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Besides habitat loss, population-biological and genetic consequences of habitat fragmentation are thought to be a major threat to species since the 1990's and thus are now in the focus of plant species conservation. Using examples, this article gives an overview on the state of the art. It aims to evaluate the relevance habitat fragmentation and the resulting small size and isolation of populations may have for Central European plant populations. Stochasticity, edge effects, pollinator limitation, genetic drift and inbreeding depression are identified as important and very widespread negative effects. Together with changed habitat quality due to eutrophication, drainage or altered land use they negatively affect the fitness of individuals and populations, resulting in an increased risk of extinction. This negative effect of small populations on the fitness of individuals is called the Allee-effect, irrespective of the underlying causes, which can only be identified by scientific experiments. Metapopulation dynamics that are supported by a habitat network may prevent a permanent extinction of plant populations and minimize the negative genetic effects of habitat fragmentation by increasing gene flow via pollen and seeds. However, existing studies from Central Europe mainly concentrated on certain plant families (Gentianaceae, Primulaceae), habitats (species- rich grasslands), insect-pollinated and outcrossing species, and species mainly relying on sexual reproduction. On the other hand, few insights exist about grasses, ruderal plants and weeds, non-indigenous, wind- and self-pollinated species, and species mainly reproducing vegetatively or via apomictic seeds. However, according to the present state of knowledge especially these plant species, and those with a high dispersal potential, have to be considered as less sensitive to habitat fragmentation. Based on these findings, habitat types are classified with regard to their sensitivity to fragmentation, and ecological characters and species traits of sensitive and less sensitive species are compared. Finally, general consequences for conservation practice are presented with regard to target species and habitats for the formation of habitat networks, minimum viable population sizes, genetic rescue of populations, and deploying plants from ex-situ conservation to natural habitats.
Aim To investigate the effect of temperature, latitude and local environment on the reproductive traits of widespread perennial forest herbs to better understand the potential impacts of rising temperatures on their population dynamics and colonization capacities. Location Six regions along a latitudinal gradient from France to Sweden. Methods Within each region, we collected data from three to five populations of up to six species. For each species, several variables were recorded in each region (temperature, latitude) and population (local abiotic and biotic environmental variables), and seed production and germination were estimated. Resource investment in reproduction (RIR) was quantified as seed number ¥ seed mass, while germinable seed output (GSO) was expressed as seed number ¥ germination percentage.We performed linear regression and mixed effect models to investigate the effects of temperature (growing degree hours), latitude and local abiotic and biotic environment on RIR and GSO. Results Temperature and latitude explained most of the variation in RIR and GSO for early flowering species with a northerly distribution range edge (Anemone nemorosa, Paris quadrifolia and Oxalis acetosella). Reproduction of the more southerly distributed species (Brachypodium sylvaticum, Circaea lutetiana and Primula elatior), in contrast, was independent of temperature/latitude. In the late summer species, B. sylvaticum and C. lutetiana, variation in RIR and GSO was best explained by local environmental variables, while none of the investigated variables appeared to be related to reproduction in P. elatior. Main conclusions We showed that reproduction of only two early flowering, northerly distributed species was related to temperature. This suggests that the potential reproductive response of forest herbs to climate warming partly depends on their phenology and distribution, but also that the response is to some extent species dependent. These findings should be taken into account when predictions about future shifts in distribution range are made.
Predictions on displacement of suitable habitats due to climate change suggest that plant species with poor colonization ability may be unable to move fast enough to match forecasted climate-induced changes in habitat distribution. However, studies on early Holocene plant migration show fast migration of many plant species that are poor colonizers today. We hypothesize that warmer temperatures during the early Holocene yielded higher seed quality, contributing to explaining the fast migration. We studied how the 3 seed quality variables, seed mass, germinability, and requirements for break of seed dormancy, vary for seeds of 11 forest herb species with varying colonization capacity collected along a 1400-km latitudinal gradient. Within species, seed mass showed a positive correlation with latitude, whereas germinability was more positively correlated with temperature (growing degree hours obtained at time of seed collection). Only slow-colonizing species increased germinability with temperature, whereas only fast-colonizing species increased germinability with latitude. These interactions were only detectable when analyzing germinability of the seeds, even though this trait and seed mass were correlated. The requirement for dormancy break did not correlate with latitude or temperature. The results indicate that seed development of slow colonizers may be favoured by a warmer climate, which in turn may be important for their migration capacity.
