This dissertation explored the way in which bumblebees' visual system helps them discover their first flower. Previous studies found bees have unlearned preferences for parts of a flower, such as its colour and shape. The first study pitted two variables against each other: pattern type: sunburst or bull's eye, versus the location of the pattern: shapes appeared peripherally or centrally. We observed free-flying bees in a flight cage using Radio-Frequency Identification (RFID) tracking. The results show two distinct behavioural preferences: Pattern type predicts landing: bees prefer radial over concentric patterns, regardless of whether the radial pattern is on the perimeter or near the centre of the flower. Pattern location predicts exploration: bees were more likely to explore the inside of artificial flowers if the shapes were displayed near the centre of the flower, regardless of whether the pattern was radial or concentric. As part of the second component, we implemented a mathematical model aimed at explaining how bees come to prefer radial patterns, leafy backgrounds and symmetry. The model was based on unsupervised neural networks used to describe cognitive mechanisms. The results captured with the results of multiple behavioural experiments. The model suggests that bees choose computationally "cheaper" stimuli, those that contain less information. The third study tested the computational load hypothesis generated by the artificial neural networks. Visual properties of symmetry, and spatial frequency were tested. Studying free-flying bees in a flight cage using motion-sensitive video recordings, we found that bees preferred 4-axis symmetrical patterns in both low and high frequency displays.

Doctor of Philosophy / Department of Horticulture, Forestry, and Recreation Resources / Kimberly A. Williams / The use of calcined clays contributes properties of nutrient and water retention to soilless root media, which varies greatly depending on the parent clay and calcining treatment. This research characterized phosphate (PO[subscript]4) sorption of various calcined clay products, including low volatile and regular volatile material (LVM and RVM) 2:1 Attasorb clays (Engelhard Corp.), 2:1 Terra Green LVM clays (Oil-Dri Co.), and Turface (Profile Products LLC) at various particle sizes; 1:1 kaolin clays (Thiele Kaolin Co.) in powder form, and diatomaceous earth (Diatomite, Eagle Picher Minerals, Inc.). Three of the calcined materials, Terra Green montmorillonite and Attasorb attapulgite (which had high PO[subscript]4-sorption based on isotherms), and diatomaceous earth (which had negligible PO[subscript]4-sorption) were evaluated as components of soilless root media in two separate greenhouse experiments. The effect of the calcined materials, rate of incorporation (0%, 5%, 10% and 20% by volume in a mix with peat and perlite), and PO[subscript]4-P application rate (0, 5, 15, 45 mg.L[superscript]-1 PO[subscript]4-P) on plant growth, effluent P content and water use were determined during production and post-production of Impatiens wallerana Hook f. 'Tempo Rose'. The calcined materials varied in their ability to adsorb PO[subscript]4-P and generally yielded L-type isotherms. Laboratory results indicated potential for substantive P retention by several of the calcined materials when used in container production. For most materials, PO[subscript]4-P sorption did not show pronounced pH dependence. During production and post-production, the test materials not only improved PO[subscript]4-P retention but also water retention and water use efficiency while still maintaining optimal physical properties at incorporation rates of 5 to 10%. Diatomaceous earth resulted in PO[subscript]4-P retention not significantly different from the calcined clays.

At certain stages of a bumblebee colony life cycle workers lay eggs. Not all workers reproduce, however, since many continue to forage and care for the nest. This leads to questions regarding what differentiates a reproductive worker from a non-reproductive one. It is hypothesized that a form of reproductive competition takes place, where the most behaviourally dominant worker becomes reproductively dominant. The behaviour of orphaned Bombus impatiens pairs was recorded and aggression, social interactions, egg-laying, and ovarian development were identified. Experiment 1 examined the association between aggression and egg-laying. Contrary to the hypothesis, the most aggressive worker did not lay more eggs. When the ovarian development of workers was manipulated and two workers with developed ovaries were paired (Experiment 3), they were more aggressive than pairs with discouraged ovarian development. This provides support for the supposition that aggression and reproduction are related, however, it is only partial support as worker pairs with encouraged ovarian development did not lay more eggs. Since aggression is believed to be only one part of behavioural dominance, Experiment 2 studied the association between social interactions and aggression and reproduction. Results showed that when two socially active bees were paired they were more aggressive than pairs including one or two socially inactive bumblebees. No significant difference in ovarian development between socially active pairs and socially inactive pairs was found. Brood presence was also predicted to affect reproductive control. Experiment 1 found egg-laying and aggression were more likely to co-occur in the absence of brood. Results from Experiment 2 supplemented the first experiment since the absence of brood increased rates of aggression and ovarian development in pairs. Whereas the results confirm aggression has a role in worker reproduction the findings also reveal that behavioural dominance does not equate to reproductive dominance under all conditions. The primary contributions of this thesis were the development of a method to distinguish behavioural dominance from reproductive dominance and determining their relationship under different environments (brood presence) and experimental manipulations (ovarian development). These contributions further define dominance in Bombus impatiens.

