Dissertationen zum Thema „Bryophytic species“

Nutrient cycling and energy flow within various ecosystems has been the subject of many studies in the past. The influence bryophyte litter has on nutrient cycling has not been investigated extensively and research on nitrogen flow, in particular, is limited. In the following study the litter quality and decomposition characteristics of material from six New Zealand moss species was investigated using both laboratory and field studies. Ptychomnion aciculare, Hypnodendron comatum and Dicranoloma dicarpum were collected from Maruia Springs in the Lewis Pass Reserve and Hypnum cupressiforme, Breutelia pendula and Racomitrium pruinosum came from Cass near Arthur's Pass. Differences between the six species and between the green (younger) and brown (older) fractions of the litter were investigated with respect to litter quality, decay rates and nitrogen mobilisation. Due to the lack of an obvious senescence cycle naturally dead moss litter was difficult to determine therefore comparisons between the older and younger material were made. Moss litter is thought to be resistant to microbial degradation due to the chemical characteristics of the material Chemical analysis of the species showed a range of high C:N and C:P ratios (53-188 and 206-815 respectively), and that approximately 90% of the acid soluble nitrogen was derived from amino acids and proteinaceous compounds. The high percentage of amino acid and proteinaceous nitrogen indicates that the litter may be of high quality if the nitrogen is not complexed with other compounds causing it to become recalcitrant. Decay rates (k) were determined from weight loss experiments giving a range of 0.23- 1.3. There was an observed increase in weight loss over time for all the litter types. Nitrogen mineralisation studies indicated that nitrogen was released (generally in the form of ammonium ions) by microbial activity over 180 days incubation. Nitrate was also observed which in some cases exceeded ammonium levels. Leaching of the litter yielded water-soluble compounds including high nitrogen levels. Water-soluble extracts were shown to support microbial growth.

Climate change is expected to have a strong impact on subarctic ecosystems. Increased temperatures as well as altered precipitation and snow cover patterns are predicted to change species distribution and affect biogeochemical processes in the subarctic tundra. Bryophytes are an essential vegetational component in northern ecosystems, due to their high abundance and importance in many ecological processes. In this study the effects of elevation and altered snow cover on the temporal dynamics of freezing damage in three subarctic bryophyte species (Hylocomium splendens, Ptilidium ciliare, and Sphagnum fuscum) were studied in a snow manipulation field experiment in Abisko, during early spring. Soil temperature and field moisture of moss shoots were collected. A freeze-thaw incubation experiment was conducted to investigate the freeze-thaw cycle resistance of H. splendens and P. ciliare originating from habitats with two differing snow-cover thicknesses. Freezing damage differed significantly between the bryophytes species with P. ciliare experiencing the least and S. fuscum the highest damage. Damage was higher at the low elevation, possibly attributable to acclimation effects. Snow removal led to higher damage in moss shoots, but no interactions of the different snow cover treatments with elevation, species or time were found. Freezing damage increased over time and no recovery occurred, likely due to temporal patterns in soil freeze-thaw cycles during early spring. Soil freeze-thaw cycles were the main factor influencing damage in bryophytes after snow melt. Measured environmental parameters could not explain the entire variation in damage. Damage might additionally be attributable to increased UV radiation or disturbances by herbivores.

A tool for management and conservation of valuable forests in Sweden are WKH:s. In this study WKH:s different in size, connectivity, amount of dead wood and quality of logs were investigated for species richness of bryophytes and brackets on coniferous logs. The aim was to clarify which scales and features that was important for the diversity of species as well as for individual species. The results showed that the amount of dead wood was most important on the site scale, and some species were affected at the landscape scale, a positive effect of valuable tracts. On the scale of individual logs, factors such as diameter, sun exposure, succession stage, contact with ground and ground bryophytes cover was most important. Red-listed species preferred logs with large diameter and late successional stages. The conclusion was that the quality of the substrate and the amount of dead wood was most important, but the amount of WKH:s on the landscape scale was also important for some species.

This study focused on the primary cell wall constituents and plasmodesmata (PD) density in three mosses and four liverwort apical cells (AC) and immediate derivatives. The three mosses have tetrahedral apical cells and the liverworts possess tetrahedral, hemidiscoid and lenticular AC geometries. The primary cell wall in the studied taxa is comprised of two layers. A fibrillar layer, which is the outer wall layer, contains compacted cellulosic fibrils, and represents the two adjacent primary walls and middle lamella, the latter of which is rarely discernible. An electron-lucent inner wall layer abuts the plasma membrane. This layer has faint fibrous materials that extend from the plasma membrane to the fibrillar layer. Generally, as the cell wall ages it thickens, the fibrillar layer increases in width while the electron-lucent wall stays more or less consistent in width. In the four liverworts, the most recent wall of the AC has the highest PD density in the apical region regardless of AC geometry. As the walls elongate, primary wall is laid down between PD, separating them and resulting in lower densities and wider PD diameters in older walls. The season of fixation and whether plants were studied from nature or culture have an influence on AC ultrastructure. A developmental study of Physcomitrella patents gametophores in four stages, bud, 2-leaved, 7-8-leaved and ~20-leaved, reveals that the primary cell wall constituents change slightly during development. Specifically, LM5 a RG-I pectin antibody against the galactan branch epitope is only localized in the fibrillar layer of young water-conducting cells in the 7-8-leaved and 20-leaved gametophores. LM20, an antibody against HG esterified pectins, does not localize in any of the cell walls during development. The distribution patterns for AGPs (JIM13 and LM2) are consistent during gametophore development and predominantly localize on the electron-lucent layer and wall/plasma membrane interface. However, LM2 is mainly localized on the fibrillar layer in 7-8-leaved cell walls. AGPs also localize on element of the cytoplasm. LM6, an antibody against an RG-I pectin with arabinan branch epitopes, also localizes AGPs and because it expressed similar distribution patterns as JIM13 and LM2 on the cell wall, it likely localizes AGP in Physcomitrella. In addition, LM6 localizes pectins on the fibrillar layer similar to LM5 and LM19 for HG unesterified pectins. Callose predominantly localizes at the PD neck region. This study provides the first documentation of changes in size and shape of AC with age in Physcomitrella patens gametophores. The PD densities of gametophytes examined in this study fall into the lineage-specific network of PD (LPD) group designated for sporophytes of monilophytes and Selaginella (heterosporous lycophyte) with single ACs. Takakia lepidozioides leafy shoot has a tetrahedral AC with a highly curved free surface. This peculiar moss has mucilage hair (MH) associated with axil of phyllids. Mucilage hair in both species are 3-celled with a forth epidermal cell as the base. However, occasional 2-celled MH is seen in T. ceratophylla. The ultrastructure of MH has similarities with other mosses and liverworts.

