Population dynamics of a threatened cactus species: general assessment and effects of matrix dimensionality

Harrisia portoricensis is an endemic Caribbean cactus currently under threatened status. In this study we used population projection matrices to evaluate the conservation status of this species and we performed a systematic analysis of the effects of matrix dimensionality on the inferred demographic parameters. Results revealed that population growth rates (λ) were 0.946 and 0.961 for the 2007–2008 and 2008–2009 periods respectively, suggesting a declining population with limited persistence ability. Even when the highest elasticity values corresponded to the survival of adults, numerical simulations suggested that increases in either seedling establishment or fecundity could render λ > 1. Our empirical-based analysis using raw demographic data revealed a clear trend for λ values to decrease with increasing matrix dimension. Stasis and fecundity elasticities were also found to decrease whereas retrogression and growth elasticitites increased with increasing matrix dimension. These results are roughly insensitive to the method used to create matrices of different dimensions. For H. portoricensis, large matrices with narrow classifications were required to minimize variations in λ, highlighting the need for large data sets to assess the convergence of results with matrix dimensionality. Our combined results emphasize that under current scenarios the ability of H. portoricensis for population growth is severely limited. Any management strategy designed for the conservation of this species should consider long-term monitoring of populations as well as programs that enhance seedling establishment and adult survival.     

高寒地区不同海拔藏茴香茎解剖结构比较研究

采用石蜡切片法,分别对5种不同海拔高度藏茴香茎初生解剖结构进行比较研究,并用一维和二维(面积)的度量指标进行观测,以期探索随海拔升高,藏茴香茎结构的变化规律,从而探讨藏茴香对高寒气候的适应性,为藏茴香这一食用、药用植物资源开发利用提供基础资料。结果表明:藏茴香茎的初生结构由表皮、皮层、维管柱构成,随着海拔升高,藏茴香茎的表皮厚度、髓厚度、髓射线宽度以及通气组织的面积逐渐增加,而维管组织与基本组织的比值不断缩小,皮层的细胞层数、厚度逐渐下降;维管束的数量、厚度不断降低,但维管束的发育程度和导管的直径有逐渐增加的趋势。藏茴香茎表皮的不断增厚,可以减少植物体内热量的散失,并有效地降低强紫外线对植物造成的伤害。基本组织比例的增加,有利于贮藏更多的生理调节性物质,增加植物的抗寒能力;通气组织逐渐发达,可极大地降低空气在植物体内的扩散阻力,形成氧气由地上部运输到地下部的通道,还可增加植物体的浮力,减轻了植物体比重,对植物体起到了良好的支撑作用。藏茴香茎解剖结构随海拔的变化,体现了植物对环境的适应性。研究中发现的随着海拔升高,茎皮层的细胞层数和厚度以及茎中维管束的数量和厚度均下降,但维管束的发育程度和导管直径增加的现象,还有待于进一步研究。     

Empirical and virtual investigation of the population dynamics of an alien plant under the constraints of local carrying capacity: Heracleum mantegazzianum in the Czech Republic

During the last decades, invasive alien species have become a global concern because of their ecological and economic impact. Heracleum mantegazzianum (Apiaceae), is a tall monocarpic perennial native to Caucasus and invasive in Europe since the 1950s. Within an interdisciplinary EU project aimed at assessing suitable management strategy, we analysed the demography and ecology of this species in its invasive range. The monitoring of population dynamics in the Czech Republic led to the result that in the observed sites the species showed decreasing populations. To find an explanation for this unexpected result, two types of models were parameterized, based on the empirical data: (1) a stage-based transition matrix model, which projected a continuous negative development, and (2) a spatially explicit individual-based model (IBM), including individual variation. This second model was able to create a population with steady individual numbers. Analyses of the simulation showed that in more than 54% of the simulated years (n=5000) the growth rate was smaller than one. Still, population increase in the remaining years was sufficient to sustain a population. Nevertheless long-term observations document an invasive behaviour of the observed populations. Hence, we could assume temporal changes in the course of an invasion and thus wanted to evaluate the probability of sampling negative growth in dependence of time since first invasion. By using a method from ‘Virtual Ecology’, we approached the question: first we create an invasive population, based on the empirical data of H. mantegazzianum and second empirical sampling techniques were mimicked using the Virtual Ecologist approach. The results demonstrate how the probability of sampling negative growth increases with time since first invasion. Hence, we assume that the studied populations have already reached a maximum of their local invasive potential and thus stagnate in their size.     