This dissertation explored the way in which bumblebees' visual system helps them discover their first flower. Previous studies found bees have unlearned preferences for parts of a flower, such as its colour and shape. The first study pitted two variables against each other: pattern type: sunburst or bull's eye, versus the location of the pattern: shapes appeared peripherally or centrally. We observed free-flying bees in a flight cage using Radio-Frequency Identification (RFID) tracking. The results show two distinct behavioural preferences: Pattern type predicts landing: bees prefer radial over concentric patterns, regardless of whether the radial pattern is on the perimeter or near the centre of the flower. Pattern location predicts exploration: bees were more likely to explore the inside of artificial flowers if the shapes were displayed near the centre of the flower, regardless of whether the pattern was radial or concentric. As part of the second component, we implemented a mathematical model aimed at explaining how bees come to prefer radial patterns, leafy backgrounds and symmetry. The model was based on unsupervised neural networks used to describe cognitive mechanisms. The results captured with the results of multiple behavioural experiments. The model suggests that bees choose computationally "cheaper" stimuli, those that contain less information. The third study tested the computational load hypothesis generated by the artificial neural networks. Visual properties of symmetry, and spatial frequency were tested. Studying free-flying bees in a flight cage using motion-sensitive video recordings, we found that bees preferred 4-axis symmetrical patterns in both low and high frequency displays.

L’environnement a un effet considérable sur les végétaux par différents facteurs abiotiques (climat, sol, urbanisation) ou biotiques (pollinisateurs, herbivores). Les fortes pressions de sélection exercées par ces facteurs sur certains traits phénotypiques aboutissent souvent à des adaptations chez les plantes. Les pollinisateurs exercent une pression de sélection sur les traits floraux qui résultent en des adaptations convergentes que l’on appelle syndromes de pollinisation. L’étude de ses syndromes, mais aussi de la performance de pollinisation de chaque visiteur, permet de mieux comprendre l’intensité de cette pression de sélection ainsi que l’évolution florale. Dans cette thèse, nous nous sommes appuyés sur l’utilisation de deux modèles d’études végétaux : la famille des Gesneriaceae des Antilles et l’Impatiente du Cap. Ces deux modèles, de par leurs stratégies de pollinisation et leurs variations florales, sont appropriés pour étudier l’adaptation florale aux pollinisateurs. Nous avons ainsi pu tester 1) si la forme florale est expliquée par la forme du bec des colibris pollinisateurs chez les Gesneriaceae antillaises ; 2) si l’espèce Rhytidophyllum bicolor Urb. est généraliste en pollinisation et si ses différents types fonctionnels de pollinisateurs ont une performance de pollinisation similaire ; 3) si l’urbanisation affecte la forme florale de l’espèce Impatiens capensis Meerb., à travers des changements dans les communautés de pollinisateurs. Pour tester ces hypothèses, des approches de morphométries géométriques ont été utilisées (hypothèses 1 et 3), ainsi que des observations de pollinisation in situ et la mesure du taux de visite (hypothèses 2 et 3). La mesure de performance de pollinisation a été réalisé via le comptage des grains de pollen déposés sur le stigmate après chaque visite, et à la mesure du taux de visite. Enfin, des tests statistiques (ANOVA, régressions linéaires) et analyses multivariées (analyses de redondance, analyse en composantes principales) ont été réalisées sur les données obtenues pour chaque étude. Les résultats de la première étude montrent une corrélation positive entre la longueur de la corolle de fleurs spécialistes aux colibris et la longueur du bec des pollinisateurs. Bien que moins significatifs, les résultats des comparaisons de formes globales, obtenues par application de morphométrie géométrique, indiquent que cette approche est prometteuse pour ce genre d’analyse. Nous avons démontré que la forme florale des fleurs généralistes est impactée par les colibris pollinisateurs, bien que d’une manière différente des spécialistes. 4 Pour l’hypothèse 2, la prédiction de stratégie de pollinisation de l’espèce Rhytidophyllum bicolor a été validée, puisque cette plante est pollinisée par des colibris, chauves-souris et abeilles. Cependant, face au déclin des populations de colibris après le passage de l’ouragan Matthew sur Haïti, seules les performances de pollinisation des abeilles et des chauves-souris ont pu être mesurées. Les résultats montrent que les chauves-souris sont des pollinisateurs efficaces et conséquents, bien que la performance des abeilles ne soit pas négligeable. Il a ainsi pu être mis de l’avant que la stratégie de pollinisation généraliste semble être un avantage pour les plantes présentes dans les zones sujettes aux fluctuations de populations de pollinisateurs, comme cela peut souvent être le cas sur les îles à la suite du passage d’un ouragan. La troisième étude montre que l’urbanisation n’a pas d’effet direct sur la forme florale de l’Impatiente du Cap, mais qu’elle a des effets indirects significatifs via les changements causés sur les communautés de pollinisateurs. Sur les six sites échantillonnées les pollinisateurs