The effect of moderate rates of nitrogen deposition on ground floor vegetation is poorly predicted by uncontrolled surveys or fertilization experiments using high rates of nitrogen (N) addition. We compared the temporal trends of ground floor vegetation in permanent plots with moderate (7–13 kg ha−1 year−1) and lower bulk N deposition (4–6 kg ha−1 year−1) in southern Sweden during 1982–1998. We examined whether trends differed between growth forms (vascular plants and bryophytes) and vegetation types (three types of coniferous forest, deciduous forest, and bog). Trends of site-standardized cover and richness varied among growth forms, vegetation types, and deposition regions. Cover in spruce forests decreased at the same rate with both moderate and low deposition. In pine forests cover decreased faster with moderate deposition and in bogs cover decreased faster with low deposition. Cover of bryophytes in spruce forests increased at the same rate with both moderate and low deposition. In pine forests cover decreased faster with moderate deposition and in bogs and deciduous forests there was a strong non-linear increase with moderate deposition. The trend of number of vascular plants was constant with moderate and decreased with low deposition. We found no trend in the number of bryophyte species. We propose that the decrease of cover and number with low deposition was related to normal ecosystem development (increased shading), suggesting that N deposition maintained or increased the competitiveness of some species in the moderate-deposition region. Deposition had no consistent negative effect on vegetation suggesting that it is less important than normal successional processes.

Bryophytes are rarely used as a food source by any animal species, but the genus Tipula (Diptera, Tipulidae) contains some of the few insect species able to feed, and complete their life-cycle, on bryophytes. Vegetation particle volumes ingested by larvae of eleven Tipula species increased only marginally between instars and not to the extent expected from the rate of growth of body mass. Early and late instars within a species frequently ingested similar sized particles. The overall efficiency of digestion of vegetation particles was low and similar between the four instars of each of the eleven species. Generally, the only method by which later larval instars can obtain a higher proportion of nutrients is by feeding on a larger number of smaller vegetation particles and not by ingesting large particles. In feeding choice experiments, Tipula confusa preferred moss species from woodland habitats, whereas Tipula subnodicornis did not show an overall preference for either woodland or moorland moss species. Tipula subnodicornis also showed a less extensive hierachical preference/avoidance than Tipula confusa for the ten moss species investigated. The moss species Campylopus paradoxus and Sphagnum papillosum accumulated Pb(^2+) ions and Zn(^2+) ions to high concentrations. There was some evidence that Tipula subnodicornis larvae were deterred from feeding on these mosses with high levels of introduced heavy metal ions. Tipula montana was able to thrive and complete its life-cycle in Britain at lower altitudes than had been previously thought. Individuals of this species show a combination of one-year and two-year life-cycles at Waskerley Common. The feeding methods employed by Tipula species can explain why some of them have remained as consumers of bryophytes.

The aim of this thesis was to increase knowledge about metapopulation and metacommunity processes in patchy, dynamic landscapes, using epiphytic bryophytes as a model system. Host trees and deciduous forest stands in the coniferous landscape are patchy, temporal and undergo changes in habitat quality during succession. Epiphytes must track this dynamic habitat network for their long-term survival. Community patterns at different spatial scales were explored and linked to regional metapopulation processes and local population dynamics. Spatial structuring in species richness both at a local and regional scale indicated stronger dispersal limitation but lower sensitivity to habitat quality in species with large asexual than in species with small sexual diaspores. In sexually dispersed species, a strong rescue effect was indicated by a bimodal frequency distribution of the species and by increasing local abundance with increasing patch connectivity. Present connectivity to other deciduous forest patches had positive effects on richness of asexually dispersed species, whereas richness of sexually dispersed species was instead related to the landscape connectivity 30 years ago. A study of local growth and reproduction suggested that this is caused by delayed sexual, but not asexual, reproduction. Habitat conditions affected the production of sporophytes, but not of asexual diaspores. No differences in either growth rates or competitive abilities among species with different dispersal and life-history strategies were found. In vitro experiments showed that establishment is higher from large asexual diaspores than from small sexual. Establishment of all diaspore types was limited by pH. There were indications of trade-offs between high germination and protonemal growth rates, desiccation tolerance and a rapid development of shoots from protonema. The results indicated that the epiphyte metacommunity is structured by two main trade-offs: dispersal distance (diaspore size) versus age at first reproduction, and dispersal distance versus sensitivity to habitat quality. Trade-offs in species traits may have evolved as a consequence of conflicting selection pressures imposed by habitat turnover, connectivity and irregular water supply rather than by species interactions. Syndromes of interrelated species traits imply that fairly small changes in habitat conditions can lead to distinct changes in metacommunity diversity: the results indicate that increasing distances among patches cause most harm to asexually dispersed species, whereas cuttings of forests of high age and quality as well as increasing patch dynamics are most harmful to sexually dispersed species.
Senarelagd disputation från: 2009-09-26, Lindahlsalen, EBC, Villavägen 9, 75236 Uppsala, Uppsala, 10:00

