Phenotypic plasticity of native vs. invasive purple loosestrife: a two-state multivariate approach

The differences in phenotypic plasticity between invasive (North American) and native (German) provenances of the invasive plant Lythrum salicaria (purple loosestrife) were examined using a multivariate reaction norm approach testing two important attributes of reaction norms described by multivariate vectors of phenotypic change: the magnitude and direction of mean trait differences between environments. Data were collected for six life history traits from native and invasive plants using a split-plot design with experimentally manipulated water and nutrient levels. We found significant differences between native and invasive plants in multivariate phenotypic plasticity for comparisons between low and high water treatments within low nutrient levels, between low and high nutrient levels within high water treatments, and for comparisons that included both a water and nutrient level change. The significant genotype x environment (G x E) effects support the argument that invasiveness of purple loosestrife is closely associated with the interaction of high levels of soil nutrient and flooding water regime. Our results indicate that native and invasive plants take different strategies for growth and reproduction; native plants flowered earlier and allocated more to flower production, while invasive plants exhibited an extended period of vegetative growth before flowering to increase height and allocation to clonal reproduction, which may contribute to increased fitness and invasiveness in subsequent years.     

Importance of individual and environmental variation for invasive species spread: a spatial integral projection model

Plant survival, growth, and flowering are size dependent in many plant populations but also vary among individuals of the same size. This individual variation, along with variation in dispersal caused by differences in, e.g., seed release height, seed characteristics, and wind speed, is a key determinant of the spread rate of species through homogeneous landscapes. Here we develop spatial integral projection models (SIPMs) that include both demography and dispersal with continuous state variables. The advantage of this novel approach over discrete-stage spread models is that the effect of variation in plant size and size-dependent vital rates can be studied at much higher resolution. Comparing Neubert-Caswell matrix models to SIPMs allowed us to assess the importance of including individual variation in the models. As a test case we parameterized a SIPM with previously published data on the invasive monocarpic thistle Carduus nutans in New Zealand. Spread rate (c*) estimates were 34% lower than for standard spatial matrix models and stabilized with as few as seven evenly distributed size classes. The SIPM allowed us to calculate spread rate elasticities over the range of plant sizes, showing the size range of seedlings that contributed most to c* through their survival, growth and reproduction. The annual transitions of these seedlings were also the most important ones for local population growth (lambda). However, seedlings that reproduced within a year contributed relatively more to c* than to lambda. In contrast, plants that grow over several years to reach a large size and produce many more seeds, contributed relatively more to lambda than to c*. We show that matrix models pick up some of these details, while other details disappear within wide size classes. Our results show that SIPMs integrate various sources of variation much better than discrete-stage matrix models. Simpler, heuristic models, however, remain very valuable in studies where the main goal is to investigate the general impact of a life history stage on population dynamics. We conclude with a discussion of future extensions of SIPMs, including incorporation of continuous time and environmental drivers.     

Scale dependence of reproductive failure in fragmented Echinacea populations

I investigated reproduction in a three-year study of Echinacea angustifolia, purple coneflower, growing in a fragmented prairie landscape. I quantified the local abundance of flowering conspecifics at individual-based spatial scales and at a population-based spatial scale. Regression analyses revealed that pollen limitation increased while seed set and fecundity decreased with isolation of individual plants. Isolation, defined as the distance to the k(th) nearest flowering conspecific, was a good predictor of pollen limitation, for all nearest neighbors considered (k = 1-33), but the strength of the relationship, as quantified by R2, peaked at intermediate scales (k = 2-18). The relationship of isolation to seed set and fecundity was similarly strongest at intermediate scales (k = 3-4). The scale dependence of individual density effects on reproduction (density of flowering plants within x meters) resembled that of isolation. Analyses at a population-based scale showed that pollen limitation declined significantly with population size. Seed set and fecundity also declined with population size, but significantly so only in 1998. Whether quantifying local abundance with population- or individual-based measures, reproductive failure due to pollen limitation is a consistent consequence of Echinacea scarcity. However, individual-based measures of local abundance predicted pollen limitation from a wider sample of plants with a simpler model than did population size. Specifically, the largest site, a nature preserve, is composed of plants with intermediate individual isolation and, as predicted, intermediate pollen limitation, but its large population size poorly predicted population mean pollen limitation.     