Questions: 1. Are there differences among species in their preference for coniferous vs. deciduous forest? 2. Are tree and shrub species better colonizers of recent forest stands than herbaceous species? 3. Do colonization patterns of plant species groups depend on tree species composition? Location: Three deciduous and one coniferous recent forest areas in Brandenburg, NE Germany. Methods: In 34 and 21 transects in coniferous and deciduous stands, respectively, we studied the occurrence and percentage cover of vascular plants in a total of 150 plots in ancient stands, 315 in recent stands and 55 at the ecotone. Habitat preference, diaspore weight, generative dispersal potential and clonal extension were used to explain mechanisms of local migration. Regression analysis was conducted to test whether migration distance was related to species’ life-history traits. Results: 25 species were significantly associated with ancient stands and ten species were significantly more frequent in recent stands. Tree and shrub species were good colonizers of recent coniferous and deciduous stands. In the coniferous stands, all herbaceous species showed a strong dispersal limitation during colonization, whereas in the deciduous stands generalist species may have survived in the grasslands which were present prior to afforestation. Conclusions: The fast colonization of recent stands by trees and shrubs can be explained by their effective dispersal via wind and animals. This, and the comparably efficient migration of herbaceous forest specialists into recent coniferous stands, implies that the conversion of coniferous into deciduous stands adjacent to ancient deciduous forests is promising even without planting of trees.
Natürliche Standorte der Waldkiefer gibt es in Deutschland nur kleinflächig. Während Kiefernforste anstelle natürlicher Laubwälder heute oft landschaftsprägend sind, bildet die konkurrenzschwache und lichtbedürftige Kiefer ausschließlich auf extrem trockenen oder nassen, nährstoffarmen Standorten naturnahe Schlusswaldgesellschaften. Regionale Schwerpunkte liegen in subkontinentalen Regionen wie dem nordostdeutschen Tiefland und Bayern, ein „natürliches Kiefernareal" lässt sich aber kaum abgrenzen. An der Trockengrenze des Waldes finden sich auf Kalk- und Dolomitgesteinen artenreiche Karbonat-Trockenkiefernwälder mit Elementen der alpinen Rasen und Kalkmagerrasen in der Bodenvegetation. Diese Wälder besiedeln steile, südexponierte Felsen und morphodynamisch aktive Bereiche wie Rutschhänge und FlussSchotterböden im Umkreis der Alpen, kommen aber auch in den Mittelgebirgen vor. Ihr Gegenstück auf sauren Standorten sind die Sand- und Silikat-Kiefernwälder der Quarzsande und Sandstein-Verwitterungsböden, deren Bodenvegetation durch Zwergsträucher, Moose und Strauchflechten geprägt ist. Hier siedelt die Kiefer in den Tieflagen besonders auf Binnendünen und Sandern, aber auch auf Küstendünen der Ostsee, in den Mittelgebirgen z. B. auf den Sandsteinriffen der Sächsischen Schweiz. Der dritte Wuchsbereich natürlicher Kiefernwälder sind saure, nährstoffarme Moore, die ganz überwiegend von Regenwasser gespeist werden. Auch die Kiefern-Moorwälder sind in Nordostdeutschland und Bayern am häufigsten. Von diesen Standorten ausgehend, wo ihr Platz kaum von anderen Baumarten streitig gemacht wird, tritt die Waldkiefer immer wieder als Pionier auf weniger extremen Standorten auf. In der Naturlandschaft kam dies etwa nach Waldbränden oder Stürmen vor, doch der Mensch förderte die Kiefer durch Auflichtung der Wälder, Waldweide und Streunutzung stark. Auch die damit verbundene Nährstoffverarmung macht eine exakte Abgrenzung natürlicher Kiefernstandorte unmöglich. Die schlechtwüchsigen und forstwirtschaftlich nicht interessanten, ästhetisch aber sehr ansprechenden natürlichen Kiefernbestände sind heute vor allem durch Stickstoff-Immissionen gefährdet. Trotz ihrer oft kargen Erscheinung besitzen sie einen hohen Wert für die Biodiversität und den Artenschutz. Neben bodenbewohnenden Flechten und regionalen Relikt-Endemiten ist vor allem die in den letzten Jahrzehnten zunehmend gefährdete Vielfalt an Mykorrhiza-Pilzen hervorzuheben, die der Kiefer das Leben auf extrem nährstoffarmen Standorten überhaupt ermöglichen. Abschließend werden mögliche Schutz- bzw. Regenerationsmaßnahmen wie das Abplaggen flechtenreicher Kiefernstandorte vorgestellt.