Abstract Understanding of natural and human induced changes to freshwater biodiversity patterns is critical for efficiently conserving and managing these highly impacted ecosystems. In this doctoral thesis, I study the natural spatial and temporal variability of littoral macroinvertebrate communities, as well as aquatic biodiversity responses (of bryophytes, macroinvertebrates and diatoms) to human impact on forested headwater streams. I also study the temporal variability of natural and human-impacted stream communities. I show that littoral macroinvertebrate communities are most variable between individual samples and to some extent between littorals, and that the diversity patterns of rare and common littoral macroinvertebrates vary at different spatial scales. Temporal (interannual) variability is of minor importance, particularly for the core species of the community. For rare species, however, the temporal component was generally more important than spatial turnover. For headwater streams, I show that most taxonomic groups responded negatively to human impact on the adjacent riparian forest. Nevertheless, pristine riparian forest is not a sufficient surrogate for stream biodiversity as also streams with relatively disturbed riparian forests supported diverse benthic communities. A rapid classification method based on the physical structure of the stream channel and its riparian forest is able to identify the reference-state communities, as well as the most severely altered communities, but a reliable identification of the intermediately-disturbed sites would need more precise information about the key stressors of stream communities. I found that fine sediments originating from forest drainage were the main factor causing reduction of stream biodiversity. I also found that forestry impacted streams varied temporally more than pristine streams, emphasizing the importance of interannual sampling of impacted streams for a reliable status assessment
Tiivistelmä Luonnollisen ja ihmisen aiheuttaman biodiversiteettivaihtelun ymmärtäminen hoito- ja suojelutoimissa on tärkeää pienvesien monimuotoisuuden turvaamiseksi. Tarkastelen väitöskirjassani järvien rantavyöhykkeen pohjaeläinten ajallista ja paikallista vaihtelua sekä latvapurojen eliöyhteisöjen (pohjaeläimet, sammalet, piilevät) vasteita ihmisen aiheuttamaan fysikaaliseen muutokseen rantametsässä ja purouomassa. Tarkastelen myös ajallisen vaihtelun eroja luonnontilaisten ja ihmisvaikutteisten latvapurojen eliöyhteisöjen välillä. Osoitan, että järven rantavyöhykkeen pohjaeläinyhteisöt vaihtelevat eniten yksittäisten näytteiden välillä ja pienessä määrin eri rantojen välillä, mutta ajallinen vaihtelu ei heikennä tulosten luotettavuutta muiden kuin harvinaisten lajien osalta yhteen vuoteen rajoittuvassa näytteenotossa. Purotutkimuksissa osoitan, että useiden vesieliöryhmien lajirunsaus muuttuu ihmisen aiheuttaman rantametsän muutoksen seurauksena, mutta luonnontilaista rantametsää ei silti voida pitää yksiselitteisesti myös puron suojelutason mittarina, koska se ei välttämättä turvaa eliöiden lajistollista eheyttä suhteessa alkuperäiseen alueelliseen lajistoon. Rantametsän ja uoman fysikaalisiin rakennepiirteisiin perustuvaa nopeaa luokittelumenetelmää voidaan käyttää eliöyhteisöltään monimuotoisimpien ja metsätalouden voimakkaimmin rasittamien purojen tunnistamisessa. Eliöyhteisöltään lievemmin muutettujen purojen luotettavaan erotteluun luokittelun pitäisi kuitenkin sisältää tarkempaa tietoa metsäojitusten aiheuttamasta liettymisestä, joka on tulosteni perusteella haitallisin metsätaloudesta johtuva puroyhteisöjä rasittava tekijä. Havaitsin myös metsätalouden rasittamien purojen tilanarvioinnin toistettavuuden olevan luonnontilaisia puroja heikompaa, joten metsätalouden rasittamien purojen pidempiaikainen seuraaminen tuntuu välttämättömältä luotettavien tilanarvioiden tuottamiseksi

