Revision of the genus Fosterella (Bromeliaceae)

Die vorliegende Arbeit stellt eine umfassende taxonomische Revision der Gattung Fosterella L.B. Sm. (Bromeliaceae) dar, die alle 31 derzeit akzeptierten Arten umfasst und einen Bestimmungsschlüssel für diese beinhaltet. Die Revision beruht auf der morphologisch-anatomischen Auswertung von Herbarmaterial (über 800 Exsikkate), Lebendpflanzen (ca. 150 Akzessionen) und eigenen vergleichenden Untersuchungen im Freiland. Die Gattung Fosterella ist seit nunmehr etlichen Jahren Forschungsgegenstand einer interdisziplinären Studie, die sowohl molekulrae, als auch anatomische, morphologische und biogeographische Untersuchungen einbezieht. Unser Interesse an der Gattung Fosterella gründet sich auf ihrer enormen ökologischen und biogeographischen Vielfalt, sie gilt als hervorragendes Modellsystem für Artbildungsmechanismen in den Anden. In den letzten Jahren wurde von verschiedenen molekularen Methoden Gebrauch gemacht, um die verwandtschaftlichen Beziehungen innerhalb der Gattung zu untersuchen, so dass mittlerweile gut aufgelöste Stammbäume vorliegen. Diese molekularen Studien, überwiegend durchgeführt von Dr. Martina Rex, wurden ergänzt durch intensive Sammelaktivitäten und eingehende taxonomische Untersuchungen im Rahmen der vorliegenden Revision. Auf diese Weise konnten die morphologische Plastizität der einzelnen Arten erfasst und schließlich ein wohlfundiertes Artkonzept vorgelegt werden. Zunächst wird ein kurzer Überblick über die Familie der Bromeliaceen als auch die Gattung Fosterella gegeben, in dem jeweils Informationen zur Verbreitung, Morphologie, Physiologie, Ökologie und Phylogenie geliefert werden. Im Anschluss an einen historischen Überblick des taxonomischen Werdegangs wird die Abgrenzung der Gattung Fosterella zu den nächstverwandten Gattungen Deuterocohnia, Dyckia und Encholirium erläutert. Die morphologischen Merkmale zur Differenzierung der Arten innerhalb der Gattung werden im Hinblick auf ihre Zuverlässigkeit und ihr Gewicht diskutiert. Der Artschlüssel basiert auf Merkmalen, die leicht auszumachen und gut zu unterscheiden sind. Bei der ausführlichen Beschreibung der Arten wird auch auf ihre jeweilige Verbreitung, Ökologie, taxonomische Abgrenzung, systematische Verwandtschaft sowie die Etymologie des Namens eingegangen. Beigefügt sind jeweils Zeichnungen, ein Foto vom Holo-/Lectotypus, Fotos von Lebendpflanzen sowie eine Verbreitungskarte. Im Rahmen der taxonomischen Arbeit wurden fünf Arten zu Synonymen reduziert: Fosterella chiquitana Ibisch, R. Vásquez & E. Gross und F. latifolia Ibisch, R. Vásquez & E. Gross wurden in die Synonymie von F. penduliflora (C.H. Wright) L.B. Sm. eingezogen; F. fuentesii Ibisch, R. Vásquez & E. Gross als Synonym zu F. albicans (Griseb.) L.B. Sm. gestellt; F. elata H. Luther in die Synonymie von F. rusbyi (Mez) L.B. Sm. verwiesen und F. nowickii Ibisch, R. Vásquez & E. Gross als Synonym zu F. weddelliana (Brongn. ex Baker) L.B. Sm. gestellt. Fosterella schidosperma (Baker) L.B. Sm. var. vestita L.B. Sm. & Read wird zum Synonym von Fosterella weberbaueri (Mez) L.B. Sm. reduziert. Sechs Arten wurden neu beschrieben: Fosterella batistana Ibisch, Leme & J. Peters; F. christophii Ibisch, R. Vásquez & J. Peters; F. elviragrossiae Ibisch, R. Vásquez & J. Peters; F. kroemeri Ibisch, R. Vásquez & J. Peters; F. nicoliana J. Peters & Ibisch und F. robertreadii Ibisch & J. Peters. Das Taxon F. gracilis (Rusby) L.B. Sm. wurde neu etabliert. Um die Evolution von einzelnen morphologischen Merkmalen zu rekonstruieren, wurden die Zustände von zehn ausgewählten Merkmalen kodiert und auf einen molekularen Stammbaum kartiert. Die folgenden Merkmalszustände wurden als ursprünglich innerhalb der Gattung ermittelt: Stammlosigkeit, ganzrandige Blattspreiten, flache Rosetten mit dem Boden aufliegenden Blättern, locker beschuppte Blattunterseiten, schildförmige Haare mit gezähntem Rand, ganzrandige Pedunkel-Brakteen, rispenförmiger Blütenstand, kahle/verkahlende Blütenstandsachsen, weiße Petalen und einfach-aufrechte Narben. Rückschlüsse bezüglich der Evolution und Ausbreitung der Gattung Fosterella werden diskutiert: Die überwiegend kleinen Verbreitungsgebiete der Arten hängen offensichtlich mit ihren fragmentierten, inselartigen Habitaten (z.B. innerandine Trockentäler) zusammen. Die Tatsache, dass die Yungas-Bergregenwälder des Departamento La Paz, Bolivia, die Region mit der größten Artenvielfalt darstellen, lässt sich mit der extrem variablen Topographie und der außerordentlich hohen Vielfalt an Habitaten dieser Region erklären. Aus folgenden Gründen erscheint es sehr wahrscheinlich, dass die Gattung Fosterella ihren Ursprung im Tiefland hat: Die Mehrheit der Arten weist einen eher mesophytischen Habitus auf und ist in mehr oder weniger humiden Habitaten zu finden. Die Gattung ist durch mehrere Arten in sehr alten Habitaten des präkambrischen Schilds im Tiefland von Zentral-Südamerika vertreten. Weiterhin betreiben, soweit bekannt, alle Fosterella Arten C3 Photosynthese, während in den Gattungen der Schwestergruppe, Deuterocohnia, Dyckia and Encholirium, CAM der verbreitete Photosyntheseweg ist. In jedem Fall ist die Besiedelung der Anden und/oder Tieflandhabitate mehrfach unabhängig voneinander geschehen, vielleicht sogar in beiden Richtungen.