Allocation in reproduction is not tailored to the probable number of matings in common toad (<Emphasis Type="Italic">Bufo bufo</Emphasis>) males

The theory of life history evolution assumes trade-offs between competing fitness traits such as reproduction, somatic growth, and maintenance. One prediction of this theory is that if large individuals have a higher reproductive success, small/young individuals should invest less in reproduction and allocate more resources in growth than large/old individuals. We tested this prediction using the common toad (Bufo bufo), a species where mating success of males is positively related to their body size. We measured testes mass, soma mass, and sperm stock size in males of varying sizes that were either (1) re-hibernated at the start of the breeding season, (2) kept without females throughout the breeding season, or (3) repeatedly provided with gravid females. In the latter group, we also estimated fertilization success and readiness to re-mate. Contrary to our predictions, the relationship between testes mass and soma mass was isometric, sperm stock size relative to testes mass was unrelated to male size, fertilization success was not higher in matings with larger males, and smaller males were not less likely to engage in repeated matings than larger males. These results consistently suggest that smaller males did not invest less in reproduction to be able to allocate more in growth than larger males. Causes for this unexpected result may include relatively low year-to-year survival, unpredictable between-year variation in the strength of sexual selection and low return rates of lowered reproductive investment.     

Bridging the generation gap in plants: pollination, parental fecundity, and offspring demography

Despite extensive study of pollination and plant reproduction on the one hand, and of plant demography on the other, we know remarkably little about links between seed production in successive generations, and hence about long-term population consequences of variation in pollination success. We bridged this "generation gap" in Ipomopsis aggregata, a long-lived semelparous wildflower that is pollinator limited, by adding varying densities of seeds to natural populations and following resulting plants through their entire life histories. To determine whether pollen limitation of seed production constrains rate of population growth in this species, we sowed seeds into replicated plots at a density that mimics typical pollination success and spacing of flowering plants in nature, and at twice that density to mimic full pollination. Per capita offspring survival, flower production, and contribution to population increase (lambda) did not decline with sowing density in this experiment, suggesting that typical I. aggregata populations freed from pollen limitation will grow over the short term. In a second experiment we addressed whether density dependence would eventually erase the growth benefits of full pollination, by sowing a 10-fold range of seed densities that falls within extremes estimated for the natural "seed rain" that reaches the soil surface. Per capita survival to flowering and age at flowering were again unaffected by sowing density, but offspring size, per capita flower production, and lambda declined with density. Such density dependence complicates efforts to predict population dynamics over the longer term, because it changes components of the life history (in this case fecundity) as a population grows. A complete understanding of how constraints on seed production affect long-term population growth will hinge on following offspring fates at least through flowering of the first offspring generation, and doing so for a realistic range of population densities.     

Effects of Habitat Size and Patch Isolation on Reproductive Success of the Serpentine Morning Glory

Abstract: We examined the effects of habitat area and patch isolation on reproductive success in serpentine morning glory ( Calystegia collina [Convolvulaceae]), a primarily self-incompatible clonal plant endemic to serpentine outcrops in northern California's coast ranges. Within a 4000-km ² region, we compared the reproductive success of C. collina on 16 small (<5 ha) and 7 large ( >300 ha) outcrops. Flower and fruit production were significantly higher on large serpentine outcrops than on small outcrops. Fruit production also was positively correlated with the soil's ratio of calcium to magnesium. Successful pollination was positively affected by flower density and the number of other flowering patches within 100 m of a C. collina patch. The number of nearby flowering patches was considerably higher on large than on small outcrops. Flowers on large outcrops did not receive significantly more bee visitors than flowers on small outcrops, suggesting that pollination success is related to the quality rather than the quantity of pollen deposited. Fruit production by plants on both small and large outcrops was enhanced by the experimental addition of pollen from other patches, but not by the addition of pollen from the same patch. These findings demonstrate that the size of habitat may have strong effects on the reproductive success of locally endemic plants by enhancing opportunities for successful sexual reproduction. They also warn against the presumption that naturally patchy plant species are invulnerable to the effects of habitat fragmentation.     