Abstract Knowing which factors govern variation in community structure is crucial for assessing and conserving biodiversity. Two main processes structuring biological communities are selection by environmental conditions and dispersal between sites. In this thesis, I studied the potential importance of these two processes in structuring stream metacommunities (i.e. a set of local communities that are connected by the dispersal of individuals). I used altogether four datasets collected from three different drainage basins located in northern Finland. For small species, such as stream diatoms, bryophytes and macroinvertebrates, measuring dispersal directly and for all the species in a community, is very difficult. Thus, I used three kinds of spatial proxies to represent the potential effects of dispersal: (1) the size of species pool, (2) spatial locations of the sample sites and (3) distances between sites. In general, local environmental characteristics explained the variation in community structure better than any of the spatial proxies. The results suggested that stream diatoms, bryophytes and macroinvertebrates should not be dispersal limited at within-basin scale. However, in some cases spatial proxies explained a relatively larger proportion of community variation. One major difficulty when using such proxies is that high and limiting dispersal rates produce similar patterns. Here, I also present and test a novel hierarchical metacommunity approach for differentiating if a significant spatial signal is produced by high or limiting dispersal rates. This approach seemed promising. The results suggested that, at very small spatial scales (i.e. within a stream section) the community composition of stream macroinvertebrates is possibly influenced by high dispersal rates. I also used the so-called deconstructive approach and assigned the whole community into smaller groups of species based on their traits (i.e. feeding mode, dispersal mode and body size). This approach showed, for example, that metacommunity organization of species with actively dispersing terrestrial adults was, in general, more strongly related to environmental variables than the metacommunity organization of more passively dispersing species. I conclude that several approaches are needed for disentangling the importance of dispersal at the metacommunity level
Tiivistelmä Eliöyhteisöjen rakenteeseen vaikuttavien tekijöiden tunteminen on välttämätöntä luonnon monimuotoisuuden hoidon ja suojelun kannalta. Kaksi merkittävää tekijää yhteisörakenteen säätelyssä ovat ympäristötekijöiden aiheuttama valinta ja levittäytyminen. Tutkin väitöskirjassani näiden kahden tekijän suhteellista merkitystä puroeliöiden metayhteisöjen rakentumiselle. Metayhteisöllä tarkoitetaan joukkoa paikallisia yhteisöjä, joiden välillä yksilöt kykenevät levittäytymään. Käytin neljää aineistoa, jotka oli kerätty yhteensä kolmelta valuma-alueelta Pohjois-Suomesta. Pienille lajeille, kuten piileville, sammalille ja pohjaeläimille, levittäytymisen mittaaminen suoraan ja kaikille yhteisön lajeille vertailukelpoisella tavalla on hyvin vaikeaa. Tässä väitöskirjassa tavoitteeni oli selvittää, kuinka hyvin epäsuoria menetelmiä voidaan käyttää levittäytymisen merkityksen selvittämisessä. Käytin kolmea erilaista niin sanottua spatiaalista muuttujaa kuvaamaan levittäytymisen potentiaalisia vaikutuksia: (1) alueellisen lajipoolin kokoa, (2) tutkimuspaikkojen spatiaalista sijaintia ja (3) paikkojen välisiä etäisyyksiä. Yleisesti ottaen tulokset osoittivat, että paikalliset ympäristötekijät selittivät yhteisörakenteen vaihtelua paremmin kuin mikään spatiaalisista muuttujista. Saadut tulokset viittasivat siihen, että valuma-alueen sisällä purojen piilevien, sammalten ja pohjaeläinten yhteisörakenteen ei pitäisi olla rajoittuneen levittäytymisen muokkaamaa. Joissain tapauksissa jokin levittäytymistä kuvaava muuttuja kuitenkin selitti suhteellisesti suuremman osuuden yhteisörakenteen vaihtelusta. Tämä tulos tuo esille yhden spatiaalisten muuttujien käyttöön liittyvän merkittävän haasteen: samankaltaiseen tulokseen voi johtaa sekä voimakas että rajoittunut levittäytyminen. Väitöskirjassani esittelen ja testaan myös uudenlaista lähestymistapaa, jonka tavoitteena on erottaa, onko merkitsevä spatiaalinen signaali voimakkaan vai rajoittuneen levittäytymisen aiheuttamaa. Tämä uusi menetelmä viittasi siihen, että erittäin pienillä spatiaalisilla mittakaavoilla, kuten saman purojakson sisällä, puropohjaeläinten yhteisörakenne voi olla voimakkaan levittäytymisen muovaamaa. Käytin myös niin kutsuttua dekonstruktiivista lähestymistapaa. Jaoin koko yhteisön pienempiin ryhmiin niiden lajiominaisuuksien perusteella (esim. ravinnonotto, levittäytymistyyppi, vartalon koko). Saadut tulokset antoivat muun muassa viitteitä siitä, että aktiivisesti levittäytyvien eliöiden metayhteisöjen rakentuminen saattaa olla passiivisesti levittäytyviä lajeja riippuvaisempi paikallisista ympäristötekijöistä. Yhteenvetona totean, että metayhteisötasolla levittäytymisen merkityksen selvittämiseksi on käytettävä samanaikaisesti hyvin monenlaisia menetelmiä, sillä jokaisella menetelmällä on puutteensa ja siten eri menetelmiä tarvitaan täydentämään toisiaan

The aims of this thesis are to (1) analyze the predictability (indicators) of plant and fungal species diversity in old-growth forests, and (2) assess the history and biodiversity of woodland key habitats (WKHs) and their potential to maintain species diversity in fragmented boreal forest landscapes.