principaux, Bombus sp. Latreille et Apis mellifera Linnaeus, sont les mêmes. Cependant les taux de diversité obtenus montrent une variation entre les sites, due à la présence de certaines espèces de pollinisateurs occasionnels dans certains sites et pas dans d’autres. Ces taux ne sont pas plus faibles dans les sites les plus urbains. Les résultats indiquent que certaines formes florales sont associées à des espèces de pollinisateurs particulières. Les différentes espèces de pollinisateurs ayant une pression de sélection différente sur les traits floraux, l’urbanisation a ainsi un impact indirect sur la forme florale chez l’espèce étudiée. À travers trois études différentes, cette thèse a mis en avant l’impact que l’environnement peut avoir sur les traits floraux, de manière indirecte, via les pollinisateurs. Alors que la deuxième et troisième étude ont montré la pression exercée par les pollinisateurs sur les traits floraux dans différents cas de perturbations de l’habitat, la première étude a permis de mieux comprendre l’adaptation remarquable des fleurs à leurs pollinisateurs, même pour des espèces généralistes en pollinisation. / The environment has a considerable effect on plants through various abiotic (climate, soil, urbanization) or biotic (pollinators, herbivores) factors. The strong selection pressures exerted by these factors on phenotypic traits often results in adaptations. Pollinators exert selection pressure on floral traits that result in converging adaptations called pollination syndromes. The study of syndromes, as well as the pollination performance of each floral visitor, allows us to better understand the intensity of a/biotic selection pressure and floral evolution. In this thesis, we relied on two plant models: the Gesneriaceae family in the Antilles, and the common Jewelweed, Impatiens capensis Meerb. Due to their pollination strategies and their floral variation, both of these models are suitable for studying floral adaptation to biotic factors, specifically, their pollinators. We were thus able to test the following hypotheses: 1) whether the floral form is explained by the beak shape of pollinating hummingbirds in West Indies Gesnericeae; 2) whether the Gesnericeae Rhytidophyllum bicolor Urb. has a generalist strategy for pollination and whether their different functional types of pollinators have similar pollination performances; 3) whether urbanization affects the floral form of the common Jewelweed, through changes in pollinator communities. To test these hypotheses, geometric morphometric approaches were used (hypotheses 1 and 3), as well as in situ pollination observations, and estimation of the visitation rate (hypotheses 2 and 3). Pollination performance was measured by counting pollen grains deposited on the stigma after each visit, and by measuring the visitation rate. Finally, statistical tests (ANOVA, linear regressions) and multivariate analyses (redundancy analysis, principal component analysis) were carried out on the data obtained to test each hypothesis. The first results show a positive correlation between the length of the corolla of flowers specialized for hummingbirds and the beak length of pollinators. Although less significant, our results of the comparisons of global shapes, obtained by applying geometric morphometry, indicate that this approach is promising for this kind of analysis. We show that the floral form of generalist flowers is impacted by pollinating hummingbirds, albeit in a different way from specialists. For the second hypothesis, our prediction that R. bicolor has a generalist pollination strategy was validated, since this plant is pollinated by hummingbirds, bats and bees. However, faced with the decline in hummingbird populations after Hurricane Matthew hit Haiti in 2016, only the pollination performance of bees and bats could be measured. Our results show that bats are efficient and consistent pollinators of R. bicolor, although the performance of bees is not negligible. Thus, it has been possible to put forward that the generalist pollination strategy seems to be an advantage for plants present in areas subject to fluctuations in their pollinator populations, as can often be the case on hurricane-prone islands. The third study shows that urbanization does not have a direct effect on the flower form of common Jewelweed, but that urbanization does have significant indirect effects through changes caused on pollinator communities. At the six sites sampled, the main pollinators, Bombus sp. Latreille and Apis mellifera Linnaeus, are the same. However, diversity rates show variation between sites, due to the presence of certain species of occasional pollinators in some sites and not in others. These rates are not lower in the most urban sites. Our results indicate that certain floral forms are associated with particular pollinator species. Since different pollinator species have different selection pressures on floral traits, urbanization has an indirect impact on the floral form in the species studied. Through three different studies, this thesis highlighted the impact that the environment can have on floral traits, indirectly, via pollinators. While the second and third studies showed the pressure exerted by pollinators on floral traits in different cases of habitat disturbance, the first study helped to better highlight the remarkable adaptation of flowers to their pollinators, even for generalist species in pollination.