The Corticiaceae (Basidiomycetes) in Taiwan

Geographically, Taiwan is an Island and situated in the northeast of Asia, on the western side of the Pacific Basin, at the southeast of main China, south of Japan, and north of the Philippines. The main topographic character is the longitudinally oriented mountainous area. More than 200 peaks rise above 3000 m. They departed Taiwan into two lowland areas, an eastern and western plain. Taiwan is departed into subtropical (north area) and tropical zone (south area), which have a warm and humid climate, due to the Tropic of Cancer passing through. The average annual temperature in the lowland amounts to 28°C (7~38°C). The temperate climate also presents in the mountainous areas. The tropical typhoons usually come in summer and bring heavy rain, while the monsoon seasons have an important effect on the regional rainfall distribution. The mean annual rainfall of Taiwan is about 2600 mm (1000~6700 mm); the mountainous areas receive more rain than the lowlands. In Taiwan, according to different temperature and vegetation, the ecological environments were given rise to vertical biotic zonations, and form five major types: highland snowfield, highland meadow, coniferous forest, deciduous forest, and tropical forest. Six National Parks in Taiwan are located in the mountainous areas, in the north, the south, and on Jinmen Island. The National Parks represent about 8.4% of the country area. In this study, the collection sites are situated in Yangmingshan, Shei-Pa, Yushan, and Kenting National Park. Due to the island isolation, the proportions of endemic species are great in Taiwan, which also presents a high biodiversity. There are 4255 species of vascular plants including 1133 endemic. 5936 species in 1276 genera of fungi are hitherto reported in Taiwan. Among them, 233 Corticiaceae species were recorded, over one third (79 species) of them are known only from Taiwan. The first fungal report in Taiwan is about Phytophthora cyperi, published by the Japanese researcher T. Kawakami in 1904. Therefore, the history of research about fungi in Taiwan is more than one hundred years old. An eminent Japanese mycologist K. Sawada made an intensive survey from 1919 to 1959, and reported 2464 fungi species in his eleven volumes of “Descriptive Catalogue of Formosan Fungi”. However, only a few species (21 species in 9 genera) of Corticiaceae were recorded. From 1973, Chen and Lin resumed the study on Corticiaceae, and also some other foreign mycologists contributed for this field after 1980. The German research group lead by Franz Oberwinkler from Tübingen University collected in Taiwan several times. They published a number of new species and new records. Since 1989, S. H. Wu, a Taiwanese mycologist, has published a great amount of reports on corticioid fungi from Taiwan. Corticioid fungi were made up by the large and heterogeneous unnatural family Corticiaceae and other resupinate fungi belonging to other natural families in the Agaricomycetes. Molecular studies have shown that corticioid genera are distributed across all major clades of Agaricomycetes indicating that the corticioid fungi represent a polyphyletic group. They have resupinate fruitbodies and similar habitats. Species are characterized by simple fruitbody, more or less effused, and present smooth, porioid, grandinioid to odontioid hymenial surface. The fruitbodies are differently colored and usually soft to tough. Most of the Corticiaceae species are wood-saprobic organisms and gain the energy from the decomposing of wood-substrate such as cellulose or lignin. Materials for this study were collected by the author and other mycologists in Taiwan during surveys in April and May 1996, and March 2007, using the spring season with its high humidity and warm climate which are optimal conditions for the development of fungi. For assembling, the convenience sampling method was used in this study. This approach was chosen because it enables to detect a high biodiversity in a short time, and also to find species with rare or patchy distribution. The collecting sites from the North to the South include four National Parks and some preserved forests. They cover many different habitats such as low lands and high mountains. Fresh specimens were dried and analysed with a light microscope. 265 specimens belonging to Corticiaceae were studied in this research. Among them, 50 species in 21 genera including 11 new records and 10 new species were described with text and drawing. Four new species are belonging to Hyphodontia (H. sp. nov. 1, H. sp. nov. 2, H. sp. nov. 3, and H. sp. nov. 4), four to Schizopora (Sch. sp. nov. 1, Sch. sp. nov. 2, Sch. sp. nov. 3, and Sch. sp. nov. 4), one in Trechispora (T. sp. nov. 1), and one in Tubulicrinis (T. sp. nov. 1). Species recorded as new are Aleurodiscus amorphus, Botryohypochnus isabellinus, Hyphodontia cineracea, Hyphodontia palmae, Hypochnicium vellereum, Merulius tremellosus, Metulodontia nivea, Paullicorticium ansatum, Phlebia radiata, Phlebiella ardosiaca, and Xylobolus frustulatus. Besides, Botryohypochnus, Merulius, Metulodontia, Paullicorticium, and Xylobolus are also newly recorded genera in Taiwan. The genus Hyphodontia presents the highest diversity with 20 out of 50 species recorded. The second important genus is Hyphoderma, however with only 5 species. This indicates that Hyphodontia and Hyphoderma have a higher ability to develop in variable environments and approximately shows the predominance of these two genera in Taiwanese Corticiaceae. There are 11 new records out of the 50 species recorded, representing 22%. Some species, e.g. Hypochnicium vellereum and Paullicorticium ansatum were in the past recorded only in Europe and North America with cold and temperate climate. The samples of them are for the first time found in the subtropical belt, and display some difference from those of temperate regions. These collections should be molecularly investigated to clarify if they represent the same species of temperate areas. Patchily distributed species, for example Phlebiella ardosiaca, previously known only in Europe, and Hyphodontia palmae collected only in Brazil, were first recorded in different continents. Two possibilities are indicated by these new records: they are worldwide species but very rare to be found, or the Taiwanese specimens are taxonomically different. More survey from other continents and molecular study for these collections should be done in the future to solve this question. The distribution of Corticiaceae in Taiwan presents the variations in the north, central, and south areas and shows the diversity in lowlands and high mountains. The results of this study provide the evidence that the temperate Corticiaceae species displays a wider distribution. Subtropical and tropical taxa probably have also high dispersal capacities, and could possibly be found in the future in neighboring areas such as China, Japan, Korea or South Asia, but this needs further researches. In the total of 50 species, 10 new taxa were described in this study, giving about 20%. Some new species (e.g. Hyphodontia sp. 1, Hyphodontia sp. 2, and Hyphodontia sp. 3) are very similar to known species (Hyphodontia sambuci and Hyphodontia formosana), and the distinctive characters of Schizopora sp. nov. 1 are intermediate between those of Schizopora paradoxa and Hyphodontia flavipora. Thus, these small differences between the new and known species, suggest that the speciation occurred when the fungi migrated into Taiwan, due to the high diversity of environment, and amounts of the endemic plants. Taiwan is an intermediate place for the south (tropical) fungal species to migrate and adapt to north (temperate) regions. The middle and high altitude environments in Taiwan offer good conditions for the fungal speciation and possibly the occurrence of physiological changes to adapt to the temperate climate. Thus Taiwan has an important position for the biogeography of Asia mycobiota. 5936 known species in Taiwan represent about only 20% of the estimated number (24000) of Taiwanese fungal taxa. In this study, the findings (22% new records and 20% new species) indicate that amounts of unknown fungi species are expected in Taiwan. The lack of knowledge indicates that many new species are awaiting description, and fungal survey in Taiwan remains in a Pioneer phase. The last three wide surveys of Corticiaceae researches took place 20 years before this study (Chen & Lin 1977, Lin & Chen 1989, Wu 1990). After previous important contributions, the present taxonomic study comprising 21 genera is the most extensive on Corticiaceae of Taiwan.