Predictability was explored in Granlandet nature reserve, an unexploited landscape composed of discrete old-growth Picea forest patches of varying size isolated by wetland, reflecting conditions of insular biota at stochastic equilibrium. Data from 46 patches (0.2-12 ha) showed that most species were rare. However, species richness and composition patterns exhibited a high degree of predictability, which strengthen the possibility to apply biodiversity indicators in old-growth forest stands. Area was a key factor. The increase in species richness starts to level out at 2-3 ha. Large patches host more Red-list species in their interiors than do small ones, i.e. stand size is an important qualitative aspect of old-growth habitat. Nestedness emerged in relation to area but also in equal-sized plots. Structural complexity and habitat quality were important for species richness and compositional patterns, and small habitats of high quality could harbor many rare species. Monitoring of wood-fungi on downed logs showed that species diversity on downed logs changed over periods of 5-10 years and that the occurrences of annual species were unpredictable. It is suggested that monitoring of species with durable fruit bodies (mainly polypores) is likely to be a feasible approach to obtain comparable data over time.

Assessments of biodiversity of WKHs were performed in two areas with contrasting histories of forest exploitation, namely in south boreal and north boreal Sweden. Analyses of the history of 15 south boreal WKHs showed that fire-suppression, selective logging until mid-20th century and abandonment by modern forestry has shaped their forest structure. These WKHs are not untouched forests, they lack key structural components and harbor few Red-list species. Artificial interventions to restore natural processes and patterns are needed to further increase their suitability for threatned species. Modeling analyses of species richness in 32 WKHs in north boreal Sweden, some of which have not been isolated by modern forestry until recently, indicated an excess of crustose lichen species, i.e. WKHs may face delayed species extinctions. By contrast, the results indicate that wood-fungi have tracked the environmental changes. Differences in substrate dynamics between epiphytes on living trees and species growing on decaying logs may explain the diffeence between species groups. The results indicate that population densities of Red-list species were low, which may result in further depletion of species diversity.

Continuing species declines and extinctions are likely if not conservation of WKHs are combined with other considerations in th managed forest landscape. Both WKHs and their surroundings must be managed and designed to maintain biodiversity over time. For a successful future conservation of boreal forest biodiversity monitoring of WKHs must be combined with monitoring of refeence areas.

Rain forest canopies are renowned for their very high biodiversity and the critical role they play in key ecological processes and their influence on global climate. Despite that New Zealand supports one of the most diverse and extensive epiphyte flora of any temperate forest system, few studies have investigated epiphyte communities and their invertebrate fauna along with factors that influence their distribution and composition. This thesis represents the first comprehensive study of entire epiphyte communities and their resident invertebrate fauna in the canopy of New Zealand’s indigenous forests. The aim of this study was to determine spatial patterns of epiphyte and invertebrate species richness, abundance and community composition in relation to abiotic variables, and in particular, the responses of these communities to elevated temperature and rainfall. This study was carried out in coastal lowland podocarp-broadleaved forests at two sites on the West Coast of the South Island of New Zealand. Samples from 120 mat-forming epiphyte assemblages located on inner canopy branches of 40 northern rata (Metrosideros robusta) trees were studied to characterise the component flora and fauna. Additionally, biomass, branch and tree characteristics and community responses to treatments designed to elevate temperature and rainfall to simulate predicted climate change were measured. This investigation revealed astonishing diversity and functional complexity of epiphyte and invertebrate life in this ecosystem. The 30.6 kg (dry weight) of epiphyte material collected contained a total of 567 species, 170 epiphyte and 397 invertebrate (excluding immature specimens and mites) species, including at least 10 species new to science and many undescribed species Epiphyte communities were found to be dominated by non-vascular plants (80 % of the total species richness), particularly liverworts and invertebrate communities were dominated with respect to abundance (~ 80 % of the total individuals) by Acari, Collembola and Hymenoptera (primarily ants) and functionally by scavengers and ants. Epiphyte and invertebrate communities were highly variable with respect to spatial patterning of species richness, abundance and composition across sites, among trees within sites and among branches within trees. Overall, a highly significant proportion, > 75 %, of the variance could be attributed to differences at the branch level, but these differences could not be explained by the environmental factors measured. There were no consistent relationships between the spatial pattern of epiphytes and invertebrates, or between vascular and non-vascular plants. However, there were significant positive correlations between epiphyte biomass and invertebrate species richness (r = 0.472; p < 0.0001) and abundance (r = -0.395; p < 0.0001), as well as non-living epiphyte biomass and scavenger species richness (r = 0.4; p < 0.0001). Microclimatic measurements taken on epiphyte mats were also highly variable with respect to temperature and relative humidity at similar physical locations within the same tree as well as across trees within sites. There was also considerable variation in the intensity and frequency of climatic extremes, although potentially harmful climatic conditions were experienced by all the epiphyte mats for which weather variables were measured. Negative correlations existed between both epiphyte and invertebrate community composition and increased temperatures expressed as cumulative degree days above 5˚C. However, variability was such that there was no direct evidence that increased temperature and rainfall treatments had an effect on invertebrate species richness, abundance or diversity. Northern rata host trees harbour an astonishingly diverse and complex canopy flora and fauna that is characterised by high spatial variability. Such variability highlights that to determine species distribution and community dynamics in canopy habitats in response to disturbance caused either by climate change or invasive species the structure of entire communities at different taxonomic and spatial scales, along with their responses to microclimatic factors, need to be studied. If such complexities are not taken into account, inappropriate interpretation may result in poor decisions concerning the conservation status, vulnerability and subsequent management of such unique ecosystems.

