Effects of range position,inter-annual variation and density on demographic transition rates of <Emphasis Type="Italic">Hornungia petraea</Emphasis> populations

The central-marginal model assumes unfavourable and more variable environmental conditions at the periphery of a species’ distribution range to negatively affect demographic transition rates, finally resulting in reduced population sizes and densities. Previous studies on density-dependence as a crucial factor regulating plant population growth have mainly focussed on fecundity and survival. Our objective is to analyse density-dependence in combination with the effect of inter-annual variation and range position on all life stages of an annual plant species, Hornungia petraea, including germination and seed incorporation into the seed bank. As previous studies on H. petraea had revealed a pattern opposite to existing theory with lower population densities at the distribution centre in Italy than at the periphery in Germany, we hypothesised that (1) demographic transition rates are lower, (2) the inter-annual variation in demographic transition rates is higher and (3) the intensity of density-dependence is weaker in Italy than in Germany. To analyse demographic transition rates, we used an autoregressive covariance strategy for repeated measures including density and inter-annual variation. All the three hypotheses were confirmed, but the impact of range position, density-dependence and inter-annual variation differed among the transition steps. All transition rates except fecundity were higher in the German populations than in the Italian populations. Germination rate and incorporation rate into the seed bank were strongly density-dependent. Central populations showed a larger inter-annual variation in fecundity and winter survival rate. Winter survival rate was the only transition step with a stronger density-dependence in peripheral populations. In most cases, these differences between distribution centre and periphery would not have emerged without taking density-dependence and inter-annual variation into account. We conclude that including range position, inter-annual variation and density-dependence in one single statistical model is an important tool for the interpretation of demographic patterns regarding the central-marginal model. Electronic Supplementary Material Supplementary material is available for this article at     

Global asymptotic stability of plant-seed bank models

Many plant populations have persistent seed banks, which consist of viable seeds that remain dormant in the soil for many years. Seed banks are important for plant population dynamics because they buffer against environmental perturbations and reduce the probability of extinction. Viability of the seeds in the seed bank can depend on the seed’s age, hence it is important to keep track of the age distribution of seeds in the seed bank. In this paper we construct a general density-dependent plant-seed bank model where the seed bank is age-structured. We consider density dependence in both seedling establishment and seed production, since previous work has highlighted that overcrowding can suppress both of these processes. Under certain assumptions on the density dependence, we prove that there is a globally stable equilibrium population vector which is independent of the initial state. We derive an analytical formula for the equilibrium population using methods from feedback control theory. We apply these results to a model for the plant species Cirsium palustre and its seed bank.     

Interactions between density-dependent processes, population dynamics and control of an invasive plant species, Tripleurospermum perforatum (scentless chamomile)

Tripleurospermum perforatum is an invasive weedy species which exhibits strong over-compensating density dependence. Interactions between density-dependent survival, probability of flowering and fecundity were modelled and their impact on the population dynamics were examined. When only fecundity was density-dependent, the dynamics were similar to those observed in the model containing all three density-dependent terms. Density-dependent survival was a stabilizing process when acting in combination with density-dependent fecundity and probability of flowering; removing density-dependent survival from the model produced two-point cycles. The addition of a seed bank was also stabilizing. Simulations of control strategies at different life-history stages indicated that full control would be difficult due to the strong over-compensating density dependence, with severe reductions in fecundity and late season survival necessary in order to reduce equilibrium seed density and biomass.     