Plant invasions represent a major ecological and socio-economical issue and understanding the drivers as well as consequences of plant invasions is thus one of the main goals of plant ecology. It is equally important to reveal general patterns underlying plant invasions and to understand the details of biology of individual invaders. In this thesis I explored plant-soil feedback (PSF) as a possible general mechanism underlying plant invasiveness, and also focused in detail on drivers and consequences of Impatiens parviflora invasion. The aims of this thesis were to i) assess the differences in intraspecific PSF between invasive and alien non-invasive species using a large set of species; ii) explore the relationship between PSF, residence time and phylogenetic novelty of the alien species; iii) compare the importance of PSF and other plant characteristics for plant invasiveness; iv) compare PSF between invasive and native congeners of similar level of dominance in the field; v) evaluate the effect of cultivating conditions on results of PSF experiments; vi) describe invasion dynamics and determine factors affecting spread of invasive I. parviflora using a method of monitoring its natural spread in several types of habitats, and vii) assess the impact of I. parviflora on native vegetation of oak-...

Les quinones sont des composés largement répandus dans la nature, aussi bien dans le règne végétal, qu'animal ou microbien. Ces molécules sont biosynthétisées à partir d'un petit nombre d'intermédiaires simples. Le noyau des naphtoquinones, en particulier, peut être synthétisé à partir de quatre précurseurs principaux l'acétate, la toluhydroquinone, le p-hydroxybenzoate et le shikimate. L'objet du présent travail est l'étude de la biosynthèse des naphtoquinones végétales ou bactériennes dérivant de l'acide shikimique (acide trihydroxy-3,4,5 cyclohexène-1 carboxylique), et la recherche des intermédiaires de cette voie de biosynthèse. Etant donné, à l'origine, l'absence quasi-totale de résultats expérimentaux dans ce domaine et le développement important qu'il a acquis en même temps que nous commencions notre travail, celui-ci a été constamment lié dans son développement aux données les plus récentes parues dans la littérature. Quelques études antérieures avaient montré que l'acide shikimique devait être le précurseur commun des acides aminés aromatiques et de plusieurs facteurs de croissance aromatiques (acide dihydroxy-2,3 benzoique, dihydroxy-3,4 benzoique, p-aminobenzoique et p-hydroxybenzoique) ainsi que de la ménaquinone de Escherichia coli (16,17). Lorsque nous commençons ce travail, on sait seulement que l'acide shikimique est incorporé in toto dans les ménaquinones bactériennes, de telle sorte que son carboxyle devient un des carbonyles de la quinone. Trois questions se posent alors: - Sur lequel des carbones 2 ou 6 adjacents au carboxyle, le cycle naphtoquinonique se ferme-t-il ? - Quelle est la molécule qui fournit les trois carbones nécessaires à la fermeture du cycle quinonique de la naphtoquinone, et quel est le mécanisme de cette fermeture ? - Le cycle naphtoquinonique ainsi formé est-il incorporé de façon symétrique ou non dans les ménaquinones, c'est-à-dire le carboxyle de l'acide shikimique est-il incorporé indifféremment dans les deux carbonyles quinoniques, ou sélectivement dans l'un des deux ? Pour résoudre la première question, LEISTNER et ZENK d'une part et M. LEDUC dans notre laboratoire d'autre part, ont utilisé l'acide shikimique [14 C-1,6]. Cette méthode nécessite une dégradation laborieuse des quinones formées. Nous avons, quant à nous, synthétisé un acide shikîmique [3H-3] marqué au tritium en une position spécifique, ce qui permet une dégradation simple et univoque. Une réponse partielle à la deuxième question a été donnée par CAMPBELL. Il montrait, en 1969, que le glutamate est capable d'apporter ses carbones 2, 3 et 4 pour la fermeture du cycle naphtoquinonique, mais il ne disposait pas alors des résultats de notre laboratoire lui permettant d'induire correctement le mécanisme de cette biosynthèse. Nous avons pu postuler la formation d'un intermédiaire, I'acide ortho-succinoylbenzoique OBS, dont nous avons pu prouver l'existence par synthèse et incorporation dans les quinones étudiées . L'existence de cet intermédiaire et la possibilité que nous avons eue de réaliser sur l'OSB différents marquages spécifiques nous ont permis de répondre à la troisième question, à