The chemical composition and antioxidant activity of ethanolic extracts of five moss species are presented in this diploma thesis. The total phenol content was estimated as gallic acid equivalents by the Folin-Ciocalteu reagent method, while the qualitative composition of the extracts were determined by high performance liquid chromatography coupled with photodiode array detection and by gas chromatography - mass spectroscopy detection. The antioxidant properties assessed included iron(III) reduction, 1, 1-diphenyl-2-picrylhydrazyl anion free radical scavenging and the ability of extracts to protect 2-deoxy-D-ribose against hydroxyl radical- mediated degradation was assessed. The extracts contained phenolic compound. Free phenolic acids as benzoic acid derivatives and glycosides as vitexin-like structures.The ethanolic extracts of five moss species demonstrated antioxidant and free radical scavenging activity; however, they were not as potent as the positive control.

Folgende Themenkomplexe wurden behandelt: Biogeographie, Pflanzendiversität, Verbreitungsmuster von Lebensformen und Arten in Mikrohabitaten sowie entlang ökologischer Gradienten, Epiphyt-Porophyt-Beziehungen, Vergesellschaftung, Sekundärsukzession und die Auswahl von Indikatorarten. Zum ersten Mal wird bei einer derartigen Untersuchung die sekundäre Sukzession von Kryptogamengesellschaften in tropischen Bergregenwäldern voll berücksichtigt. Die Untersuchung wurde im Einzugsgebiet des Río Savegre auf der pazifikseitigen Abdachung der westlichen Cordillera de Talamanca (Costa Rica) durchgeführt.Kapitel 1 gibt eine generelle Einführung in Tropische Bergregenwälder und die Bedeutung von Moosen und Makroflechten als wichtige Komponenten dieser Ökosysteme. Der Leser wird in das Untersuchungsgebiet, sein Klima, seine Geologie und seine Vegetation eingeführt.Im Kapitel 2 werden die pflanzengeographischen Muster der Moose montaner Eichenwälder der Cordillera de Talamanca dargestellt und diskutiert.Kapitel 3 diskutiert Diversität, Mikrohabitatdifferenzierung und Lebensformenspektren der Moose in einem hochmontanen Eichenwald Costa Ricas. Es wurde versucht, die Moosflora aller Mikrohabitate eines Eichenwaldes, inklusive der Strauch- und Bodenschicht zu erfassen. Kapitel 4 und 5 beinhalten die Beschreibung epiphytischer Kryptogamengesellschaften (Moose und Makroflechten) in Primär- und Sekundärwäldern. Mit Hilfe einer aus dem Alpinismus abgeleiteten Klettertechnik wurden Bäume von der Stammbasis bis in die Zweige der äußeren Krone beprobt. Die Vegetationsaufnahmen repräsentieren die Variation der epiphytischen Vegetation. Prozentuale Deckungsgrade der rindenbewohnenden Moose und Makroflechten wurden geschätzt und mit Hilfe einer Detrended Correspondence Analysis (DCA)' und nicht-metrischer Multi-response Permutation Procedure (MRPP)' verglichen. Rindenproben der untersuchten Bäume wurden mit deionisiertem Wasser extrahiert und der pH-Wert bestimmt.Kapitel 4 behandelt die epiphytischen Kryptogamengemeinschaften auf den dominierenden Baumarten (Quercus copeyensis und Q. costaricensis) primärer, hochmontaner Eichenwälder Costa Ricas. Kapitel 5 vergleicht Artenreichtum, Vergesellschaftung und Ökologie der epiphytischen Kryptogamen in primären und sekundären hochmontanen Eichenwäldern Costa Ricas. Quercus copeyensis-Bäume der oberen Baumschicht von Primär- und Sekundärwäldern wurden beprobt mit dem Ziel einen Einblick in Regenerationspotential, -muster und -prozesse der Epiphytensukzession nach anthropogener Störung zu erhalten.

The aim of this thesis was to create potential distribution models for Dicranum majus (Greater Fork Moss) and Polytrichum alpinum (Alpine Haircap) in Bohemian Switzerland National Park. In the Czech Republic these bryophyte species occur in cold climatic regions typically with higher altitudes. In Bohemian and Saxon Switzerland they can occur in really low altitudes thanks to unique microclimatic conditions of deep inversion ravines. These bryophyte species had low number of occurence records in studied area before the start of my research (4 occurence localities for Dicranum majus, 8 occurence localities for Polytrichum alpinum). Predictive habitat suitability models can be an effective tool for selecting potential new occurence localities, planning field research or management design. During field research I recorded 34 new occurence localities for Dicranum majus and 29 new occurence localities for Polytrichum alpinum in Bohemian Switzerland National Park. I used 8 topographic parameters derived from digital elevation model with 1 m resolution as environmental data. Using these data I created models of potential distribution of the most suitable habitats for both species with algorithms Artificial neural networks (ANN), Generalised linear model (GLM) and Random forest (RF). RF algorithm had the...

