLP data was useful in separating the different sections of the genus as well as differentiating the different genomic groups of section Eumusa. Section Rhodochlamys (x = 11) appeared as a distinct entity and clustered closely with the Musa acuminata Colla complex of section Eumusa that has the same basic chromosome number. This relationship is congruent with previous studies. However, unlike previous proposals that questioned the identity of Rhodochlamys as a separate taxonomic unit, PCO analysis of the AFLP data showed that it is a distinct entity. Musa laterita Cheesman (Rhodochlamys) and Musa schizocarpa Simmonds clustered with the M. acuminata complex suggesting that they may be sources of useful genes for the improvement of the cultivated bananas. Callimusa formed a distinct unit and was closer to Australimusa than to the other sections. Although both sections share the same basic chromosome number of x = 10 these sections are genetically distinct     

Age and growth of Siberian sculpin (Cottus poecilopus) and young brown trout (Salmo trutta) in a subalpine Norwegian river

Age, growth and density of Siberian sculpin (Cottus poecilopus) and young brown trout (Salmo trutta) within two sections of River Atna; above Lake Atnsjøen [Section 1 at altitudes between 739 and 715 m] and below Lake Atnsjøen [Section 2 at altitudes between 430 and 370 m] was studied during a 6-year period (1986–91). The water temperature was considerably lower in Section 1 than in Section 2, as the number of days with a water temperature above 10?°C (T _{D ≥ 10?° C} ) from spring to August 1 ranged between 2–26 and 26–52 days, respectively. Juvenile brown trout (age 0+) attained a significantly smaller body size in Section 1 than in Section 2; mean length ±SD was 35 ± 8 mm (ranged 27–46) and 43 ± 7 mm (range 38–46), respectively. In Section 2, there was a highly positive correlation between the body length of 0+ brown trout and mean water temperature in June (p<0.005), and also to some extent in Section 1 (p=0.11). Individuals of age 1+ did not exhibit any such difference, while fish in age group 2+ were larger in Section 1 than in Section 2. By using the number of days with a water temperature between the range 5–10?°C (T _{D ≥ 5 ? 10?° C} ) as test variables, we found a highly positive correlation between the August 1 body length of 0+ brown trout and T _{D ≥ 9}?°C from spring to August 1 in Section 2 (p<0.05), as opposed to T _{D ≥ 7}?°C for trout in Section 1 (p=0.11). Young Siberian sculpin (age 0+ and 1+) also exhibited slower growth in Section 1 than in Section 2, but this was not the case among older specimens. In the year with the lowest temperature measured (1987), no 0+ Siberian sculpin were caught in any of the two sections, indicating that low temperature affects their survival. Both species exhibited large spatial and temporal variation in density. Thus, data on abundance and growth sampled on one occasion at one site can not be regarded as representative for these two fish populations.     

Seed proteins and systematics of Tripsacum

The genus Tripsacum (Gramineae) is distributed between the latitude 42°N and 24°S in the New World. It is divided into two sections. Section Tripsacum includes 11 species with T. dectyloides (L) L. extending across the range of the genus. Section Fasciculata includes five Meso-American species with T. lanceolatum Rupr. ex Fourn. extending into southern Arizona. The genus displays considerable diversity in seed proteins. Variation patterns are of limited use in distinguishing sections, but are species and habitat specific. Protein data are particularly useful in subspecific classification, and consistently distinguish diploid from polyploid races of T. zopilotense Hern. and Randolph and T. bravum Gray. The tetraploid ectotype of T. bravum deserves specific rank and the robust ecotype of T. dactyloides var. meridonate de Wet and Timothy deserves varietal rank.     

PHYLOGENETIC IMPLICATIONS OF DIFFERENCES IN NUMBER OF PLASTID PHOSPHOGLUCOSE ISOMERASE ISOZYMES IN NORTH AMERICAN COREOPSIS (ASTERACEAE: HELIANTHEAE: COREOPSIDINAE)