Spatially structured population dynamics in feral oilseed rape

We studied the population dynamics of feral oilseed rape (Brassica napus) for 10 years (1993-2002) in 3658 adjacent permanent 100 m quadrats in the verges of the M25 motorway around London, UK. The aim was to determine the relative importance of different factors affecting the observed temporal patterns of population dynamics and their spatial correlations. A wide range of population dynamics was observed (downward or upward trends, cycles, local extinctions and recolonizations), but overall the populations were not self-replacing (lambda < 1). Many quadrats remained unoccupied throughout the study period, but a few were occupied at high densities for all 10 years. Most quadrats showed transient oilseed rape populations, lasting 1-4 years. There were strong spatial patterns in mean population density, associated with soil conditions and the successional age of the plant community dominating the verge, and these large-scale spatial patterns were highly consistent from year to year. The importance of seed spilled from trucks in transit to the processing plant at Erith in Kent was confirmed: rape populations were significantly higher on the 'to Erith' verge than the 'from Erith' verge (overall mean 2.83-fold greater stem density). Quadrats in which lambda > 1 were much more frequent in the 'to Erith' verge, indicating that seed immigration can give the spurious impression of self-replacing population dynamics in time-series analysis. There was little evidence of a pervasive Moran effect, and climatic forcing did not produce widespread large-scale synchrony in population dynamics for the motorway as a whole; just 23% of quadrats had significant rank correlations with the mean time-series. There was, however, significant local spatial synchrony of population dynamics, apparently associated with soil disturbance and seed input. This study draws attention to the possibility that different processes may impose population synchrony at different scales. We hypothesize that synchrony in this system is driven by at least three processes: small-scale, local forcing caused by soil disturbance, intermediate-scale forcing as a result of seed input, and large-scale climatic forcing (e.g. winter rainfall) that affects the motorway as a whole.     

Plant Pattern Development during Early Post‐Restoration Succession in Grasslands—A Case Study of Arabis nemorensis

The main objective of this study was to analyze whether plant material transfer is a suitable approach for establishing plant populations with spatial patterns and population structures resembling those of remnant populations. We studied pattern formation and population characteristics in three remnant and two restored populations of the biennial plant species Arabis nemorensis in the upper Rhine valley in southwestern Germany over a period of 2 years. We investigated spatial patterns of seedlings, juveniles, and adults as well as the small‐scale horizontal and vertical distribution of seeds in the soil, followed the fate of individual plants and recorded structural habitat parameters such as vegetation and litter cover. Population dynamics differed between the study sites, but there was no pronounced difference between restored and remnant sites. Seedlings, juveniles, and adults as well as seeds in the seed bank showed aggregated spatial patterns on all study sites, with positive autocorrelation on a scale of 20–60 cm. Within sites, patterns remained approximately stable through time. Restored sites experienced rapid seed bank formation as a result of the restoration measures. Our results suggest that the restoration measures were not only successful in transferring the target species but also triggered rapid formation of spatially structured populations that, years after restoration, closely resembled those of remnant sites.     

Belowground bud banks and meristem limitation in tallgrass prairieplant populations

Rhizome meristem populations were sampled in tallgrass prairie to quantify the size, grass?:?forb composition, and temporal and spatial variability of the soil bud bank and to compare fire effects on bud bank and seed bank composition. Soil cores (10.5 cm diameter, 15 cm deep) were collected from replicate annually and infrequently burned tallgrass prairie sites, and intact rhizomes and rhizome buds were censused. Bud bank densities ranged from approximately 600 to 1800 meristems/m(2) among sites and had high spatial and seasonal variability. In annually burned prairie, the total bud bank density was two-fold greater and the grass?:?forb meristem ratio was more than 30-fold greater than that of infrequently burned prairie. These patterns are opposite those observed in soil seed banks at this site. The rhizome population in annually burned prairie was 34% larger than the established aboveground tiller population. By contrast, the bud bank density in unburned prairie was significantly lower than aboveground stem densities, indicating possible belowground meristem limitation of stem density and net primary production on infrequently burned prairie. The patterns observed in this study suggest that the densities and dynamics of tallgrass prairie plant populations, as well as their response to disturbance (e.g., fire and grazing) and climatic variability, may be mediated principally through effects on the demography of belowground bud populations. Patterns of seed reproduction and seed bank populations have little influence on short-term aboveground population dynamics of tallgrass prairie perennials.     

Introduced populations of <Emphasis Type="Italic">Genista monspessulana</Emphasis> (French broom) are more dense and produce a greater seed rain in California,USA, than native populations in the Mediterranean Basin of Europe