Agricultural practices and horizontal nutrient balances in urban gardens and the alternative use of urban agricultural land in Khartoum, Sudan

The surge in the urban population evident in most developing countries is a worldwide phenomenon, and often the result of drought, conflicts, poverty and the lack of education opportunities. In parallel with the growth of the cities is the growing need for food which leads to the burgeoning expansion of urban and peri-urban agriculture (UPA). In this context, urban agriculture (UA) contributes significantly to supplying local markets with both vegetable and animal produce. As an income generating activity, UA also contributes to the livelihoods of poor urban dwellers. In order to evaluate the nutrient status of urban soils in relation to garden management, this study assessed nutrient fluxes (inputs and outputs) in gardens on urban Gerif soils on the banks of the River Nile in Khartoum, the capital city of Sudan. To achieve this objective, a preliminary baseline survey was carried out to describe the structure of the existing garden systems. In cooperation with the author of another PhD thesis (Ms. Ishtiag Abdalla), alternative uses of cow dung in brick making kilns in urban Khartoum were assessed; and the socio-economic criteria of the brick kiln owners or agents, economical and plant nutritional value of animal dung and the gaseous emission related to brick making activities were assessed. A total of 40 household heads were interviewed using a semi-structured questionnaire to collect information on demographic, socio-economic and migratory characteristics of the household members, the gardening systems used and the problems encountered in urban gardening. Based on the results of this survey, gardens were divided into three groups: mixed vegetable-fodder gardens, mixed vegetable-subsistence livestock gardens and pure vegetable gardens. The results revealed that UA is the exclusive domain of men, 80% of them non-native to Khartoum. The harvested produce in all gardens was market oriented and represented the main source of income for 83% of the gardeners. Fast growing leafy vegetables such as Jew’s mallow (Corchorous olitorius L.), purslane (Portulaca oleracea L.) and rocket (Eruca sativa Mill.) were the dominant cultivated species. Most of the gardens (95%) were continuously cultivated throughout the year without any fallow period, unless they were flooded. Gardeners were not generally aware of the importance of crop diversity, which may help them overcome the strongly fluctuating market prices for their produce and thereby strengthen the contributions of UA to the overall productivity of the city. To measure nutrient fluxes, four gardens were selected and their nutrients inputs and outputs flows were monitored. In each garden, all plots were monitored for quantification of nutrient inputs and outputs. To determine soil chemical fertility parameters in each of the studied gardens, soil samples were taken from three selected plots at the beginning of the study in October 2007 (gardens L1, L2 and H1) and in April 2008 (garden H2) and at the end of the study period in March 2010. Additional soil sampling occurred in May 2009 to assess changes in the soil nutrient status after the River Nile flood of 2008 had receded. Samples of rain and irrigation water (river and well-water) were analyzed for nitrogen (N), phosphorus (P), potassium (K) and carbon (C) content to determine their nutrient inputs. Catchment traps were installed to quantify the sediment yield from the River Nile flood. To quantify the nutrient inputs of sediments, samples were analyzed for N, P, K and organic carbon (Corg) content, cation exchange capacity (CEC) and the particle size distribution. The total nutrient inputs were calculated by multiplying the sediment nutrient content by total sediment deposits on individual gardens. Nutrient output in the form of harvested yield was quantified at harvest of each crop. Plant samples from each field were dried, and analyzed for their N, P, K and Corg content. Cumulative leaching losses of mineral N and P were estimated in a single plot in garden L1 from December 1st 2008 to July 1st 2009 using 12 ion exchange resins cartridges. Nutrients were extracted and analyzed for nitrate (NO3--N), ammonium (NH4+-N) and phosphate PO4-3-P. Changes in soil nutrient balance were assessed as inputs minus outputs. The results showed that across gardens, soil N and P concentrations increased from 2007 to 2009, while particle size distribution remained unchanged. Sediment loads and their respective contents of N, P and Corg decreased significantly (P < 0.05) from the gardens of the downstream lowlands (L1 and L2) to the gardens of the upstream highlands (H1 and H2). No significant difference was found in K deposits. None of the gardens received organic fertilizers and the only mineral fertilizer applied was urea (46-0-0). This equaled 29, 30, 54, and 67% of total N inputs to gardens L1, L2, H1, and H2, respectively. Sediment deposits of the River Nile floods contributed on average 67, 94, 6 and 42% to the total N, P, K and C inputs in lowland gardens and 33, 86, 4 and 37% of total N, P, K and C inputs in highland gardens. Irrigation water and rainfall contributed substantially to K inputs representing 96, 92, 94 and 96% of total K influxes in garden L1, L2, H1 and H2, respectively. Following the same order, total annual DM yields in the gardens were 26, 18, 16 and 1.8 t ha-1. Annual leaching losses were estimated to be 0.02 kg NH4+-N ha-1 (SE = 0.004), 0.03 kg NO3--N ha-1 (SE = 0.002) and 0.005 kg PO4-3-P ha-1 (SE = 0.0007). Differences between nutrient inputs and outputs indicated negative nutrient balances for P and K and positive balances of N and C for all gardens. The negative balances in P and K call for adoptions of new agricultural techniques such as regular manure additions or mulching which may enhance the soil organic matter status. A quantification of fluxes not measured in our study such as N2-fixation, dry deposition and gaseous emissions of C and N would be necessary to comprehensively assess the sustainability of these intensive gardening systems. The second part of the survey dealt with the brick making kilns. A total of 50 brick kiln owners/or agents were interviewed from July to August 2009, using a semi-structured questionnaire. The data collected included general information such as age, family size, education, land ownership, number of kilns managed and/or owned, number of months that kilns were in operation, quantity of inputs (cow dung and fuel wood) used, prices of inputs and products across the production season. Information related to the share value of the land on which the kilns were built and annual income for urban farmers and annual returns from dung for the animal raisers was also collected. Using descriptive statistics, budget calculation and Gini coefficient, the results indicated that renting the land to brick making kilns yields a 5-fold higher return than the rent for agriculture. Gini coefficient showed that the kiln owners had a more equal income distribution compared to farmers. To estimate emission of greenhouse gases (GHGs) and losses of N, P, K, Corg and DM from cow dung when used in brick making, samples of cow dung (loose and compacted) were collected from different kilns and analyzed for their N, P, K and Corg content. The procedure modified by the Intergovernmental Panel on Climate Change (IPCC, 1994) was used to estimate the gaseous emissions of cow dung and fuel wood. The amount of deforested wood was estimated according to the default values for wood density given by Dixon et al. (1991) and the expansion ratio for branches and small trees given by Brown et al. (1989). The data showed the monetary value of added N and P from cow dung was lower than for mineral fertilizers. Annual consumption of compacted dung (381 t DM) as biomass fuel by far exceeded the consumption of fuel wood (36 t DM). Gaseous emissions from cow dung and fuel wood were dominated by CO2, CO and CH4. Considering that Gerif land in urban Khartoum supports a multifunctional land use system, efficient use of natural resources (forest, dung, land and water) will enhance the sustainability of the UA and brick making activities. Adoption of new kilns with higher energy efficiency will reduce the amount of biomass fuels (cow dung and wood) used the amount of GHGs emitted and the threat to the few remaining forests.

Systematics and evolution of the genus Deuterocohnia Mez (Bromeliaceae)