Enzyme electrophoresis was employed to ascertain the number of loci encoding plastid phosphoglucose isomerase (PGI) in species representing all sections of North American Coreopsis. Several species from each of the closely related genera Bidens, Coreocarpus, Cosmos, and Thelesperma were also examined. Species in nine of the 11 sections of North American Coreopsis have two isozymes for plastid PGI, and nearly all species examined in the four other genera also have two (one species has three) isozymes. Since most diploid vascular plants have one plastid PGI isozyme, a gene duplication probably occurred in an ancestor that is common to Coreopsis and the other four genera. That is, two isozymes represent the ancestral number for Coreopsis. The two sections (Electra and Anathysana) apparently lacking the duplication are closely related woody plants restricted largely to Mexico. One gene encoding plastid PGI ostensibly was silenced in a common ancestor of these two sections. This is concordant with other data suggesting a close relationship between the two sections, i.e., they appear to represent a monophyletic group. The electrophoretic data also indicate that 1) the enigmatic monotypic section Silphidium is more closely related to eastern North American sections and not derived from section Electra; and 2) section Anathysana is not ancestral to the three California sections Leptosyne, Pugiopappus, and Tuckermannia; rather, it represents a terminal element closely related to and possibly derived from section Electra.     

Age and distribution of the Late Devonian brachiopod genus Dzieduszyckia Siemiradzki, 1909 in southern China

Conodont biostratigraphical work was done at four sections recently found with occurrence of the rhynchonellide brachiopod genus, Dzieduszyckia Siemiradzki, in southern Guangxi and in the border area between Dushan County of Guizhou and Nandan County of Guangxi, South China. These sections represent two different types of facies, i.e., carbonate platform and intraplatform basin. The conodont analysis reveals that this genus occurs in the Upper triangularis Zone and the Middle crepida Zone at the Dazhai Section, through the Lower to Upper crepida zones at Dalong, and is restricted within the Upper rhomboidea Zone at the two intraplatform basin sections (Changtang and Duli). This result demonstrates that the occurrence of these peculiar rhynchonellide brachiopods in South China, regardless of the depositional environments, is within the Lower Famennian instead of the previously suggested Upper Famennian. Furthermore, this brachiopod genus in South China began to inhabit on the carbonate platform almost since the beginning of the Famennian and did not extend to the intraplatform basin facies until the late Early Famennian.Available biostratigraphic data indicate that during the Early and Middle Famennian, Dzieduszyckia is widely distributed not only in South China, but also throughout the world, such as Morocco and southern Ural.Observation on the new collections from the four studied sections reveals that the peculiar rhynchonellide brachiopods have a great morphological variation within each section. Significant differences existed among the collections from different sedimentary settings and localities, probably reflecting the environmental and geographic constraint on the morphology of Dzieduszyckia. Samples from different layers in the same section have nearly identical morphological variation, suggesting the temporal inheritance in morphology of the rhynchonellide brachiopod.     

Conodont biostratigraphy and carbon isotope chemostratigraphy of the Middle to Upper Ordovician on the western Yangtze Platform,South China

Middle and Late Ordovician strata in the nearshore area of the western Yangtze Platform are investigated. In total, 241 δ¹³C_carb samples and 108 conodont samples from the Songliang and Tuanjie sections in the northeastern Yunnan Province were collected for chemostratigraphic and biostratigraphic studies. Altogether, 21 species belonging to 16 conodont genera were recovered from the two sections, among which Ansella jemtlandica, Drepanoistodus basiovalis, Hamarodus brevirameus, and Microzarkodina hagetiana are important age indicators within the Dapingian to early Katian interval. Four carbon isotope shifts are recognized in the studied sections: a negative shift in the early Darriwilian, the middle Darriwilian carbon isotope excursion (MDICE), the early Katian Guttenberg carbon isotope excursion (GICE), and a further positive carbon isotope shift later in the early Katian which is a new record of this event in South China. Integrated carbon chemostratigraphy and conodont biostratigraphy indicate early Darriwilian and early Katian ages for the base and top, respectively, of the Huadan Formation at the Songliang section, and a Darriwilian to early Katian age for the Shihtzupu Formation at the Tuanjie section.     

The systematics ofLinum sect.Linopsis (Linaceae)

SectionLinopsis of the genusLinum is the largest and probably the most widespread of the five commonly recognized sections. Using a number of traits, but especially heterostyly, style union, development of false septa in the fruit, pollen morphology, and chromosome number, together with petal pigmentation, number of floral parts, and stigma shape, the section is divided into five subsections:Dichrolinum, with one series;Halolinum with one;Linopsis, with four;Keniense, with one; andRigida, with two. Of these, subsect.Keniense and two series are new, the remaining four subsections and four series are published in a new rank. The inferred relationships of all of the groups and the general distribution of each are discussed. The section ranges from southern Europe and the Mediterranean region to India, eastern and southern Africa and throughout much of North and South America. The primitive species of the section are believed to be in the Mediterranean region.