Some invasive plants perform better in their area of introduction than in their native region, and this is often attributed either to phenotypic responses and/or to adaptive evolution following exposure to new environmental conditions. Genista monspessulana (French broom) is native to Europe, but highly invasive and abundant along the Pacific Coast of the USA. In this study, the population density and age structure, plant growth and reproductive traits, and seed bank characteristics of 13 native (Mediterranean Basin) and 15 introduced (California, USA) field populations of G. monspessulana were compared. Mean population density, plant height and stem diameter were greater in introduced populations, with the latter two traits explained by a greater mean plant age. Age structure also showed a greater percentage of seedling plants in introduced populations. Fecundity was higher in introduced populations when measured in terms of mature seeds per pod, but lower when comparing seed production per plant (number of pods and mature seeds). Thus, seed rain and seed bank size was considerably higher in introduced populations. Results from this study indicate that G. monspessulana performs better in its introduced region. We hypothesize that release from natural enemies and competitors together with more favorable environmental conditions in the introduced region may explain the invasion success of G. monspessulana. As a result, an integrated management approach using introduced seed predators to suppress seed production and selected management practices to reduce seed banks may be needed for effective long-term control in California.     

Development of vegetation patterns in early primary succession

Question: We investigated colonisation filters in early plant community development on a glacial outwash plain. We asked if these were related to seed limitation or to a lack of safe sites, if topographical heterogeneity affected species patchiness and how species life cycles influence successional trajectories. Location: An outwash plain (Skeiðarársandur) in southeast Iceland. Methods: We identified surface heterogeneity at two different scales, ca. 10–15 cm (larger stones and established plants) and ca. 50 m (shallow depressions representing dry river beds) at two study sites. We quantified species cover, flowering plant density, seed production, seed rain, seed bank density, seedling emergence and seedling survival from June 2005 to June 2007 for the whole plant community, and measured seed production for five species. Results: Mean vegetation cover was <2.5% at the sites. Low emergence rates and high seedling mortality were the two main recruitment filters. Only 1.4% of seedlings emerging in 2005 survived into the 2007 growing season. Topographical heterogeneity had little effect on plant colonisation. High annual variation was recorded, and the two study sites (ca. 2 km apart) differed in their colonisation success. Of the five species, establishment of Cerastium alpinum and Silene uniflora was most limited by lack of seeds, whereas establishment of Luzula spicata, Poa glauca and Rumex acetosella was most limited by safe sites. Conclusions: We conclude that colonisation processes and patterns in early primary succession on Skeiðarársandur were largely influenced by stochastic factors.     

Ecological relationships between seedling emergence and litter cover in the earliest stage of plant succession on sandy soil

The relationships between seedling emergence and litter cover were studied in the earliest successional stage of a plant community. During a period of 3 years, changes in vegetation cover and species composition were assessed on three permanent quadrats with virgin sandy soil substrates A successional pattern from summer annual grasses to winter annual herbacoeus plants to biennial and perennial plants was confirmed, and invasion and replacement of the component species were conspicuous. The spatial distributions of seedlings and litter were heterogeneous in winter, and some patches consisting of both seedlings and litter were observed. There was a strong positive correlation between distribution patterns of seedlings and litter; seedling density increased with increasing litter cover. The results suggest that the litter cover of the previous stage plays an imporrant role in promoting successional changes in the early stage of plant succession.     

Populational dynamics of three annual species of alpine plants in the Rocky Mountains

Summary Survivorship, fecundity, and seed bank size were measured over two years in alpine populations of the annuals, Koenigia islandica, Polygonum confertiflorum, and P. douglasii. The major loss of individuals from all populations occurred between seed dispersal and germination. Survival of vegetative plants to maturity was high in all species, usually above 80%, and the average number of seeds produced per plant was less than 10 in all species in both years. Seed banks existed for all three species but were small since more than 80% of the viable seeds in the soil germinated in the spring. Both survival and fecundity were negatively correlated with density in P. confertiflorum. These patterns of population dynamics are similar to those found in many annuals of temperate environments.     

Demographic response of plant populations to habitat fragmentation and temporal environmental variability