The present study investigates the systematics and evolution of the Neotropical genus Deuterocohnia Mez (Bromeliaceae). It provides a comprehensive taxonomic revision as well as phylogenetic analyses based on chloroplast and nuclear DNA sequences and presents a hypothesis on the evolution of the genus. A broad morphological, anatomical, biogeographical and ecological overview of the genus is given in the first part of the study. For morphological character assessment more than 700 herbarium specimens from 39 herbaria as well as living plant material in the field and in the living collections of botanical gardens were carefully examined. The arid habitats, in which the species of Deuterocohnia grow, are reflected by the morphological and anatomical characters of the species. Important characters for species delimitation were identified, like the length of the inflorescence, the branching order, the density of flowers on partial inflorescences, the relation of the length of the primary bracts to that of the partial inflorescence, the sizes of floral bracts, sepals and petals, flower colour, the presence or absence of a pedicel, the curvature of the stamina and the petals during anthesis. After scrutinizing the nomenclatural history of the taxa belonging to Deuterocohnia – including the 1992 syonymized genus Abromeitiella – 17 species, 4 subspecies and 4 varieties are accepted in the present revision. Taxonomic changes were made in the following cases: (I) New combinations: A. abstrusa (A. Cast.) N. Schütz is re-established – as defined by Castellanos (1931) – and transfered to D. abstrusa; D. brevifolia (Griseb.) M.A. Spencer & L.B. Sm. includes accessions of the former D. lorentziana (Mez) M.A. Spencer & L.B. Sm., which are not assigned to D. abstrusa; D. bracteosa W. Till is synonymized to D. strobilifera Mez; D. meziana Kuntze ex Mez var. carmineo-viridiflora Rauh is classified as a subspecies of D. meziana (ssp. carmineo-viridiflora (Rauh) N. Schütz); D. pedicellata W. Till is classified as a subspecies of D. meziana (ssp. pedicellata (W. Till) N. Schütz); D. scapigera (Rauh & L. Hrom.) M.A. Spencer & L.B. Sm ssp. sanctae-crucis R. Vásquez & Ibisch is classified as a species (D. sanctae-crucis (R. Vásquez & Ibisch) N. Schütz); (II) New taxa: a new subspecies of D. meziana Kuntze ex Mez is established; a new variety of D. scapigera is established; (the new taxa will be validly published elsewhere); (III) New type: an epitype for D. longipetala was chosen. All other species were kept according to Spencer and Smith (1992) or – in the case of more recently described species – according to the protologue. Beside the nomenclatural notes and the detailed descriptions, information on distribution, habitat and ecology, etymology and taxonomic delimitation is provided for the genus and for each of its species. An key was constructed for the identification of currently accepted species, subspecies and varieties. The key is based on easily detectable morphological characters. The former synonymization of the genus Abromeitiella into Deuterocohnia (Spencer and Smith 1992) is re-evalutated in the present study. Morphological as well as molecular investigations revealed Deuterocohnia incl. Abromeitiella as being monophyletic, with some indications that a monophyletic Abromeitiella lineage arose from within Deuterocohnia. Thus the union of both genera is confirmed. The second part of the present thesis describes and discusses the molecular phylogenies and networks. Molecular analyses of three chloroplast intergenic spacers (rpl32-trnL, rps16-trnK, trnS-ycf3) were conducted with a sample set of 119 taxa. This set included 103 Deuterocohnia accessions from all 17 described species of the genus and 16 outgroup taxa from the remainder of Pitcairnioideae s.str. (Dyckia (8 sp.), Encholirium (2 sp.), Fosterella (4 sp.) and Pitcairnia (2 sp.)). With its high sampling density, the present investigation by far represents the most comprehensive molecular study of Deuterocohnia up till now. All data sets were analyzed separately as well as in combination, and various optimality criteria for phylogenetic tree construction were applied (Maximum Parsimony, Maximum Likelihood, Bayesian inferences and the