S.H. Luijten

Integrating demographic and genetic approaches in plant conservation

Abstract We summarize the problems that populations of formerly common plants may encounter when habitat fragmentation isolates them and reduces population size. Genetic erosion, inbreeding depression, Allee-effects on reproductive success, catastrophes and environmental stochasticity are illustrated with studies on species that have recently become rare in The Netherlands due to habitat fragmentation. These clearly indicate that population viability is negatively affected. We also show that in the recent literature (since 1980), most studies on the conservation of rare plants have addressed population genetic structure and relationships between genetic variation and population size. Though important, these studies are not suitable for assessing the importance of genetics for population viability. In turn, demographic studies can detect changes in vital rates in small populations, but cannot reveal underlying genetic causes. Fitness and demographic studies are also well-represented in the literature, but remarkably few studies have attempted to integrate empirical demographic and genetic studies. We discuss two approaches to fill this very important lacuna in our knowledge. One of these constructs matrix-projection models on the basis of demographic censuses of—if possible—large and viable populations, and combines these with the results of experiments to determine inbreeding effects on demographic transitions and, subsequently, population growth and extinction. The other approach is to demographically monitor experimentally created small and large populations with low and high genetic variation and measure their actual growth rates and probabilities of extinction. We conclude that demography and demographic-genetic experiments should play a central role in plant conservation genetics.

Future prospects for the rare, late-flowering Gentianella germanica and Gentianopsis ciliata in Dutch nutrient-poor calcareous grasslands

We discuss the population biology of two calcareous grassland gentians, Gentianella germanica and Gentianopsis ciliata, in relation to the habitat management currently practiced in The Netherlands. There, at the margin of their range, both species are rare. Gentianella germanica persists on six remaining locations, whereas Gentianopsis ciliata, with two populations, is nearly extinct. Gentianella germanica is a strict biennial, Gentianopsis ciliata an iteroparous perennial. Both species depend on insects for seed production and su!er from low insect visitation. Pollination experiments in one Dutch population demonstrated that Gentianopsis ciliata is self-compatible, but hardly sets any seed under natural conditions due to pollen limitation. The low reproductive success of both species is partly due to the low pollinator densities at their late flowering time, partly caused by the small population size of the gentians themselves, and partly a result of mowing too early. The latter has destroyed the seed crop of several subsequent years in one population of Gentianopsis ciliata and some of Gentianella germanica. In Gentianella germanica, the early mowing and low insect visitation seems to have resulted in selection of less herkogamous and consequently more autofertile individuals. The perspectives for Gentianopsis ciliata are currently extremely poor in The Netherlands. Under the present circumstances, extinction will most likely occur within 10‐20 years. Population reinforcement (seeding, artificial cross-pollination with nearby populations) should be considered if we want to conserve this species. For both gentians, but also of other flagship species of nutrient-poor calcareous grasslands, the total grassland area needs to be enlarged and must constitute an interconnected network of reserves. The traditional management method, sheep grazing, is to be preferred over mowing, but only if the flock visits each grassland patch or reserve for only a short time of less than a day, and with intervals of more than 2 weeks between visits. If mowing is preferred for other reasons, it should be done rotationally, and not before October. # 2002 Elsevier Science Ltd. All rights reserved.

Variable herkogamy and autofertility in marginal populations ofGentianella germanica in the Netherlands

Gentianella germanica(Willd.) Börner, a rare plant species, occurs at the margin of its distribution area in the Netherlands. It is found only in isolated patches of chalk grassland surrounded by intensively-used agricultural land. In 1997 and 1998, we investigated the spatial and temporal separation of anthers and stigma in relation to autofertility. We performed our experiments in one large and two small populations to investigate the possible selection against herkogamy and for increased autofertility in marginal populations with pollination limitation. Dichogamy appeared to be very weak and varied within individuals. Homogamous and protandrous flowers were most frequent. Dichogamy is apparently no effective barrier against self-fertilization in this species. Herkogamy varied significantly between individuals and between populations, and plant and flower size were not correlated with anther-stigma separation. Both observations suggest a genetic basis for herkogamy. Autofertility was generally high in all three populations. Only in one small population was a correlation between autofertility and herkogamy found. In this population, most plants had the stigma positioned above the anthers (mean herkogamy +0.8 mm), while in the other two populations the stigma was positioned mainly in between or below the anthers (mean herkogamy −0.55 mm). In comparison with earlier data from 1991 and 1992, plants in the large population were less herkogamous and more self-fertile in 1998. Since 1993 this population was mown each year one month earlier than usual, leading to dramatic reductions (bottlenecks) in population size. In addition, the early mowing time may have caused a further reduction of the already scarce visitation by pollinating insects. We hypothesize that both adverse conditions may have caused a selection of plants with smaller anther-stigma separation and higher autofertility and thus showing that human influence can have profound effects on the reproductive success and evolution of rare plant species, even if it occurs in the context of nature conservation.

Matrix population models from 20 studies of perennial plant populations

Demographic transition matrices are among the most commonly applied population models for both basic and applied ecological research. The relatively simple framework of these models and simple, easily interpretable summary statistics they produce have prompted the wide use of these models across an exceptionally broad range of taxa. Here, we provide annual transition matrices and observed stage structures/population sizes for 20 perennial plant species which have been the focal species for long-term demographic monitoring. These data were assembled as part of the “Testing Matrix Models” working group through the National Center for Ecological Analysis and Synthesis (NCEAS). The data represent 82 populations with >460 total population-years of observations. It is our hope that making these data available will help promote and improve our ability to monitor and understand plant population dynamics. The complete data sets corresponding to abstracts published in the Data Papers section of the journal are publ...

Outbreeding depression and breeding system evolution in small, remnant populations of Primula vulgaris: consequences for genetic rescue

Many species suffer from anthropogenic habitat fragmentation. The resulting small and isolated populations are more prone to extinction due to, amongst others, genetic erosion, inbreeding depression and Allee-effects. Genetic rescue can help mitigate such problems, but might result in outbreeding depression. We evaluated offspring fitness after selfing and outcrossing within and among three very small and isolated remnant populations of the heterostylous plant Primula vulgaris. We used greenhouse-grown offspring from these populations to test several fitness components. One population was fixed for the pin-morph, and was outcrossed with another population in the field to obtain seeds. Genetic diversity of parent and offspring populations was studied using microsatellites. Morph and population-specific heterosis, inbreeding and outbreeding depression were observed for fruit and seed set, seed weight and cumulative fitness. Highest fitness was observed in the field-outcrossed F1-population, which also showed outbreeding depression following subsequent between-population (back)crossing. Despite outbreeding depression, fitness was still relatively high. Inbreeding coefficients indicated that the offspring were more inbred than their parent populations. Offspring heterozygosity and inbreeding coefficients correlated with observed fitness. One population is evolving homostyly, showing a thrum morph with an elongated style and high autonomous fruit and seed set. This has important implications for conservation strategies such as genetic rescue, as the mating system will be altered by the introduction of homostyles.