Many plant species currently exist in fragmented populations of different sizes, while they also experience unpredictable climatic fluctuation over time. However, we still understand little about how plant demography responds to such spatial and temporal environmental variability. We studied population dynamics of an understory perennial herb Trillium camschatcense in the Tokachi plain of Hokkaido, Japan, where a significant effect of forest fragmentation on seedling recruitment was previously reported. Four populations across a range of fragment sizes were studied for 6 years, and the data were analyzed using matrix population models. Per capita fecundity (the number of recruits per plant) varied greatly among populations, but the variation in population growth rates (λ) was mainly driven by the variation in stasis and growth rates, suggesting that the general trend of reduced fecundity in fragmented populations may not be readily translated into subsequent dynamics. Temporal variation in λ among years was more than 2 times larger than spatial variation among populations, and this result was likely attributable to the contrasting response of correlation structures among demographic rates. The among-population variation in λ was dampened by negative covariation between matrix elements possibly due to density-dependent regulation as well as an inherent constraint that some elements are not independent, whereas positive covariation between matrix elements resulted in large temporal variation in λ. Our results show that population dynamics responded differently to habitat fragmentation and temporal variability of the environment, emphasizing the need to discriminate these spatial and temporal variations in demographic models. Although no populations were projected to be declining in stochastic simulations, correlation between current habitat size and plant density implies historical λ is positively related to habitat size.     

不同密度条件下芨芨草空间格局对环境胁迫的响应

基于小尺度上植物间相互作用与空间格局的高度相关性,选用可避免环境异质性影响的K2点格局函数,研究了沙枣-芨芨草群落中63个芨芨草样方(3密度条件×3生境条件×7重复)中芨芨草种群在小尺度下(0—0.5 m)的空间格局,探讨了在3种密度条件(高、中、低)及3种生境条件下(冠盖区、过渡区、空旷区),芨芨草种群空间格局对土壤理化性质胁迫(盐分、有机质及容重)程度的响应。研究结果表明,就土壤盐分,养分含量及土壤容重而言,土壤理化性质胁迫程度沿冠盖区、过渡区和空旷区增加的趋势。相应地,芨芨草种群呈聚集分布的样方比例在沿冠盖区(6/21)较低,而在过渡区(11/21)和空旷区(11/21)较高。然而,在不同密度条件下,芨芨草种群空间格局对土壤性质胁迫的响应不同。在低密度条件下,芨芨草种群在冠盖区多数为聚集分布(4/7),在过渡区和空旷区全部为聚集分布;中密度条件下,芨芨草聚集分布样方比例沿冠盖区-过渡区-空旷区梯度增加(分别为2/7,3/7,4/7),但低于同等胁迫条件下低密度样方中聚集分布数量;高密度条件下,在各胁迫条件下,除了一个过渡区高密度样方,其余芨芨草种群均为随机分布。总体上,随着环境胁迫增强,在中低密度下,芨芨草种群趋向于聚集分布;但在高密度下,芨芨草种群均以随机分布为主。此外,芨芨草种群空间格局随密度变化趋势比随土壤理化性质胁迫梯度变化趋势更加明显,可能表明相对于土壤理化性质胁迫程度,芨芨草种群密度对其空间格局影响更大。因此,在考虑芨芨草种群空间格局对环境胁迫的响应时,应当考虑种群密度因素。     

Population biology of <Emphasis Type="Italic">Kosteletzkya pentacarpos</Emphasis> (Malvaceae) in the Llobregat delta (Catalonia,NE of Spain)

Field experiments and wild population monitoring have been performed to study the population biology of the rare long-lived Kosteletzkya pentacarpos (Malvaceae) in the Llobregat delta (Catalonia, NE Spain). Field experiments explored the fate of seeds in soil at different depths, seedling emergence, and seedling survival, growth and flowering with and without canopy cover during the first 2 years of life. They also were used to ascertain the size-related pattern of seedling survivorship and flowering. Field data concerning mortality, growth and fecundity of adult plants were collected yearly in three wild populations for 7–9 years. In old adults (reproducing long before the beginning of the study), ANOVAR tests were performed to compare maximum diameter, total and fertile shoots, and viable seeds per plant between years and populations. New adults (starting flowering the first year of study or in subsequent years) were used to explore, using linear and polynomial regressions, the association of RGR and both total and fertile shoot production with (i) plant size (maximum basal diameter or its logarithm); (ii) plant age (years in adult stage); and (iii) plant age after removing the effect of size and year-to-year fluctuations. In this case, we examined the age-related pattern of the residuals obtained from the regressions with size and year. The study identified the following main demographic features of K. pentacarpos: (i) transient, shallow soil seed bank; (ii) shade tolerance of seedling emergence; (iii) canopy-facilitation of seedling survival and bolting during the first two years of life; (iv) size-related pattern for seedling survivorship but not flowering; (v) exclusive dependence on a fluctuating seed output for reproduction; (vi) rapid adult growth; and (vii) high adult longevity but (viii) rapid depletion of fecundity with age. Seed output was highly constrained by mining insects. The changing size-structure and the decreasing reproductive success of old adults in several populations suggest that K. pentacarpos might undergo a dynamics of population establishment and extinction in the Ricarda marshes. Because of fluctuating reproduction and the lack of a persistent seed bank, the conservation of standing adult populations appears to be a key factor to ensure the persistence of the species.     