distance method Neighbour Joining). Congruent topologies were generally obtained with different algorithms and optimality criteria, but individual clades received different degrees of statistical support in some analyses. The rps16-trnK locus was the most informative among the three spacer regions examined. The results of the chloroplast DNA analyses revealed a highly supported paraphyly of Deuterocohnia. Thus, the cpDNA trees divide the genus into two subclades (A and B), of which Deuterocohnia subclade B is sister to the included Dyckia and Encholirium accessions, and both together are sister to Deuterocohnia subclade A. To further examine the relationship between Deuterocohnia and Dyckia/Encholirium at the generic level, two nuclear low copy markers (PRK exon2-5 and PHYC exon1) were analysed with a reduced taxon set. This set included 22 Deuterocohnia accessions (including members of both cpDNA subclades), 2 Dyckia, 2 Encholirium and 2 Fosterella species. Phylogenetic trees were constructed as described above, and for comparison the same reduced taxon set was also analysed at the three cpDNA data loci. In contrast to the cpDNA results, the nuclear DNA data strongly supported the monophyly of Deuterocohnia, which takes a sister position to a clade of Dyckia and Encholirium samples. As morphology as well as nuclear DNA data generated in the present study and in a former AFLP analysis (Horres 2003) all corroborate the monophyly of Deuterocohnia, the apparent paraphyly displayed in cpDNA analyses is interpreted to be the consequence of a chloroplast capture event. This involves the introgression of the chloroplast genome from the common ancestor of the Dyckia/ Encholirium lineage into the ancestor of Deuterocohnia subclade B species. The chloroplast haplotypes are not species-specific in Deuterocohnia. Thus, one haplotype was sometimes shared by several species, where the same species may harbour different haplotypes. The arrangement of haplotypes followed geographical patterns rather than taxonomic boundaries, which may indicate some residual gene flow among populations from different Deuteroccohnia species. Phenotypic species coherence on the background of ongoing gene flow may then be maintained by sets of co-adapted alleles, as was suggested by the porous genome concept (Wu 2001, Palma-Silva et al. 2011). The results of the present study suggest the following scenario for the evolution of Deuterocohnia and its species. Deuterocohnia longipetala may be envisaged as a representative of the ancestral state within the genus. This is supported by (1) the wide distribution of this species; (2) the overlap in distribution area with species of Dyckia; (3) the laxly flowered inflorescences, which are also typical for Dyckia; (4) the yellow petals with a greenish tip, present in most other Deuterocohnia species. The following six extant lineages within Deuterocohnia might have independently been derived from this ancestral state with a few changes each: (I) D. meziana, D. brevispicata and D. seramisiana (Bolivia, lowland to montane areas, mostly reddish-greenish coloured, very laxly to very densely flowered); (II) D. strobilifera (Bolivia, high Andean mountains, yellow flowers, densely flowered); (III) D. glandulosa (Bolivia, montane areas, yellow-greenish flowers, densely flowered); (IV) D. haumanii, D. schreiteri, D. digitata, and D. chrysantha (Argentina, Chile, E Andean mountains and Atacama desert, yellow-greenish flowers, densely flowered); (V) D. recurvipetala (Argentina, foothills of the Andes, recurved yellow flowers, laxly flowered); (VI) D. gableana, D. scapigera, D. sanctae-crucis, D. abstrusa, D. brevifolia, D. lotteae (former Abromeitiella species, Bolivia, Argentina, higher Andean mountains, greenish-yellow flowers, inflorescence usually simple). Originating from the lower montane Andean regions, at least four lineages of the genus (I, II, IV, VI) adapted in part to higher altitudes by developing densely flowered partial inflorescences, shorter flowers and – in at least three lineages (II, IV, VI) – smaller rosettes, whereas species spreading into the lowlands (I, V) developed larger plants, laxly flowered, amply branched inflorescences and in part larger flowers (I).