Vegetation und Standort bodensaurer Buchenwälder am Arealrand : am Beispiel Mittelbrandenburgs
(2008)
Different from NW Germany, the northern part of NE Germany and the "Hohe Flaeming" region, central Brandenburg is considered as being largely devoid of natural beech forests because of its subcontinental, dry climate. In the present study the vegetation ecology of beech forests of the region is comprehensively documented for the first time, and they are compared with NW German stands in Lower Saxony. In the study area beech forests are concentrated in the Berlin-Potsdam region along the Havel river lakes which is characterised by relatively high precipitation and a specific land use history. All belong to the Luzulo-Fagetum growing on acid soils. Four subtypes are distinguished according to nutrient availability and soil moisture. The central Brandenburg Luzulo-Fagetum does not markedly deviate from other beech forests in the northern German lowlands with respect to vegetation structure and edaphic subtypes. However, numerous indicator species for humid or moist conditions are less frequent than under atlantic climate conditions in the lowlands of Lower Saxony, a pattern occurring also in other forest communities. On the other hand, nitrogen and disturbance indicators are more frequent in central Brandenburg. As expected, podzolisation of the soils and humus accumulation is lower in beech forests under subcontinental climate, but surprisingly the soils are more sandy and thus drier. However, beech forests are lacking on south-exposed slopes, and they are notably occurring in northern exposition. A combined analysis of distribution patterns and climatic data, postglacial vegetation history and forest use history, and actual rejuvenation dynamics reveals that the present-day beech forests in central Brandenburg have to be considered as near-natural relics, which are currently spreading. The range of potentially natural beech forests is larger than assumed until now, but further on it is not clearly to define.
