Johan Ehrlén

Seed and microsite limitation of recruitment in plant populations

SummaryAvailability of seed and microsites, respectively, are two factors that potentially may limit recruitment in plant populations. Microsites are small-scale sites suitable for germination and survival of seedlings. We discuss this dichotomy of recruitment limitation both from a theoretical and empirical point of view. Investigations of recruitment in 14 woodland species showed that 3 species were seed limited, 6 species were limited by a combination of seed and microsite availability, and 5 species were found not to be seed limited, but the limiting factor was not identified. A “combination of seed and microsite limitation” implies that recruitment is promoted by increasing both seed and microsite availability. We suggest that the importance of seed limitation in plant populations has been underestimated, and that the operating limiting factors may be dependent on spatial and temporal scale. We expect that many species, if adequately studied, will turn out to be both seed and microsite limited. Experimental field studies that incorporate a range of seed and microsite “densities” in various spatial and temporal scales are needed to examine the extent to which plant populations are seed and microsite limited.

ELASTICITIES: A REVIEW OF METHODS AND MODEL LIMITATIONS

Elasticity is a perturbation measure in matrix projection models that quantifies the proportional change in population growth rate as a function of a proportional change in a demographic transition (growth, survival, reproduction, etc.). Elasticities thus indicate the relative “importance” of life cycle transitions for population growth and maintenance. In this paper, we discuss the applications of elasticity analysis, and its extension, loop analysis, in life history studies and conservation. Elasticity can be interpreted as the relative contribution of a demographic parameter to population growth rate. Loop analysis reveals the underlying pathway structure of the life cycle graph. The different kinds of results of the two analyses in studies of life histories are emphasized. Because elasticities quantify the relative importance of life cycle transitions to population growth rate, it is generally inferred that management should focus on the transitions with the largest elasticities. Such predictions based on elasticities seem robust, but we do identify three situations where problems may arise. The mathematical properties and biological constraints that underlie these pitfalls are explained. Examples illustrate the additional information that needs to be taken into account for a sensible use of elasticities in population management. We conclude by indicating topics that are in need of research.

DISPERSAL LIMITATION AND PATCH OCCUPANCY IN FOREST HERBS

The distribution of species depends on the availability of suitable habitats, the capacity to disperse to these habitats, and the capacity of populations to persist after establishment. Dispersal limitation implies that not all suitable habitat patches will be occupied by a species. However, the extent to which dispersal limits local distribution is poorly known. In this study, we transplanted seeds, bulbils, and juvenile plants to examine patterns of dispersal limitation and patch occupancy in seven temperate-forest herbs. Recruitment was recorded during four years in 48 patches. The investigated species varied considerably in their natural abundance in the patches. Patterns of seedling emergence and establishment among patches were not related to any of nine investigated abiotic factors. In contrast, the availability of seeds or bulbils was found to limit recruitment in six of the investigated species. Establishment was also successful in many patches where the species did not occur naturally. Estimated patch occupancy in the investigated species ranged from 17.2% to 94.6%. Seed size was positively correlated with the probability of successful establishment of seeds and negatively correlated with patch occupancy. The results suggest that dispersal limitation is an important structuring factor in temperate-forest herb communities. The distribution of species can be perceived as the result of processes operating both among and within patches. Seed size is a key trait in these processes.

Diversity of ageing across the tree of life

Evolution drives, and is driven by, demography. A genotype moulds its phenotype’s age patterns of mortality and fertility in an environment; these two patterns in turn determine the genotype’s fitness in that environment. Hence, to understand the evolution of ageing, age patterns of mortality and reproduction need to be compared for species across the tree of life. However, few studies have done so and only for a limited range of taxa. Here we contrast standardized patterns over age for 11 mammals, 12 other vertebrates, 10 invertebrates, 12 vascular plants and a green alga. Although it has been predicted that evolution should inevitably lead to increasing mortality and declining fertility with age after maturity, there is great variation among these species, including increasing, constant, decreasing, humped and bowed trajectories for both long- and short-lived species. This diversity challenges theoreticians to develop broader perspectives on the evolution of ageing and empiricists to study the demography of more species.

How do plant ecologists use matrix population models

Matrix projection models are among the most widely used tools in plant ecology. However, the way in which plant ecologists use and interpret these models differs from the way in which they are presented in the broader academic literature. In contrast to calls from earlier reviews, most studies of plant populations are based on < 5 matrices and present simple metrics such as deterministic population growth rates. However, plant ecologists also cautioned against literal interpretation of model predictions. Although academic studies have emphasized testing quantitative model predictions, such forecasts are not the way in which plant ecologists find matrix models to be most useful. Improving forecasting ability would necessitate increased model complexity and longer studies. Therefore, in addition to longer term studies with better links to environmental drivers, priorities for research include critically evaluating relative/comparative uses of matrix models and asking how we can use many short-term studies to understand long-term population dynamics.