Experimental Demonstration of an Allee Effect in American Ginseng

Abstract: Harvesting of wild American ginseng (   Panax quinquefolius ) for the herbal trade has lowered natural population sizes. We tested for reproductive limitation due to small population size (a form of the Allee effect) by experimentally planting "natural" populations numbering 4, 16, and 64 using 4-year-old cultivated plants. Plant size traits and reproductive traits ( bud, flower, green fruit, and mature fruit) were recorded through the ensuing summer. Fruit production per flower and per plant increased in proportion to flowering population size (  p = 0.0063 and p = 0.0017, respectively), strongly suggesting that an Allee effect occurs in very small populations. The increase in fruit production was not explained by either plant or inflorescence size differences. Although population size-dependent pollination, through insufficient pollinator visitation rate or pollen transfer rate, seems the most likely cause of the observed effects, our limited observations of pollinators were not sufficient to demonstrate a change in pollination rates as a function of population size. Knowledge of the presence as well as the mechanism underlying this Allee effect may be especially useful for management and determination of minimum viable population size of the species in the wild.     

Ontogenetic shifts in fusion–rejection thresholds in a colonial marine hydrozoan,<Emphasis Type="Italic"> Hydractinia symbiolongicarpus</Emphasis>

Like many modular organisms, genetically distinct colonies of the hydrozoan Hydractinia symbiolongicarpus naturally fuse to produce chimeras. One of the principal cooperative benefits of fusion arises from the increased size of the resulting chimeric individual, which may enhance survivorship. However, fusion also promotes conflict through competition between cell lineages for representation in reproductive tissues. Previous studies on H. symbiologicarpus show that, consistent with kin selection theory, a highly polymorphic self/non-self recognition system limits fusion to close kin. However, these recognition systems are intrinsically subject to error. Conspecific acceptance threshold theory predicts that as the costs and benefits of making recognition errors change, or the frequencies of encounters between acceptable and unacceptable kin vary, the recognition system should respond. Specifically, as the benefits of acceptance decline or the frequency of encounters with unacceptable individuals increases, the acceptance threshold should become more restrictive. We tested this hypothesis by monitoring changes in the expression of fusion/rejection behaviors of H. symbiolongicarpus during colony establishment, a period of high mortality when the size-dependent benefits of fusion may be changing most rapidly, and the frequency of encounters with close kin declines. Across seven full-sib families, fusion frequencies between pairs of sibling colonies declined from 73% for 3-day-old colonies to 58% by day 12. This decline is consistent with optimal acceptance threshold theory. However, the period of maximum decline also corresponds to an interval during which the recognition effector mechanism becomes fully functional, suggesting that the shift to a more restrictive conspecific acceptance threshold may reflect an intrinsic constraint on recognition system maturation.Communicated by T. Czeschlik     

Energy allocation in a reef coral under varying resource availability

An organism’s pattern of resource allocation to reproduction and growth over time critically impacts on its lifetime reproductive success. During times of low resource availability, there are two fundamental, mutually exclusive strategies of energy investment: maintenance of somatic tissues to support survival and later reproduction or investment into an immediate reproductive event at the risk of subsequent death. Here, we examine energy allocation patterns in the coral Montipora digitata to determine whether energy investment during periods of resource shortage favours growth or reproduction in a sessile, modular marine species. We manipulated light regimes (two levels of shading) on plots within a shallow reef flat habitat (Orpheus Island, Great Barrier Reef, Australia) and quantified energy uptake (rates of net photosynthesis), energy investment into reproduction (E _R ), tissue growth per unit surface area (E _T ) and energy channelled into calcification (E _C ). With declining resource availability (i.e. reduced photosynthesis), relative energy investment shifted from high (~80%) allocation to tissue growth (E _R :E _T :E _C  = 11:81:8%) to an increasing proportion channelled into reproduction and skeletal growth (20:31:49%). At the lowest light regime, calcification was maintained but reproduction was halted and thus energy content per unit surface area of tissue declined, although no mortality was observed. The changing hierarchy in energy allocation among life functions with increasing resource limitation found here for an autotrophic coral, culminating in cessation of reproduction when limitations are severe, stands in contrast to observations from annual plants. However, the strategy may be optimal for maximising fitness components (growth, reproduction and survival) through time in marine modular animals.     

Seasonal decline of offspring quality in the European starling Sturnus vulgaris: an immune challenge experiment