savoir, l'incorporation orientée ou non de l'OSB dans les quinones. En même temps, différentes équipes montraient que d'autres quinones existant dans les végétaux telles que la lawsone, la juglone et différentes anthraquinones dérivaient aussi de l'acide shikimique; il nous a semblé intéressant d'appliquer nos techniques à l'étude très voisine de la biosynthèse de ces quinones. Pour la clarté de l'exposé, nous présentons dans le premier chapître les méthodes de synthèse des précurseurs radioactifs, dans le deuxième, les méthodes d'isolement et de dégradation des quinones. Les trois chapîtres suivants rapportent l'incorporation dans les ménaquinones, les naphtoquinones végétales et les anthraquinones végétales. Dans le sixième chapître, nous discutons nos résultats et proposons un schéma général de biosynthèse des quinones dérivant de l'acide shikimique. La partie expérimentale de ce travail est rassemblée après l'exposé théorique.

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Buchenwälder sind die großflächigste potenziell natürliche Vegetationsform Deutschlands und ein nach EU-FFH-Richtlinie besonders zu schützender Biotoptyp. Eine hohe Naturnähe ist auch in Wirtschaftswäldern (WiWald) notwendig, um die typischen Lebensgemeinschaften naturnaher Wälder langfristig zu erhalten, doch mangelt es an praktikablen/verifizierten Indikatoren, wie die nutzungsbedingte Abweichung vom Naturzustand ermittelt werden kann. In >100 Jahre alten und ~40 ha großen Tiefland-Buchenwäldern (Mecklenburg-Vorpommern/Brandenburg) wurde anhand von 13 WiWäldern, vier seit <20 Jahren (k20) und drei seit >50 Jahren (r50) unbewirtschafteten Beständen den folgenden Fragen nachgegangen: Wie groß sind die strukturellen, vegetationskundlichen und carabidologischen Unterschiede zwischen bewirtschafteten, kurz- und langfristig unbewirtschafteten Buchenwäldern? Gibt es strukturelle Indikatoren und quantitative Größen zur Abschätzung des Einflusses forstlicher Bewirtschaftung auf die Biozönosen von Tiefland-Buchenwäldern? In Probekreisen (Pk) von 500 m² an Rasterpunkten (100 m x 100 m) wurden strukturelle und in Pk von 314 m² vegetationskundliche Daten erhoben. In fünf Pk/Bestand wurde jeweils eine Barberfalle über die Vegetationsperiode installiert. Ganzflächig wurden die Verteilung der Waldentwicklungsphasen (WEP)und zusätzlich zu den Pk-Aufnahmen hektarweise Sonderstrukturen aufgenommen. U. a. wurden folgende Sonderstrukturen aufgenommen: Zunderschwamm, Kronen- und Zwieselbrüche, Ersatzkronen, Blitzrinnen, Risse/Spalten, Höhlen, Mulmkörper/-taschen. Diese naturschutzfachlich wichtigen Sonderstrukturen wurden aus den Habitatansprüchen der typischen Buchenwaldfauna abgeleitet.Es konnten große Unterschiede zwischen WiWald und r50-Flächen (v. a. >100 Jahre unbewirtschafteten Flächen) aufgezeigt werden. Die k20-Flächen unterscheiden sich nicht wesentlich vom WiWald. Die Anzahl verschiedener WEP/ha und WEP-Patches/ha liegt in den r50-Flächen signifikant höher als im WiWald. Der Holzvorrat der r50-Flächen liegt mit ~600 m³/ha (Terminal- ~800 m³/ha, Zerfallsphase 450 m³/ha) deutlich höher als im WiWald. Charakteristisch für die r50-Flächen ist das Vorkommen von in ihrer Vitalität eingeschränkten Bäume ab 80 cm BHD und ein inhomogeneres Lichtmosaik im Bestand. Die Stammqualitäten (u. a. Astigkeit) in r50-Flächen unterscheiden sich kaum von denen in WiWald. In den r50-Flächen kommt bedeutend mehr Totholz (>142 m³/ha) als im WiWald (max. 34 m³/ha) vor. Im WiWald können Stubben dominieren. Verschiedene Totholzqualitäten sind im WiWald nur unvollständig vorhanden. Etwa 40 % des Totholzes besitzt keine Totholznachbarn (r50-Flächen: <2 %) und die Lichtverhältnisse am Totholz sind nicht so vielfältig (wenig sonnenexponiert und wenig gering besonnt). In den >100 Jahre unbewirtschafteten Flächen kommen ~12 Sonderstrukturtypen mit >200 Sonderstrukturen/ha vor. 19 von 20 Sonderstrukturen sind im WiWald signifikant seltener und 11 Sonderstrukturen sind als Naturnähe-Indikatoren geeignet.Vegetation: In der Krautschicht sind höhere Deckungsgrade, mehr (lichtanzeigende) Arten, weniger Waldarten und eine höhere Diversität zu verzeichnen. Im WiWald wird u. a. das Vorkommen von Calamagrostis epigeios, Impatiens parviflora und Rubus idaeus gefördert. Stark gefährdete Moosarten sind im WiWald seltener als in den Referenzwäldern, da sie vor allem auf liegendem Totholz und auf den Stammanläufen vorkommen. Carabiden: Im WiWald gibt es weniger Individuen und