COLONIZATION–EXTINCTION DYNAMICS OF AN EPIPHYTE METAPOPULATION IN A DYNAMIC LANDSCAPE

Metapopulation dynamics have received much attention in population bi- ology and conservation. Most studies have dealt with species whose population turnover rate is much higher than the rate of patch turnover. Models of the dynamics in such systems have assumed a static patch landscape. The dynamics of many species, however, are likely to be significantly affected by the dynamics of their patches. We tested the relative im- portance of local conditions, connectivity, and dynamics of host tree patches on the meta- population dynamics of a red-listed epiphytic moss, Neckera pennata, in Sweden. Repeated surveys of the species and its host trees were conducted at three sites over a period of six years. There was a positive effect of connectivity, and colonizations mainly occurred in the vicinity of occupied trees. Colonizations were also less likely on strongly leaning trees. Local extinctions sometimes occurred from small trees with low local abundances but were most often caused by treefall. Simulations of the future (100 years) dynamics of the system showed that the metapopulation size will be overestimated unless the increased local ex- tinction rate imposed by the dynamics of the trees is accounted for. The simulations also suggested that local extinctions from standing trees may be disregarded in dynamic models for this species. This implies that the dynamics of N. pennata can be characterized as a patch-tracking metapopulation, where local extinctions are caused by patch destruction.

Reproductive effort and herbivory timing in a perennial herb: Fitness components at the individual and population levels

We experimentally investigated how pollinator- and herbivore-induced changes influence the performance of the long-lived herb Primula veris. Eight treatments that corresponded to natural factors normally affecting this species were designed to enhance or reduce reproductive success and resource availability (flower removal, supplementary pollination, defoliation). During the experimental season and in the following year we quantified responses in terms of survival, growth, and seed production of reproductive plants. Matrix population models were used to calculate population growth rate using the demographic parameters recorded in permanent plots and respective treatment groups. Seed production was not limited by pollen availability, and we found no evidence of a cost of reproduction. Leaf removal had either no effect or a negative effect on future performance, depending on the timing of removal. Defoliation early in the season reduced current seed production and future growth, whereas removal during fruit development affected performance in the following year. Demographic models suggest that leaf damage has a smaller negative impact than flower removal on overall performance in this population. Our results suggest that the source-sink paths vary over the season and that the timing of herbivory may influence the extent to which effects are carried over to subsequent reproductive seasons.

Causes and consequences of variation in plant population growth rate: a synthesis of matrix population models in a phylogenetic context

Explaining variation in population growth rates is fundamental to predicting population dynamics and population responses to environmental change. In this study, we used matrix population models, which link birth, growth and survival to population growth rate, to examine how and why population growth rates vary within and among 50 terrestrial plant species. Population growth rates were more similar within species than among species; with phylogeny having a minimal influence on among-species variation. Most population growth rates decreased over the observation period and were negatively autocorrelated between years; that is, higher than average population growth rates tended to be followed by lower than average population growth rates. Population growth rates varied more through time than space; this temporal variation was due mostly to variation in post-seedling survival and for a subset of species was partly explained by response to environmental factors, such as fire and herbivory. Stochastic population growth rates departed from mean matrix population growth rate for temporally autocorrelated environments. Our findings indicate that demographic data and models of closely related plant species cannot necessarily be used to make recommendations for conservation or control, and that post-seedling survival and the sequence of environmental conditions are critical for determining plant population growth rate.

The mechanisms causing extinction debts

Extinction debts can result from many types of habitat changes involving mechanisms other than metapopulation processes. This is a fact that most recent literature on extinction debts pays little attention to. We argue that extinction debts can arise because (i) individuals survive in resistant life-cycle stages long after habitat quality change, (ii) stochastic extinctions of populations that have become small are not immediate, and (iii) metapopulations survive long after that connectivity has decreased if colonization-extinction dynamics is slow. A failure to distinguish between these different mechanisms and to simultaneously consider both the size of the extinction debt and the relaxation time hampers our understanding of how extinction debts arise and our ability to prevent ultimate extinctions.

Proximate Limits to Seed Production in a Herbaceous Perennial Legume, Lathyrus Vernus

Resources, pollen, and predation limit fruit and seed set in Lathyrus vernus. Removal of flowers increased fruit and seed set among remaining ovaries and ovules, but did not affect seed mass. The distributions of the increased fruit and seed set among remaining ovaries indicated specific and variable patterns of translocation. The importance of pollen relative to resource deficiency varied between two experi- mental seasons, among individuals, and among flowers within shoots. Pollen supplemen- tation increased fruit and seed set in one of two experimental seasons. When shoots were supplied with extra pollen a greater proportion produced fruits, but the number of fruits per fruit-producing shoot did not increase. Differences in fruit set among flowers with different phonology and different position within inflorescences persisted also among pollen- supplemented flowers. Furthermore, while pollen availability sometimes may limit fruit initiation, maturation seems to be limited primarily by resources. Genets that received supplementary pollen in 1988 produced fewer flowers in 1989, indicating that long-term fecundity is not necessarily pollen limited. Lastly, exclusion of the main ovary predators increased fruit set.