In seasonally fluctuating environments, timing of reproduction is a crucial determinant of fitness. Studies of birds show that late breeding attempts generally result in offspring of lower reproductive value, with lower recruitment and long-term survival prospects. Several proximate mechanisms, including a seasonal decline of immune system functioning, may lead to a seasonal decline of offspring fitness. We investigated seasonal variation in offspring quality by subjecting first- and second-brood chicks of a sexually size dimorphic species, the European starling Sturnus vulgaris, to an immune challenge with a bacterial endotoxin (LPS), and evaluated their growth and physiological response in terms of total plasma antioxidant capacity (TAC), concentration of reactive oxygen metabolites and hematocrit. LPS challenge did not affect chick growth or oxidative status. However, hematocrit of second-brood chicks was higher in LPS chicks compared to controls. Body mass halfway through the rearing period (days?8–9 post-hatching), TAC and hematocrit were lower among second- vs. first-brood chicks. Interestingly, sexual dimorphism in body mass at days?8–9 post-hatching markedly differed between broods, first-brood males being 4.7% and second-brood males 22.7% heavier than their sisters, respectively. Pre-fledging mortality occurred among second-brood chicks only and was strongly female-biased. Our findings suggest that starling chicks, even if in poor conditions, are little affected by a bacterial challenge, at least in the short-term. Moreover, our study indicates that sex differences in body size, possibly mediated by sex-specific maternal investment in egg size, may heavily impact on pre-fledging survival in a different way in the course of the breeding season, resulting in sex-specific seasonal decline of offspring fitness. Finally, we suggest that levels of circulating antioxidants should be regarded among the proximate causes of the association between timing of fledging and long-term survival in avian species.     

Reproductive Success Increases with Local Density of Conspecif ics in a Desert Mustard (Lesquerella fendleri)

We investigated the effects of plant density on reproduction for an insect-pollinated desert mustard (Lesquerella fendleri [Brassicaceae]). Individual reproductive success, as measured by seeds per fruit, proportion of flowers setting fruit, and total seed production, increased with the density of conspecifics within 1 m. However, including the density of conspecifics at greater distances (1–3 m) did not significantly increase the amount of variation in reproductive success explained by the regression model. This implies that processes occurring on a scale of 1 m or less have important effects on reproduction. Total seed production also was greater for high-density plants than for otherwise similar plants with a low-density of conspecifics. We argue that increased pollinator visitation is the most likely cause of this facilitation and that investigations of the effects of rarity on reproductive success should directly consider density along with more commonly used attributes such as population size and fragmentation.     

Optimal fishery policy with artificial enhancement through stocking: California's white sturgeon as an example

Although optimal fishery policy has been derived from different kinds of economic and biological models, the interaction of fishing policy with artificial stocking policy has not been explicitly considered. We here determine optimal size limits, fishing effort, and stocking rate for three cases of interest: (1) recruitment-limited population, pre-recruitment stocking; (2) adult biomass-limited population, post-recruitment stocking; and (3) adult biomass-limited and recruitment-limited population, post-recruitment stocking. Results show that lower size limits should be set at the size at which the current market value exceeds the total future value of an individual, both to the fishery and to reproduction. Imposition of upper size limits is rarely optimal. Stocking is advisable when the hatchery cost times the relative contribution of stocking to recruitment is less than the contribution to the value of the catch. Optimal policy ranges from infinite effort at a specific size limit with maximum stocking when the cost of stocking is zero, to lower values of size limit and effort as stocking costs increase, the amount of stocking decreases, and more natural reproduction is optimal. Thus, as hatchery costs decline (or value of captured fish increases), optimal stocking/fishery policy varies from an unstocked fishery to a “put and take” fishery. The results are applied to the sturgeon fishery in the San Francisco Bay Estuary as an example. They imply that a reduced lower size limit and greater fishing mortality together with stocking would be optimal, but that current levels are conservative. The stocking decision depends critically on the values of parameters that are currently poorly known, such as: hatchery costs, survival to the fishery and the mechanisms controlling the sturgeon population.     

Continuous growth facilitates feeding and reproduction: impact of size on energy allocation patterns for organisms with indeterminate growth

Body size has great influence on feeding, reproduction, and ecological importance. This study measures growth, reproduction, and feeding for several northeastern Pacific intertidal invertebrates that have indeterminate growth. In all species studied, linear size (length, diameter) showed asymptotic growth fit by the von Bertalanffy growth function, supporting the notion that less energy is allocated to growth with age because of increased reproduction. However, these same species displayed a continuous, roughly linear increase in volume with age. Both reproductive output and food intake were shown to scale proportionally with volume. This indicates that some species with indeterminate growth do not reduce energy allocation to growth with age but instead display continuous volumetric growth that facilitates increases in feeding rate and reproductive output with age and size. A simple allometric model is proposed to describe constant volumetric growth rates and linear increases in reproduction with age.     