Plant litter feedback and population dynamics in an annual plant,<Emphasis Type="Italic"> Cardamine pensylvanica</Emphasis>

The presence of litter has the potential to alter the population dynamics of plants. In this paper, we explore the effects of litter on population dynamics using a simple experimental laboratory system with populations of the annual crucifer, Cardamine pensylvanica. Using a factorial experiment with four densities and three litter levels, we determined the effect of litter on biomass and plant fecundity, and the life stages responsible for these changes in yield. Although litter had significant effects on seed germination and on seedling survivorship, we show, using a population dynamics model, that these effects were not demographically significant. Rather, the potential effect of litter on population dynamics resulted almost entirely from its effect on biomass. Persistent litter suppressed plant biomass and apparently removed the direct density effect present in the absence of litter. Thus, litter changed the shape of the recruitment curve from slightly humped to asymptotic. In addition to changing the shape of the recruitment curve, litter reduced the carrying capacity of the populations. Thus, the population dynamics model indicated that not all statistically significant responses were dynamically significant. Given the potential complexity of litter effects, simple population models provide a powerful tool for understanding the potential consequences of short-term responses. Received: 8 September 1999 / Accepted: 5 April 2000     

Linking traits of foraging animals to spatial patterns of plants: social and solitary ants generate opposing patterns of surviving seeds

Foraging traits of seed predators are expected to impact the spatial structure of plant populations, community dynamics and diversity. Yet, many of the key mechanisms governing distance- or density-dependent seed predation are poorly understood. We designed an extensive set of field experiments to test how seed predation by two harvester ant species interact with seed dispersal in shaping the spatial patterns of surviving seeds. We show that the Janzen–Connell establishment pattern can be generated by central-place foragers even if their focal point is located away from the seed source. Furthermore, we found that differences in the social behaviour of seed predators influence their sensitivity to seed density gradients and yield opposing spatial patterns of surviving seeds. Our results support the predictions of a recent theoretical framework that unifies apparently opposing plant establishment patterns, and suggest that differences in foraging traits among seed predators can drive divergent pathways of plant community dynamics.     

Red environmental noise and the appearance of delayed density dependence in age-structured populations

Previous work suggests that red environmental noise can lead to the spurious appearance of delayed density dependence (DDD) in unstructured populations regulated only by direct density dependence. We analysed the effect of noise reddening on the pattern of spurious DDD in several variants of the density-dependent age-structured population model. We found patterns of spurious DDD in structured populations with either density-dependent fertility or density-dependent survival of the first age class, inconsistent with predictions from unstructured population models. Moreover, we found that nonspurious negative DDD always emerges in populations with deterministic chaotic dynamics, regardless of population structure or the type of environmental noise. The effect of noise reddening in generating spurious DDD is often negligible in the chaotic region of population deterministic dynamics. Our findings suggest that differences in species' life histories may exhibit different patterns of spurious DDD (owing to noise reddening) than predicted by unstructured models.     

Ocean temperature and density dependence as key drivers of the population dynamics of an intertidal crab at the Brazilian oceanic islands