Neben dem Habitatverlust gelten Konsequenzen der Habitatfragmentierung seit den 1990er Jahren als wesentliche Ursache der Gefaehrdung von Pflanzen und stehen damit nun auch im Fokus des botanischen Artenschutzes. Der vorliegende Beitrag gibt einen ueberblick ueber den Stand der populationsbiologischen und genetischen Forschung und versucht abzuschaetzen, welche Bedeutung Habitatfragmentierung und die dadurch entstehenden kleinen, isolierten Populationen auf heimische Pflanzenarten haben koennen. Als wesentliche und offenbar sehr weit verbreitete negative Effekte werden Zufallsereignisse, Randeffekte, Bestaeuberlimitierung, Gendrift und Inzuchtdepression identifiziert. Zusammen mit verringerter Habitatqualitaet durch Eutrophierung, Entwaesserung oder Nutzungsaenderung wirken sie zumeist negativ auf die Fitness der Individuen und Populationen und erhoehen so deren Aussterberisiko. Dieser negative Effekt kleiner Populationen auf die individuelle Fitness wird unabhaengig von der Ursache als Allee-Effekt bezeichnet. Eine durch einen Biotopverbund gefoerderte Metapopulationsdynamik kann das dauerhafte Aussterben von Pflanzenpopulationen verhindern und mindert die negativen genetischen Effekte der Habitatfragmentierung ueber einen erhoehten Genfluss durch Pollen und Samen. Die bisherigen wissenschaftlichen Studien in Mitteleuropa beruhen allerdings in ueberproportionaler Weise auf bestimmten Pflanzenfamilien (Gentianaceae, Primulaceae), Habitaten (Trocken- und Magerrasen, Wirtschaftsgruenland), insekten- und obligat fremdbestaeubten sowie weitgehend auf sexuelle Fortpflanzung angewiesenen Arten, waehrend etwa ueber Grasartige, Ruderalpflanzen, wind- und selbstbestaeubte sowie an vegetative Fortpflanzung angepasste Arten nur wenige Erkenntnisse vorliegen. Gerade diese und Pflanzenarten mit hohem Ausbreitungspotenzial muessen aber nach derzeitigem Wissensstand als weniger sensitiv gegenueber Habitatfragmentierung eingestuft werden. Auf diesen Befunden aufbauend werden fuer die Naturschutzpraxis Biotoptypen hinsichtlich ihrer Sensitivitaet gegenueber Habitatfragmentierung klassifiziert und ein auf biologischen Merkmalen basierender Kriterienkatalog zur Auswahl von Zielarten des Biotopverbunds vorgestellt. Schließlich wird eroertert, was bei Maßnahmen zur Regeneration kleiner bzw. bereits ausgestorbener Populationen zu beachten ist, und es werden allgemeine Folgerungen zur Ausgestaltung eines Biotopverbundskonzepts fuer Pflanzen gezogen.
Sand- und Silikat-Kiefernwälder (Dicrano-Pinion) in Deutschland : Gliederungskonzept und Ökologie
(2007)
In preparation for the „Synopsis of plant communities of Germany“ a comprehensive classification concept for the Scots pine forests on sandy and silicate soils is presented. On the basis of 2699 relevés from all natural provinces with important occurrences this classification for the first time integrates both northern and southern German forest stands. Pine forests are stable (“climax”) communities on three distinct habitat types at the drought and wetness limits of forest growth. In the phytosociological system these are reflected by the clearly separated syntaxa Erico-Pinetea (dry-calcareous), Dicrano-Pinion (dry-acidic) and Vaccinio uliginosi- Pinetea (wet-acidic). However, Pulsatillo-Pinetea (dry-moderate basicity) described in earlier publications cannot be separated floristically. In addition to the stable communities on extreme habitats pine forests of the mentioned syntaxa are widespread on potential mixed deciduous forest stands, especially after anthropogenic devastation and even beyond their original range. Six communites of the Dicrano-Pinion which also includes such secondary pine forest stands are occurring in Germany. They are presented in detail and classified according to their dynamic and edaphic differentiation. Lichen-rich pine forests (Cladonio- Pinetum) which grow on extremely dry and nutrient-poor sites are ecologically and floristically well-defined, though closely connected with other Dicrano-Pinion communities by forest succession. After separation of the Cladonio-Pinetum the Leucobryo-Pinetum is a speciespoor “central association” within the alliance. The Deschampsia flexuosa-Pinus-sylvestriscommunity is the most widespread forest type and dynamically and floristically passes into the mixed oak forests on acidic soils (Quercion roboris). On base-rich habitats the Empetro- Pinetum as endemic community of the southern Baltic Sea coasts, and the Peucedano-Pinetum in the northeastern and southern German inland are distinguished. The latter is found both on calcareous sands and primarily acidic sands which are secondary limed by calciferous pollutions. Finally, differences and similarities between the geographically separated northern and southern German Dicrano-Pinion forests are discussed in a biogeographic context, emphasising the advantages of the presented nation-wide classification concept.