Biomasse von mesophilen Waldarten und eine geringere Anzahl von flugunfähigen Individuen. Als Indikatoren für naturnahe Tiefland-Buchenwälder können die drei Arten Carabus glabratus, C. hortensis und Cychrus caraboides bezeichnet werden. Indikatoren: Es wurden Zielgrößen für 29 Struktur-Indikatoren für naturnahe Wälder vorgeschlagen. Für WiWälder wurden gesonderte Zielgrößen festgelegt, die die nutzungsbedingte, nicht zu vermeidende Abweichung vom Naturzustand berücksichtigen. / Beech forests are the most important natural vegetation type of Germany,and they are included in annex II of the EU-FFH-Directive,which requests nature conservation for the listed habitat types.High naturalness is necessary in managed forests (w-sites) to maintain the typical biocoenosis of forests near nature. But there is a lack of practicable/verified indicators to determine the degree of alteration managed forests have compared to natural forests. In >100 year old and ~40 ha big lowland beech forests in Mecklenburg-Vorpommern and Brandenburg, 13 w-sites, 4 study sites which are unmanaged since <20 years (k-sites) and 3 sites which are unmanaged since >50 years (r50-sites) were investigated to answer these questions: What the differences are between w-, k- and r-sites according to forest structure, vegetation and carabids? Are there valid structural indicators with thresholds to assess the impact of forestry use on the biocoenosis of lowland beech forests? At grid points(distance 100 mx 100 m),on circular sample plots (SP) of 500 m² the structural data and on SP of 314 m² the vegetation was investigated. At five SP/study site a pitfall trap was installed during the entire vegetation period. On the whole study site the distribution of forest development phases (FDP) was mapped, and on full one ha plots the special structures were investigated. The following special structures were mapped e.g. Fomes fomentarius trees, crown and crotch breakage, substitute crowns, lightning shakes,gutters/rifts, cavities, mould and bark bag. These special structures have been derived from the habitat needs of the typical beech forest fauna.The results revealed tremendous differences between w- and r50-sites. The k-sites show no clear differences to the managed sites.In the r50-sites, the number of different FDP/ha and FDP units/ ha is significant higher than in w-sites. The timber stock of the r50-sites is ~600 m³/ha (terminal phase ~800 m³/ha, decay phase ~450³/ha). A characteristic feature of the r50-sites is the occurrence of trees with 80 cm bhd or more with reduced vitality. The timber trunk) qualities of r-sites differ only slightly from managed stands. In the r50-sites the dead wood volume (>142 m³/ha) is much higher than in the w-sites (max. 34 m³/ha). Many different features of dead wood occur only fragmentary within w-sites. About 40 % of the dead wood objects have no "dead wood neighbour" (r50-sites: <2 %), and the light distribution is much less diverse. In >100 years unmanaged r-sites ~12 different types of special structures and 200 single special structures occur per ha. 19 out of 20 special structures are significantly less frequent in w-sites; 11 special structures are specifically valuable to be used as naturalness indicators.Vegetation: In the herb layer occur higher coverage values, more (light-indicating) species, but only few species indicating ancient forests and a higher diversity index value. In w-sites, the occurrence of e. g. Calamagrostis epigeios, Impatiens parviflora and Rubus idaeus is supported. reduced. Threatened moss species are rare in w-sites compared to r-sites, since they mainly grow on laying dead wood, which is rare in forests in use, and on inclined/rough-barked stem bases. Ground beetles: The forestry use of lowland beech forests leads to less individuals and lower biomass of so-called mesophilous forest species. Furthermore, the number of flightless individuals is lower. As proper indicators for near-natural lowland beech forests, the three species Carabus glabratus, C. hortensis und Cychrus caraboides could be identified. Indicators: 29 structural indicators were identified and thresholds were given. But even in lowland beech forests managed in a conservation-friendly way, these target values for near-natural and natural forests are unlikely to be reached. Therefore, for w-sites special threshold values have been defined, which consider the inevitable difference between managed and natural forests.

