Summary 1.The relationship between large herbivore numbers and landscape cover over time is poorly understood. There are two schools of thought: one views large herbivores as relatively passive elements upon the landscape and the other as ecosystem engineers driving vegetation succession. The latter relationship has been used as an argument to support reintroductions of large herbivores onto many landscapes in order to increase vegetation heterogeneity and biodiversity through local-scale disturbance regimes. Most of the research examining the relationship between large herbivores and their impact on landscapes has used extant studies. An alternative approach is to estimate the impact of variations in herbivore populations through time using fossil dung fungal spores and pollen in sedimentary sequences. However, to date, there has been little quantification of fossil dung fungal spore records and their relationship to herbivore numbers, leaving this method open to varied interpretations. 2. In this study, we developed further the dung fungal spore method and determined the relationship between spore abundance in sediments (number cm À2 year À1) and herbivore biomass densities (kg ha À1). To establish this relationship, we used the following: (i) the abundance of Sporormiella spp., Sordaria spp. and Podospora spp. spores in modern sediments from ponds and (ii) weekly counts of contemporary wildlife over a period of 5 years from the rewilded site, Oostvaardersplassen, in the Netherlands. 3.Results from this study demonstrate that there is a highly significant relationship between spore abundance and local biomass densities of herbivores that can be used in the calibration of fossil records. Mammal biomass density (comprising Konik horses, Heck cattle and red deer) predicts in a highly significant way the abundance of all dung fungal spores amalgamated together. This relationship is apparent at a very local scale (<10 m), when the characteristics of the sampled ponds are taken into account (surface area of pond, length of shoreline). In addition, we identify that dung fungal spores are principally transported into ponds by surface run-off from the shores. 4. These results indicate that this method provides a robust quantitative measure of herbivore population size over time.

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In a progeny of a selfed individual of the dark red-flowered cultivar 'Roter Vogel' some white-flowered plants appeared as the result of a mutation of the genetic factor Anl involved in anthocyanin synthesis. The white flowers of these plants had red spots owing to back-mutations in the dermal cells of the young corolla.Owing to a striking unstability of the new allele of Anl, a number of mutants originated which differ mutually in the frequency of reversion, which expressed itself in the very substantial differences in the spot density of the limb of the corolla. Between a mean number of less than one spot per cm(2) of the limb and a mean number of over 10.000 spots/cm(2), a series of transitions was found.The reversions did not remain restricted to the young epidermis but also occurred in sporogenous tissues. This resulted in the appearance of selfcoloured red descendants of plants with red-spotted white flowers. There is a positive correlation between the spot density of the parent plants and the percentage of plants with completely red corollas.The red spots on the corolla usually have the same colour as the wild type ('Roter Vogel'), but occasionally mutants occur with paler spots, the colour varying from a very pale pink to a red nearly as deep as in the wild type. The selfcoloured descendants of such mutants also show this colour variation from pale pink to red.On the grounds of these observations a theory was formulated which postulates that the Anl locus consists of a structural gene responsible for an enzyme active during anthocyanin synthesis and a regulatory element built up from intermediate repetitive DNA. This regulatory element in turn is built up of two components, one of which, the 'mutator', decides the activation of the structural gene while the other, the 'expressor', modifies the rate of activation. The mutations must be considered representative of larger or smaller deletions within one or both of these components. Reversions are the result of the restoration of the deletions by means of an amplification of the repetitive DNA in dividing cells of the developing flower buds.

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