Understanding how climatic and density-dependent processes affect demography is crucial for predicting population responses to climate change. For marine invertebrates with complex life cycle such as decapod crustaceans, increasing temperatures might affect survival and development of early pelagic stages, whereas high density can increase competition and thus reduce growth and fecundity of older life stages. In this study, we investigate the effects of warm ocean events, body size and density on the population dynamics of the intertidal Sally lightfoot crab (Grapsus grapsus) at the Brazilian oceanic islands. Firstly, we assessed the trends of marine heatwaves (MHW) and positive temperature anomalies (ΔSST+) at the equatorial St Peter and St Paul (SPSP) Archipelago and Rocas Atoll and the subtropical Trindade Island. We then jointly analyzed short-term count, capture-recapture and fecundity data, and long-term population monitoring data (2003–2019) using an integrated population model. Warm ocean events have become more frequent and intense only at the equatorial islands. Increasing MHW frequency positively influenced recruitment in the high-density SPSP population, while MHW intensity and ΔSST+ frequency had negative impacts. Conversely, no climatic effects were observed for the low-density Rocas population, which has the largest crabs. Despite a lack of warming in Trindade, this subtropical population with intermediate density and body size was negatively affected by ΔSST+. Our findings revealed population-specific responses to climate change when accounting for local life history and ecology. Thus, environmental and density-dependent effects should be broadly considered in future conservation studies regarding ocean warming impacts on marine invertebrate populations.     

Prediction of cutting effects on a population of Chaerophyllum aureum– a demographic approach

Abstract. A demographic study of the undesirable perennial weed Chaerophyllum aureum in extensified French Pyrenean hay meadows was carried out. For two successive years the effect of date and intensity of cutting on the population demography of this colonizer. Population density is sensitive to cutting practices essentially through fecundity, seedling survival rate and seed dispersal between fields. Cutting reduces fecundity by preventing seed formation (early cutting) or by exportation of some mature seeds with hay (late cutting). Nevertheless, the number of seeds transported between field populations by hay harvest, organic manuring and associated cutting practices compensates for the lack of seed production in early‐cut populations, maintaining them at relative high densities. For each of three cutting regimes, the number of immigrant and emigrant seeds has been indirectly estimated from a prediction of population density at equilibrium from demographic parameters measured in field populations and compared with observed population densities.     

Floater dynamics can explain positive patterns of density-dependent fecundity in animal populations

After some 70 years of debate on density-dependent regulation of animal populations, there is still poor understanding of where spatial and temporal density dependence occurs. Clearly defining the portion of the population that shapes density-dependent patterns may help to solve some of the ambiguities that encircle density dependence and its patterns. In fact, individuals of the same species and population can show different dynamics and behaviors depending on their locations (e.g., breeding vs. dispersal areas). Considering this form of intrapopulation heterogeneity may improve our understanding of density dependence and population dynamics in general. We present the results of individual-based simulations on a metapopulation of the Spanish imperial eagle Aquila adalberti. Our results suggest that high rates of floater mortality within settlement areas can determine a shift in the classical relationship (from negative to positive) between the fecundity (i.e., fledglings per pair) and density (i.e., number of pairs) of the breeding population. Finally, we proved that different initial conditions affecting the breeder portion of the population can lead to the same values of fecundity. Our results can represent a starting point for new and more complex approaches studying the regulation of animal populations, where the forgotten and invisible component--the floater--is taken into account.     

植物种群更新的补充限制

补充限制基于生态位理论, 从种子萌发、幼苗存活和生长、繁殖体扩散等生活史阶段的种群统计特征及环境因素(土壤水分、养分、凋落物等)着手, 探讨种群的更新问题。种源限制和微生境限制是补充限制理论研究的核心内容, 但是哪个更为重要并没有统一的结论。种源限制与种子生产、土壤种子库和地下芽库中的繁殖体数量不足有关。其中, 气候的年际波动、土壤种子库寿命和动物的捕食都会影响种子生产在种群更新中的作用; 土壤种子库常被视为种群更新的保险库, 与地上种子雨共同促进种群更新, 但是, 如果土壤里种子具有较高的死亡率和休眠率, 将会降低种子库的作用; 地下芽库及其产生的无性分株对于种群更新的意义更多地体现在干扰后种群更强的恢复能力上。扩散限制是种群更新中的普遍现象, 与种子产量、散布能力、传播媒介、幼苗密度等因素有关。微生境限制主要表现为水分、养分、凋落物等非生物因素以及竞争、捕食等生物因素对种子的活力、萌发性、幼苗的存活力、物质分配等过程的影响, 其重要性随着植物生活史阶段而发生变化。未来需要进行综合的、长期的实验, 并应着重加强种源限制及相关生态过程的进化与生态相结合的机理性研究, 从而更深刻地认识和理解种群更新问题, 建立更为综合、系统的种群更新理论体系。