Dry grassland are among the most endangered plant communities mainly due to the overgrowth of a number of localities. That is why from the beginning of the 21st century pasture with mixed herds of sheep and goats is restored on the most valuable dry grassland sites in the Bohemian Karst as the way of maintaining these habitats. Concurrently, research sites each 1x1 m in size were set up at four pastured localities, and were paired with control sites equipped with a fence for the research of grazing effect on vegetation. The main aim of this work was to evaluate the effect of grazing on the diversity of bryophytes at the forty pairs of the above-mentioned areas in three types of calcareous dry grasslands. In addition to the inventory of bryophytes, data on the most important abiotic and biotic factors of the environment were recorded on individual sites. The total number of bryophyte species found at the research sites is 46. 37 species were found at the grazing areas and the same number was at the fenced areas. Minimal differences in the number of species of grazed and unspoilt areas were also found in the comparison of individual localities. The grazing effect was shown up more significantly on species richness in the comparison of individual types of communities, especially in broad-leaved dry grassland (T3.4). There was a 40% increase in the number of species on grazing areas. The environmental factors have a positive effect on the species abundance mainly due to the higher slope of the terrain and the correlation with the larger occurrence of the stones, the smaller coverage of the herb layer and the litter. These conditions prevailed in areas with rock outcrop vegetation with Festuca pallens (T3.1), where the largest species richness of bryophytes in area was found. In the obtained data, the influence of grazing on the species composition of bryophytes on grazed and unspoilt areas was also assessed with the statistical methods and the Sörensen test of beta-diversity. The results show that grazing significantly influences the species composition towards open grassland and supports the growth of the bryophytes T3.1 plant community at the expense of the T3.4 plant community

The life of bryophytes is closely connected with their surrounding environment. Changes in atmospheric conditions on the microscale directly affect the physiological functions of bryophytes, which in turn determine their distribution. The current development of technologies allows us to measure the microclimate affecting bryophytes directly in the field. Nevertheless, there have not been many studies published examining the response of bryophytes to in situ measured microclimate. This diploma thesis is one of the first in Europe to provide data from continuous field microclimatic measurement performed on a scale relevant to bryophytes. It deals with the influence of the microclimate on the bryophyte species richness and community composition within a single gorge in the Bohemian Switzerland National Park. For 17 months, 38 HOBO Pro v2 Onset dataloggers were recording air temperature and air humidity 10 cm above the ground. I performed detailed bryological survey on two differently sized plots (circle with a radius of 1 or 2 m) around each of the dataloggers. The microclimate is influenced by the topography and character of the vegetation, therefore I derived topographical data from a digital terrain model (resolution of 1 m), calculated the canopy openness using hemispherical photographs and...