Extremes, plasticity, and invariance in vertebrate life history traits: insights from coral reef fishes

Life history theory predicts a range of directional generic responses in life history traits with increasing organism size. Among these are the relationships between size and longevity, mortality, growth rate, timing of maturity, and lifetime reproductive output. Spanning three orders of magnitude in size, coral reef fishes provide an ecologically diverse and species-rich vertebrate assemblage in which to test these generic responses. Here we examined these relationships by quantifying the life cycles of three miniature species of coral reef fish from the genus Eviota (Gobiidae) and compared their life history characteristics with other reef fish species. We found that all three species of Eviota have life spans of < 100 days, suffer high daily mortality rates of 7-8%, exhibit rapid linear growth, and matured at an earlier than expected size. Although lifetime reproductive output was low, consistent with their small body sizes, short generation times of 47-74 days help overcome low individual fecundity and appear to be a critical feature in maintaining Eviota populations. Comparisons with other coral reef fish species showed that Eviota species live on the evolutionary margins of life history possibilities for vertebrate animals. This addition of demographic information on these smallest size classes of coral reef fishes greatly extends our knowledge to encompass the full size spectrum and highlights the potential for coral reef fishes to contribute to vertebrate life history studies.     

Why run and hide when you can divide? Evidence for larval cloning and reduced larval size as an adaptive inducible defense

Predator-induced cloning (asexual reproduction), with reduced size as consequence of cloning, suggests a novel adaptation to the threat of predation. Although cloning is a common reproductive strategy of many plants and animals, cloning in response to stimuli from predators has, at present, been documented only in the larvae (plutei) of the sand dollar, Dendraster excentricus. Other studies report larval cloning in echinoderms under optimal conditions of food and temperature. A burst of asexuality should be favored when environmental conditions are conducive to growth, but it is less clear that cloning is advantageous when conditions indicate risk from predators. This study tested the hypothesis that the small size of predator-induced clones reduces vulnerability during encounters with planktivorous fish. Successful cloning was inferred from an increase in larval density, a reduction in larval size and stage, and some direct observations of budding. All clones were smaller than uncloned sibling larvae, suggesting an advantage against visual predators. Pair-wise predation trials demonstrated that planktivorous fish ate more uncloned sibling plutei than small clones. These results offer a new ecological context for asexual reproduction: rapid size reduction as a defense. If the identifiable cues for cloning in echinoderm larvae (food and predators) are linked in nature, then larval cloning may be a response to a single ecological scenario rather than two separate and unrelated conditions.     

Growth rate of the bathypelagic crustacean Gnathophausia ingens (Mysidacea: Lophogastridae). I. Dimensional growth and population structure

Gnathophausia ingens has 13 instars, each with a distinct range of sizes which does not overlap the sizes of adjacent instars. The intermolt interval, measured in the laboratory at 5.5°, 6.5° and 7.5°C, increases with increasing size and decreases with increasing temperature. At 5.5°C it varies from 166 days for the smallest individuals to 253 days for the oldest. The period of larval development in the marsupium of a female is estimated to be 530 days. The life span of females is estimated to be 2,950 days with the onset of reproduction at 2,400 days. It is sugquested that this species is semelparous. The population structure data suggest that there is low mortality through the first 7 instars, progressively higher mortality from Instar 8 through Instar 11, and slightly lower mortality in the remaining 2 instars. These life-history characteristics appear to be directed toward maximizing absolute fecundity (as opposed to time-specific fecundity) in a stable environment. These characteristics may have been selected for by low available food energy and made possible by the stability of the deep sea.     

A Possible Method for the Rapid Assessment of Biodiversity

Inventories of vertebrate and flowering plants are frequently used as surrogates for estimates of total biodiversity. This is in part because the inclusion of invertebrates and nonflowering plants is perceived as being too time-consuming, costly, and difficult because of the shortage of specialists. Estimates of the species richness of field samples of spiders, ants, polychaetes, and mosses made by a biodiversity technician and by specialist taxonomists were compared. The biodiversity technician received a few hours training in the taxonomy of each group and separated specimens into recognizable taxonomic units (RTUs). The specialists sorted to species. For the three animal groups the biodiversity technician recorded 165 taxa and the specialists 147, with the error for the ants and spiders being 13% or less. A small amount of splitting and lumping of species was detected. The concordance of estimates remained very similar when small subsamples were used. The procedure was repeated by 13 undergraduates using a subsample of spiders. Their average error was 14.4%. The greatest similarity in estimates was for the mosses, but with high levels of splitting and lumping this result was entirely fortuitous. The results suggest that RTU estimates made by biodiversity technicians may be sufficiently close to formal taxonomic estimates of species richness to be useful for the rapid assessment of biodiversity. They also show, however, that the procedures outlined here should be used on invertebrate and nonflowering plant groups before they can be confidently included in biodiversity surveys.     