Major-elements trends in cenozoic volcanites of Hungary

Hungary lies entirely within the Carpatho-Pannonian Region (CPR), a dominant tectonic unit of eastern Central Europe. The CPR consists of the Pannonian Basin system, and the arc of the Carpathian Mountains surrounding the lowlands in the north, east, and southeast. In the west, the CPR is bounded by the Eastern Alps, whereas in the south, by the Dinaridic belt. (...)

Modeling the Stream Water Nitrate Dynamics in 60,000-km(2) European Catchment, the Garonne, Southwest France

The spatial and temporal dynamics in the stream water NO3-N concentrations in a major European river-system, the Garonne (62,700 km(2)), are described and related to variations in climate, land management, and effluent point-sources using multivariate statistics. Building on this, the Hydrologiska Byrans Vattenbalansavdelning (HBV) rainfall-runoff model and the Integrated Catchment Model of Nitrogen (INCA-N) are applied to simulate the observed flow and N dynamics. This is done to help us to understand which factors and processes control the flow and N dynamics in different climate zones and to assess the relative inputs from diffuse and point sources across the catchment. This is the first application of the linked HBV and INCA-N models to a major European river system commensurate with the largest basins to be managed tinder the Water Framework Directive. The simulations suggest that in the lowlands, seasonal patterns in the stream water NO3-N concentrations emerge and are dominated by diffuse agricultural inputs, with an estimated 75% of the river load in the lowlands derived from arable farming. The results confirm earlier European catchment studies. Namely, current semi-distrubuted catchment-scale dynamic models, which integrate variations in land cover, climate, and a simple representation of the terrestrial and in-stream N cycle, are able to simulate seasonal NO3-N patterns at large spatial (> 300 km(2)) and temporal (>= monthly) scales using available national datasets.

An eclectic morphostratigraphic model for the sedimentary response to Holocene sea-level rise in northwest Europe

The improved empirical understanding of silt facies in Holocene coastal sequences provided by such as diatom, foraminifera, ostracode and testate amoebae analysis, combined with insights from quantitative stratigraphic and hydraulic simulations, has led to an inclusive, integrated model for the palaeogeomorphology, stratigraphy, lithofacies and biofacies of northwest European Holocene coastal lowlands in relation to sea-level behaviour. The model covers two general circumstances and is empirically supported by a range of field studies in the Holocene deposits of a number of British estuaries, particularly, the Severn. Where deposition was continuous over periods of centuries to millennia, and sea level fluctuated about a rising trend, the succession consists of repeated cycles of silt and peat lithofacies and biofacies in which series of transgressive overlaps (submergence sequences) alternate with series of regressive overlaps (emergence sequences) in association with the waxing and waning of tidal creek networks. Environmental and sea-level change are closely coupled, and equilibrium and secular pattern is of the kind represented ideally by a closed limit cycle. In the second circumstance, characteristic of unstable wetland shores and generally affecting smaller areas, coastal erosion ensures that episodes of deposition in the high intertidal zone last no more than a few centuries. The typical response is a series of regressive overlaps (emergence sequence) in erosively based high mudflat and salt-marsh silts that record, commonly as annual banding, exceptionally high deposition rates and a state of strong disequilibrium. Environmental change, including creek development, and sea-level movement are uncoupled. Only if deposition proceeds for a sufficiently long period, so that marshes mature, are equilibrium and close coupling regained. (C) 2002 Elsevier Science B.V. All rights reserved.