Buchenwälder sind die großflächigste potenziell natürliche Vegetationsform Deutschlands und ein nach EU-FFH-Richtlinie besonders zu schützender Biotoptyp. Eine hohe Naturnähe ist auch in Wirtschaftswäldern (WiWald) notwendig, um die typischen Lebensgemeinschaften naturnaher Wälder langfristig zu erhalten, doch mangelt es an praktikablen/verifizierten Indikatoren, wie die nutzungsbedingte Abweichung vom Naturzustand ermittelt werden kann. In >100 Jahre alten und ~40 ha großen Tiefland-Buchenwäldern (Mecklenburg-Vorpommern/Brandenburg) wurde anhand von 13 WiWäldern, vier seit <20 Jahren (k20) und drei seit >50 Jahren (r50) unbewirtschafteten Beständen den folgenden Fragen nachgegangen: Wie groß sind die strukturellen, vegetationskundlichen und carabidologischen Unterschiede zwischen bewirtschafteten, kurz- und langfristig unbewirtschafteten Buchenwäldern? Gibt es strukturelle Indikatoren und quantitative Größen zur Abschätzung des Einflusses forstlicher Bewirtschaftung auf die Biozönosen von Tiefland-Buchenwäldern? In Probekreisen (Pk) von 500 m² an Rasterpunkten (100 m x 100 m) wurden strukturelle und in Pk von 314 m² vegetationskundliche Daten erhoben. In fünf Pk/Bestand wurde jeweils eine Barberfalle über die Vegetationsperiode installiert. Ganzflächig wurden die Verteilung der Waldentwicklungsphasen (WEP)und zusätzlich zu den Pk-Aufnahmen hektarweise Sonderstrukturen aufgenommen. U. a. wurden folgende Sonderstrukturen aufgenommen: Zunderschwamm, Kronen- und Zwieselbrüche, Ersatzkronen, Blitzrinnen, Risse/Spalten, Höhlen, Mulmkörper/-taschen. Diese naturschutzfachlich wichtigen Sonderstrukturen wurden aus den Habitatansprüchen der typischen Buchenwaldfauna abgeleitet.Es konnten große Unterschiede zwischen WiWald und r50-Flächen (v. a. >100 Jahre unbewirtschafteten Flächen) aufgezeigt werden. Die k20-Flächen unterscheiden sich nicht wesentlich vom WiWald. Die Anzahl verschiedener WEP/ha und WEP-Patches/ha liegt in den r50-Flächen signifikant höher als im WiWald. Der Holzvorrat der r50-Flächen liegt mit ~600 m³/ha (Terminal- ~800 m³/ha, Zerfallsphase 450 m³/ha) deutlich höher als im WiWald. Charakteristisch für die r50-Flächen ist das Vorkommen von in ihrer Vitalität eingeschränkten Bäume ab 80 cm BHD und ein inhomogeneres Lichtmosaik im Bestand. Die Stammqualitäten (u. a. Astigkeit) in r50-Flächen unterscheiden sich kaum von denen in WiWald. In den r50-Flächen kommt bedeutend mehr Totholz (>142 m³/ha) als im WiWald (max. 34 m³/ha) vor. Im WiWald können Stubben dominieren. Verschiedene Totholzqualitäten sind im WiWald nur unvollständig vorhanden. Etwa 40 % des Totholzes besitzt keine Totholznachbarn (r50-Flächen: <2 %) und die Lichtverhältnisse am Totholz sind nicht so vielfältig (wenig sonnenexponiert und wenig gering besonnt). In den >100 Jahre unbewirtschafteten Flächen kommen ~12 Sonderstrukturtypen mit >200 Sonderstrukturen/ha vor. 19 von 20 Sonderstrukturen sind im WiWald signifikant seltener und 11 Sonderstrukturen sind als Naturnähe-Indikatoren geeignet.Vegetation: In der Krautschicht sind höhere Deckungsgrade, mehr (lichtanzeigende) Arten, weniger Waldarten und eine höhere Diversität zu verzeichnen. Im WiWald wird u. a. das Vorkommen von Calamagrostis epigeios, Impatiens parviflora und Rubus idaeus gefördert. Stark gefährdete Moosarten sind im WiWald seltener als in den Referenzwäldern, da sie vor allem auf liegendem Totholz und auf den Stammanläufen vorkommen. Carabiden: Im WiWald gibt es weniger Individuen und Biomasse von mesophilen Waldarten und eine geringere Anzahl von flugunfähigen Individuen. Als Indikatoren für naturnahe Tiefland-Buchenwälder können die drei Arten Carabus glabratus, C. hortensis und Cychrus caraboides bezeichnet werden. Indikatoren: Es wurden Zielgrößen für 29 Struktur-Indikatoren für naturnahe Wälder vorgeschlagen. Für WiWälder wurden gesonderte Zielgrößen festgelegt, die die nutzungsbedingte, nicht zu vermeidende Abweichung vom Naturzustand berücksichtigen. / Beech forests