Age,size, dominance and reproduction among male kudus: mating enhancement by attrition of rivals

Summary Size dimorphism with males larger than females has been related to the benefits for males of enhanced dominance and hence reproductive success. However, mating gains must outweigh the fitness costs of deferred reproduction and the mortality associated with further growth. The relationships between male age, size and reproduction were assessed for greater kudus (Tragelaphus strepsiceros) in the Kruger National Park in South Africa. Individually identifiable animals were monitored over 10 years, with detailed observations made during six breeding seasons. In the non-breeding season males formed loose all-male groups. Horn grappling and low intensity agonistic interactions fostered dominance rankings. Dominance was age-graded, until males reached full weight at 6 years of age. Males aged 6 and 7 years monopolized courtship and mating, but 5-year-old males secured about 10% of mating opportunities. Few males survived beyond 7 years. Male mortality rate rose steeply with age, so that the functional sex ratio of fertile females per mature male was about 14:1. During the breeding season many female groups remained unattended by a mature male. Reproductive sorting among males occurred largely through variation in survival to full size and maturity. Increased size enhances fighting success and hence dominance. Further growth ceases when the functional sex ratio exceeds the number of mating opportunities that males can effectively achieve during a breeding season. Predation amplifies the mortality cost of continued growth. In the absence of large predators, male-male interactions may be atypically exaggerated.     

温带落叶林的植物物候特征及其对气候变化的响应

植被物候是气候变化对生物圈产生长期或短期影响的重要指示因子。气候变化已经明显改变了许多物种的营养生长和繁殖物候,尤其是在温带地区。研究温带森林物候变化及其对全球变暖的响应,对认识森林物种共存,协同进化以及森林保护和经营等有重要意义。通过概述温带森林下物候研究的进展发现,光照和积温是影响木本植物展叶及繁殖物候的关键因素,林下层树木通过更早展叶,以尽量减少生长季林冠层遮阴对下层树木生长的影响,更早时期开花的树木具有从顶部向四周次第开花的时空格局,林冠层树种开花具有较好的同步性。而草本植物的物候通常受融雪时间和冠层动态的影响更大,并且,温带森林下不同生活史对策的草本植物的物候特征对气候变化的响应也不尽相同,存在明显的季节动态。繁殖物候、光照的季节变化、光合特征、授粉成功之间的联系决定了林下不同繁殖特性的草本植物的繁殖成功率。量化的、多指标、多对象的定位监测是精准物候研究的基础,物候变化的机理和建立可预测的物候模型将是未来研究的重点。     

Habitat Change and Plant Demography: Assessing the Extinction Risk of a Formerly Common Grassland Perennial

Abstract:  An important aim of conservation biology is to understand how habitat change affects the dynamics and extinction risk of populations. We used matrix models to analyze the effect of habitat degradation on the demography of the declining perennial plant Trifolium montanum in 9 calcareous grasslands in Germany over 4 years and experimentally tested the effect of grassland management. Finite population growth rates (λ) decreased with light competition, measured as leaf-area index above T. montanum plants. At unmanaged sites λ was <1 due to lower recruitment and lower survival and flowering probability of large plants. Nevertheless, in stochastic simulations, extinction of unmanaged populations of 100 flowering plants was delayed for several decades. Clipping as a management technique rapidly increased population growth because of higher survival and flowering probability of large plants in managed than in unmanaged plots. Transition-matrix simulations from these plots indicated grazing or mowing every second year would be sufficient to ensure a growth rate ≥1 if conditions stayed the same. At frequently grazed sites, the finite growth rate was approximately 1 in most populations of T. montanum . In stochastic simulations, the extinction risk of even relatively small grazed populations was low, but about half the extant populations of T. montanum in central Germany are smaller than would be sufficient for a probability of survival of >95% over 100 years. We conclude that habitat change after cessation of management strongly reduces recruitment and survival of established individuals of this perennial plant. Nevertheless, our results suggest extinction processes may take a long time in perennial plants, resulting in an extinction debt. Even if management is frequent, many remnant populations of T. montanum may be at risk because of their small size, but even a slight increase in size could considerably reduce their extinction risk.