Autocyclic erosion in tidal marshes

A common mode whereby destruction of coastal lowlands occurs is frontal erosion. The edge cliffing, nonetheless, is also an inherent aspect of salt marsh development in many northwest European tidal marshes. Quite a few geomorphologists in the earlier half of the past century recognized such edge erosion as a definite repetitive stage within an autocyclic mode of marsh growth. A shift in research priorities during the past decades (primarily because of coastal management concerns, however) has resulted in an enhanced focus on sediment-flux measurement campaigns on salt marshes. This, somewhat "object-oriented" strategy hindered any further development of the once-established autocyclic growth concept, which virtually has gone into oblivion in recent times. This work makes an attempt to resurrect the notion of autocyclicity by employing its premises to address edge erosion in tidal marshes. Through a review of intertidal morphosedimentology the underlying framework for autocyclicity is envisaged. The phenomenon is demonstrated in the Holocene salt marsh plain of Moricambe basin in NW England that displays several distinct phases of marsh retreat in the form of abandoned clifflets. The suite of abandoned shorelines and terraces has been identified in detailed field mapping that followed analysis of topographic maps and aerial photographs. Vertical trends in marsh plain sediments are recorded in trenches for signs of past marsh front movements. The characteristic sea level history of the area offers an opportunity to differentiate the morphodynamic variability induced in the autocyclic growth of the marsh plain in scenarios of rising and falling sea level and the accompanied change in sediment budget. The ideas gathered are incorporated to construct a conceptual model that links temporal extent of marsh erosion to inner tidal flat sediment budget and sea level tendency. The review leads to recognition of the necessity of adopting an holistic approach in the morphodynamic investigations where marshes should be treated as a component within the "marsh-mudflat system" as each element apparently modulates evolution of the other, with an eventual linkage to subtidal channels. (C) 2009 Elsevier B.V. All rights reserved.

Modeling the Stream Water Nitrate Dynamics in 60,000-km(2) European Catchment, the Garonne, Southwest France

The spatial and temporal dynamics in the stream water NO3-N concentrations in a major European river-system, the Garonne (62,700 km(2)), are described and related to variations in climate, land management, and effluent point-sources using multivariate statistics. Building on this, the Hydrologiska Byrans Vattenbalansavdelning (HBV) rainfall-runoff model and the Integrated Catchment Model of Nitrogen (INCA-N) are applied to simulate the observed flow and N dynamics. This is done to help us to understand which factors and processes control the flow and N dynamics in different climate zones and to assess the relative inputs from diffuse and point sources across the catchment. This is the first application of the linked HBV and INCA-N models to a major European river system commensurate with the largest basins to be managed tinder the Water Framework Directive. The simulations suggest that in the lowlands, seasonal patterns in the stream water NO3-N concentrations emerge and are dominated by diffuse agricultural inputs, with an estimated 75% of the river load in the lowlands derived from arable farming. The results confirm earlier European catchment studies. Namely, current semi-distrubuted catchment-scale dynamic models, which integrate variations in land cover, climate, and a simple representation of the terrestrial and in-stream N cycle, are able to simulate seasonal NO3-N patterns at large spatial (> 300 km(2)) and temporal (>= monthly) scales using available national datasets.

Bergmann's rule and body size in mammals

Aim: To describe the geographical pattern of mean body size of the non-volant mammals of the Nearctic and Neotropics and evaluate the influence of five environmental variables that are likely to affect body size gradients. Location: The Western Hemisphere. Methods: We calculated mean body size (average log mass) values in 110 × 110 km cells covering the continental Nearctic and Neotropics. We also generated cell averages for mean annual temperature, range in elevation, their interaction, actual evapotranspiration, and the global vegetation index and its coefficient of variation. Associations between mean body size and environmental variables were tested with simple correlations and ordinary least squares multiple regression, complemented with spatial autocorrelation analyses and split-line regression. We evaluated the relative support for each multiple-regression model using AIC. Results: Mean body size increases to the north in the Nearctic and is negatively correlated with temperature. In contrast, across the Neotropics mammals are largest in the tropical and subtropical lowlands and smaller in the Andes, generating a positive correlation with temperature. Finally, body size and temperature are nonlinearly related in both regions, and split-line linear regression found temperature thresholds marking clear shifts in these relationships (Nearctic 10.9 °C; Neotropics 12.6 °C). The increase in body sizes with decreasing temperature is strongest in the northern Nearctic, whereas a decrease in body size in mountains dominates the body size gradients in the warmer parts of both regions. Main conclusions: We confirm previous work finding strong broad-scale Bergmann trends in cold macroclimates but not in warmer areas. For the latter regions (i.e. the southern Nearctic and the Neotropics), our analyses also suggest that both local and broad-scale patterns of mammal body size variation are influenced in part by the strong mesoscale climatic gradients existing in mountainous areas. A likely explanation is that reduced habitat sizes in mountains limit the presence of larger-sized mammals.

Reconstruction of historical pollination rates reveals linked declines of pollinators and plants

Widespread reports of low pollination rates suggest a recent anthropogenic decline in pollination that could threaten natural and agricultural ecosystems. Nevertheless, unequivocal evidence for a decline in pollination over time has remained elusive because it was not possible to determine historical pollination rates. Here we demonstrate a widely applicable method for reconstructing historical pollination rates, thus allowing comparison with contemporary rates from the same sites. We focused on the relationship between the oil-collecting bee Rediviva peringueyi (Melittidae) and the guild of oil-secreting orchid species (Coryciinae) that depends on it for pollination. The guild is distributed across the highly transformed and fragmented lowlands of the Cape Region of South Africa. We show that rehydrated herbarium specimens of Pterygodium catholicum, the most abundant member of the guild, contain a record of past pollinator activity in the form of pollinarium removal rates. Analysis of a pollination time series showed a recent decline in pollination on Signal Hill, a small urban conservation area. The same herbaria contain historical species occurrence data. We analyzed this data and found that there has been a contemporaneous shift in orchid guild composition in urban areas due to the local extirpation of the non-clonal species, consistent with their greater dependence on seeds and pollination for population persistence.