are the most important natural vegetation type of Germany,and they are included in annex II of the EU-FFH-Directive,which requests nature conservation for the listed habitat types.High naturalness is necessary in managed forests (w-sites) to maintain the typical biocoenosis of forests near nature. But there is a lack of practicable/verified indicators to determine the degree of alteration managed forests have compared to natural forests. In >100 year old and ~40 ha big lowland beech forests in Mecklenburg-Vorpommern and Brandenburg, 13 w-sites, 4 study sites which are unmanaged since <20 years (k-sites) and 3 sites which are unmanaged since >50 years (r50-sites) were investigated to answer these questions: What the differences are between w-, k- and r-sites according to forest structure, vegetation and carabids? Are there valid structural indicators with thresholds to assess the impact of forestry use on the biocoenosis of lowland beech forests? At grid points(distance 100 mx 100 m),on circular sample plots (SP) of 500 m² the structural data and on SP of 314 m² the vegetation was investigated. At five SP/study site a pitfall trap was installed during the entire vegetation period. On the whole study site the distribution of forest development phases (FDP) was mapped, and on full one ha plots the special structures were investigated. The following special structures were mapped e.g. Fomes fomentarius trees, crown and crotch breakage, substitute crowns, lightning shakes,gutters/rifts, cavities, mould and bark bag. These special structures have been derived from the habitat needs of the typical beech forest fauna.The results revealed tremendous differences between w- and r50-sites. The k-sites show no clear differences to the managed sites.In the r50-sites, the number of different FDP/ha and FDP units/ ha is significant higher than in w-sites. The timber stock of the r50-sites is ~600 m³/ha (terminal phase ~800 m³/ha, decay phase ~450³/ha). A characteristic feature of the r50-sites is the occurrence of trees with 80 cm bhd or more with reduced vitality. The timber trunk) qualities of r-sites differ only slightly from managed stands. In the r50-sites the dead wood volume (>142 m³/ha) is much higher than in the w-sites (max. 34 m³/ha). Many different features of dead wood occur only fragmentary within w-sites. About 40 % of the dead wood objects have no "dead wood neighbour" (r50-sites: <2 %), and the light distribution is much less diverse. In >100 years unmanaged r-sites ~12 different types of special structures and 200 single special structures occur per ha. 19 out of 20 special structures are significantly less frequent in w-sites; 11 special structures are specifically valuable to be used as naturalness indicators.Vegetation: In the herb layer occur higher coverage values, more (light-indicating) species, but only few species indicating ancient forests and a higher diversity index value. In w-sites, the occurrence of e. g. Calamagrostis epigeios, Impatiens parviflora and Rubus idaeus is supported. reduced. Threatened moss species are rare in w-sites compared to r-sites, since they mainly grow on laying dead wood, which is rare in forests in use, and on inclined/rough-barked stem bases. Ground beetles: The forestry use of lowland beech forests leads to less individuals and lower biomass of so-called mesophilous forest species. Furthermore, the number of flightless individuals is lower. As proper indicators for near-natural lowland beech forests, the three species Carabus glabratus, C. hortensis und Cychrus caraboides could be identified. Indicators: 29 structural indicators were identified and thresholds were given. But even in lowland beech forests managed in a conservation-friendly way, these target values for near-natural and natural forests are unlikely to be reached. Therefore, for w-sites special threshold values have been defined, which consider the inevitable difference between managed and natural forests.