Marine Inseln beherbergen einen großen Teil der biologischen Vielfalt unseres Planeten und weisen gleichzeitig einen hohen Anteil endemischer Arten auf. Inselbiota sind allerdings zudem besonders anfällig für anthropogene Einflüsse wie den globalen Klimawandel, Habitatverlust und invasive Arten. Für ihren Erhalt ist es daher wichtig, die ökologischen Prozesse auf Inseln detailliert zu verstehen. Aufgrund ihrer definierten Größe und isolierten Lage eignen sich Inseln als Modellsysteme in der ökologischen und evolutionären Forschung. Der Großteil der bisherigen Inselstudien hat sich allerdings mit kleinräumigen Mustern befasst, so dass standardisierte globale Daten zu den biogeographischen Eigenschaften und eine makroökologische Synthese ihrer Biota bislang fehlen. In dieser Arbeit stelle ich eine physische und bioklimatische Charakterisierung der Inseln der Welt vor und behandle die Frage, wie abiotische Inseleigenschaften die Diversität von Inselfloren beeinflussen. Ich bearbeite zwei Hauptaspekte dieser Fragestellung: Zuerst konzentriere ich mich auf historische und heutige Klimabedingungen und physische Inseleigenschaften als Triebfedern von Pflanzendiversitätsmustern auf Inseln. Hierbei setze ich einen Schwerpunkt auf die räumliche Anordnung von Inseln und Struktur von Archipelen. Als Zweites behandle ich taxon-spezifische Unterschiede in der Antwort von Diversitätsmustern auf abiotische Faktoren. Hierzu stelle ich eine globale Datenbank mit historischen und heutigen Klimabedingungen und physischen Eigenschaften, wie Fläche, Isolation und Geologie, von 17883 Inseln größer als 1 km² vor. Mit Hilfe von Ordinations- und Klassifikationsverfahren charakterisiere und klassifiziere ich die Inseln in einem multidimensionalen Umweltraum. Außerdem entwickele ich einen Satz von ökologisch relevanten Maßen zur Beschreibung von Isolation von Inseln und ihrer räumlichen Anordnung in Archipelen, darunter Maße zu Trittstein-Inseln, Wind- und Meeresströmungen, klimatischer Ähnlichkeit, Distanzen zwischen Inseln und umgebender Landfläche. Diese Maße berücksichtigen verschiedene Aspekte von Isolation, welche Immigration, Artbildung und Aussterben auf Inseln sowie Austausch zwischen Inseln beeinflussen. Um abiotische Bedingungen mit biotischen Eigenschaften von Inselfloren in Verbindung zu bringen, nutze ich eine für diese Arbeit erstellte Datenbank aus 1295 Insel-Artenlisten, die insgesamt ca. 45000 heimische Gefäßpflanzenarten umfassen. Dies ist der umfassendste und erste globale Datensatz für Pflanzen auf Inseln, der Artidentitäten anstatt lediglich Artenzahlen beinhaltet. Die globale Insel-Charakterisierung bestätigt quantitativ, dass sich Inseln in bioklimatischen und physischen Eigenschaften vom Festland unterscheiden. Inseln sind im Durchschnitt signifikant kühler, feuchter und weniger saisonal geprägt als das Festland. Die weiteren Ergebnisse zeigen, dass eine sorgfältige Beschreibung der räumlich-physischen Eigenschaften von Inseln und Archipelen nötig ist, um die Diversitätsmuster ihrer Biota zu verstehen. Isolation ist nach Inselfläche der zweitwichtigste Einflussfaktor für den Gefäßpflanzenartenreichtum auf Inseln. Von den verglichenen Isolationsmaßen eignet sich der Anteil an umgebender Landfläche am besten zur Erklärung der Artenzahlen. Außerdem erhöht sich durch die Berücksichtigung von Trittsteininseln, großen Inseln als Quell-Landflächen und klimatischer Ähnlichkeit der Quell-Landflächen die Vorhersagekraft der Modelle. Isolation spielt eine geringere Rolle auf großen Inseln, wo in situ Diversifizierung den negativen Effekt von Isolation auf Immigration ausgleicht. Die räumliche Struktur innerhalb von Archipelen ist von besonderer Bedeutung für β-Diversität, d.h. für den Unterschied in der Artenzusammensetzung der Inseln. Außerdem beeinflusst sie indirekt, durch den Effekt auf die β-Diversität, auch die γ-Diversität, d.h. die Diversität des gesamten Archipels. Die Ergebnisse heben die enorme Bedeutung der relativen räumlichen Position von Inseln zueinander für Diversitätsmuster auf Inseln hervor und zeigen die Notwendigkeit für Inselforschung und Naturschutz, Inseln im Kontext ihres Archipels zu betrachten. Die Ergebnisse für Farne auf südostasiatischen Inseln zeigen, dass die Bedeutung von physischen Inseleigenschaften für Diversität kontinuierlich mit der Größe der betrachteten Untersuchungsfläche von der Insel- bis zur Plotebene abnimmt, wohingegen der Einfluss von lokalen Umweltbedingungen zunimmt. Lokale Artgemeinschaften sind häufig gesättigt, wodurch die Anzahl an Arten, die aus dem regionalen Artenbestand einwandern können, limitiert wird. Um Vorhersagen über lokalen Artenreichtum zu machen, ist es daher wichtig, die Skalenabhängigkeit der Effekte des regionalen Artenbestandes zu berücksichtigen. Großgruppen von Pflanzen unterscheiden sich in ihrer Ausbreitungsfähigkeit, ihrem Genfluss, Artbildungsraten und Anpassungen an das Klima. Dementsprechend zeigen die vergleichenden Analysen zwischen taxonomischen Pflanzengruppen deutliche Unterschiede in der Reaktion von Artenreichtum und phylogenetischen Diversitätsmustern auf abiotische Faktoren. Die Arten-Fläche-Beziehung, d.h. die Zunahme von Artendiversität mit zunehmender Fläche, variiert zwischen den Pflanzengruppen. Die Steigung der Arten-Fläche-Beziehung ist für Spermatophyten größer als für Pteridophyten und Bryophyten, wohingegen der y-Achsenabschnitt kleiner ist. Unter der Annahme, dass Merkmale und klimatische Anpassungen innerhalb von taxonomischen Gruppen phylogenetisch konserviert sind, führen die Filterwirkung von Ausbreitungsbarrieren und Umwelteigenschaften sowie in situ Artbildung zu Gemeinschaften eng verwandter Arten (phylogenetic clustering). Die Ergebnisse zeigen, dass physische und bioklimatische Inseleigenschaften, die mit der Filterwirkung und Artbildung in Verbindung stehen, die phylogenetische Struktur von Inselgemeinschaften beeinflussen. Die Stärke und Richtung der Zusammenhänge variieren zwischen taxonomischen Gruppen. Abiotische Faktoren erklären mehr Variation in phylogenetischer Diversität für alle Angiospermen und Palmen als für Farne, was auf Grund höherer Ausbreitungsfähigkeit und größerer Verbreitungsgebiete von Farnen den Erwartungen entspricht. Die abiotische Charakterisierung und Klassifizierung der weltweiten Inseln und die zugehörigen Daten ermöglichen eine integrativere Berücksichtigung von Inseln in der makroökologischen Forschung. In dieser Arbeit präsentiere ich die ersten Vorhersagen globaler Pflanzenartenvielfalt auf Inseln und die ersten Analysen zu unterschiedlichen Diversitätskomponenten (α, β, γ und phylogenetische Diversität) von Inselsystemen und ihren abiotischen Einflussfaktoren auf globalem Maßstab. Ich zeige, dass Zusammenhänge zwischen Umweltfaktoren und Artenzahl sowie phylogenetischen Eigenschaften von Inselgemeinschaften zwischen unterschiedlichen taxonomischen Gruppen in Abhängigkeit ihrer vorwiegenden Ausbreitungs- und Artbildungseigenschaften variieren können. Dies ist eine neue Sichtweise in der makroökologischen Inselforschung, die Rückschlüsse auf die Mechanismen hinter Diversitätsmustern von Pflanzen auf Inseln erlaubt. Ein detailliertes Verständnis davon, wie Diversität unterschiedlicher Pflanzengruppen durch Immigration und Diversifizierung auf Inseln entsteht, dürfte auch das Verständnis globaler Diversitätsmuster im Allgemeinen verbessern.