The implications of the 2003 Common Agricultural Policy reforms for land use and landscape quality in England

The assessment of the potential landscape impacts of the latest Common Agricultural Policy reforms constitutes a challenge for policy makers and it requires the development of models that can reliably project the likely spatial distribution of land uses. The aim of this study is to investigate the impact of 2003 CAP reforms to land uses and rural landscapes across England. For this purpose we modified an existing economic model of agriculture, the Land-Use Allocation Model (LUAM) to provide outputs at a scale appropriate for informing a semi-quantitative landscape assessment at the level of ‘Joint Character Areas’ (JCAs). Overall a decline in the cereal and oilseed production area is projected but intensive arable production will persist in specific locations (East of England, East Midlands and South East), having ongoing negative effects on the character of many JCAs. The impacts of de-coupling will be far more profound on the livestock sector; extensification of production will occur in traditional mixed farming regions (e.g. the South West), a partial displacement of cattle by sheep in the upland regions and an increase in the sheep numbers is expected in the lowlands (South East, Eastern and East Midlands). This extensification process will affect positively those JCAs of mixed farming conditions, but it will have negative impacts on the JCAs of historically low intensity farming (e.g. the uplands of north-west) because they will suffer from under-management and land idling. Our analysis shows that the territorialisation between intensively and extensively agricultural landscapes will continue.

The determination of total nitrogen and total phosphorus concentrations in freshwaters from land use, stock headage and population data: testing of a model for use in conservation and water quality management

1. Nutrient concentrations (particularly N and P) determine the extent to which water bodies are or may become eutrophic. Direct determination of nutrient content on a wide scale is labour intensive but the main sources of N and P are well known. This paper describes and tests an export coefficient model for prediction of total N and total P from: (i) land use, stock headage and human population; (ii) the export rates of N and P from these sources; and (iii) the river discharge. Such a model might be used to forecast the effects of changes in land use in the future and to hindcast past water quality to establish comparative or baseline states for the monitoring of change. 2. The model has been calibrated against observed data for 1988 and validated against sets of observed data for a sequence of earlier years in ten British catchments varying from uplands through rolling, fertile lowlands to the flat topography of East Anglia. 3. The model predicted total N and total P concentrations with high precision (95% of the variance in observed data explained). It has been used in two forms: the first on a specific catchment basis; the second for a larger natural region which contains the catchment with the assumption that all catchments within that region will be similar. Both models gave similar results with little loss of precision in the latter case. This implies that it will be possible to describe the overall pattern of nutrient export in the UK with only a fraction of the effort needed to carry out the calculations for each individual water body. 4. Comparison between land use, stock headage, population numbers and nutrient export for the ten catchments in the pre-war year of 1931, and for 1970 and 1988 show that there has been a substantial loss of rough grazing to fertilized temporary and permanent grasslands, an increase in the hectarage devoted to arable, consistent increases in the stocking of cattle and sheep and a marked movement of humans to these rural catchments. 5. All of these trends have increased the flows of nutrients with more than a doubling of both total N and total P loads during the period. On average in these rural catchments, stock wastes have been the greatest contributors to both N and P exports, with cultivation the next most important source of N and people of P. Ratios of N to P were high in 1931 and remain little changed so that, in these catchments, phosphorus continues to be the nutrient most likely to control algal crops in standing waters supplied by the rivers studied.

Diversity of cultivars and other plant resources used at habitation sites in the Llanos de Mojos, Beni, Bolivia: Evidence from macrobotanical remains, starch grains, and phytoliths

Although sparsely populated today, the Llanos de Mojos, Bolivia, sustained large sedentary societies in the Late Holocene (ca. 500 to 1400 AD). In order to gain insight into the subsistence of these people, we undertook macrobotanical and phytolith analyses of sediment samples, and starch grain and phytolith analyses of artifact residues, from four large habitation sites within this region. Macrobotanical remains show the presence of maize (Zea mays), squash (Cucurbita sp.), peanut (Arachis hypogaea), cotton (Gossypium sp.), and palm fruits (Arecaceae). Microbotanical results confirm the widespread use of maize at all sites, along with manioc (Manihot esculenta), squash, and yam (Dioscorea sp.). These integrated results present the first comprehensive archaeobotanical evidence of the diversity of plants cultivated, processed, and consumed, by the pre-Hispanic inhabitants of the Amazonian lowlands of Bolivia.

Pollen-based differentiation of Amazonian rainforest communities and implications for lowland palaeoecology in tropical South America

An ongoing controversy in Amazonian palaeoecology is the manner in which Amazonian rainforest communities have responded to environmental change over the last glacial–interglacial cycle. Much of this controversy results from an inability to identify the floristic heterogeneity exhibited by rainforest communities within fossil pollen records. We apply multivariate (Principal Components Analysis) and classification (Unweighted Pair Group with Arithmetic Mean Agglomerative Classification) techniques to floral-biometric, modern pollen trap and lake sediment pollen data situated within different rainforest communities in the tropical lowlands of Amazonian Bolivia. Modern pollen rain analyses from artificial pollen traps show that evergreen terra firme (well-drained), evergreen terra firme liana, evergreen seasonally inundated, and evergreen riparian rainforests may be readily differentiated, floristically and palynologically. Analogue matching techniques, based on Euclidean distance measures, are employed to compare these pollen signatures with surface sediment pollen assemblages from five lakes: Laguna Bella Vista, Laguna Chaplin, and Laguna Huachi situated within the Madeira-Tapajós moist forest ecoregion, and Laguna Isirere and Laguna Loma Suarez, which are situated within forest patches in the Beni savanna ecoregion. The same numerical techniques are used to compare rainforest pollen trap signatures with the fossil pollen record of Laguna Chaplin.