Potential impact of harvesting on the population dynamics of two epiphytic bromeliads

Large numbers of epiphytes are extracted from cloud forests for ornamental use and illegal trade in Latin America. We examined the potential effects of different harvesting regimes on the population dynamics of the epiphytic bromeliads Tillandsia multicaulis and Tillandsia punctulata. The population dynamics of these species were studied over a 2-year period in a tropical montane cloud forest in Veracruz, Mexico. Prospective and retrospective analyses were used to identify which demographic processes and life-cycle stages make the largest relative contribution to variation in population growth rate (λ). The effect of simulated harvesting levels on population growth rates was analysed for both species. λ of both populations was highly influenced by survival (stasis), to a lesser extent by growth, and only slightly by fecundity. Vegetative growth played a central role in the population dynamics of these organisms. The λ value of the studied populations did not differ significantly from unity: T. multicaulis λ (95% confidence interval) = 0.982 (0.897–1.060) and T. punctulata λ = 0.967 (0.815–1.051), suggesting population stability. However, numerical simulation of different levels of extraction showed that λ would drop substantially even under very low (2%) harvesting levels. Matrix analysis revealed that T. multicaulis and T. punctulata populations are likely to decline and therefore commercial harvesting would be unsustainable. Based on these findings, management recommendations are outlined.     

Population dynamics of the perennial Plantago media in semi‐natural grasslands

Abstract. In this study we investigated population dynamics of a perennial forb, Plantago media, in semi‐natural grasslands in southeastern Sweden. Plantago media is a rather common inhabitant of grasslands in this region, but it has been shown to experience dispersal limitations both among managed grassland sites and at potential dispersal routes along road verges. The demography of P. media was studied in 22 permanent plots at two sites over five years. A seed sowing experiment was also performed at each site. The life‐cycle of P. media includes seedlings, juveniles, small vegetative rosettes, large vegetative rosettes and flowering rosettes. The population growth rates (λ‐values) were negative for both populations, with one exception: the first year transition matrix at one site. The projected life span for individuals surviving from seedlings to flowering rosettes varied between 19.6 yr and 227.8 yr. Elasticity analysis showed that remaining in the large rosette stages, both vegetative and flowering, was the most important factor influencing population growth rate. LTRE analysis gave similar results, indicating that spatial and temporal variation have similar effects on the population growth rate. The expected time to extinction in populations with an initial size of between 100 and 1000 individuals varied between 60 and 200 yr. The seed sowing experiment showed that seedling emergence was enhanced by both seed addition and disturbance, suggesting that recruitment is limited by seed and microsite availability. Recruitment was not found to have much impact on population growth rate. However, long‐term population maintenance may depend on occasional small‐scale disturbances that enhance pulses of recruitment. Fragmented landscape is likely to effect plant populations including rare and endangered species as well as more common species which have limited dispersal mechanisms.     

Spatial data replacing temporal data in population viability analyses: An empirical investigation for plants

In conservation management, there is an urgent need for estimates of population viability and for knowledge of the contributions of different life-history stages to population growth rates. Collection of long-term demographic data from a study population is time-consuming and may considerably delay the start of proper management actions. We examined the possibility of replacing a long-term temporal data set (demographic data from several years within a population) with a short-term spatial data set (demographic data from different populations for the same subset of two continuous years) for stochastic estimates of population viability. Using matrix population models for ten perennial plant species, we found that the matrix elements of spatial data sets often deviated from those of temporal data sets and that matrix elements generally varied more spatially than temporally. The appropriateness of replacing temporal data with spatial data depended on the subset of years and populations used to estimate stochastic population growth rates (log λ_s). Still, the precision of log λ_s estimates measured as variation in the yearly change of logarithmic population size rarely differed significantly between the spatial and temporal data sets. Since a spatiotemporal comparison of matrix elements and their variation cannot be used to assess whether spatial and temporal data sets are interchangeable, we recommend further research on the topic.     

Population dynamics of Agrimonia eupatoria and Geum rivale,two perennial grassland species

The population dynamics of two polycarpic perennials,Agrimonia eupatoria and Geum rivale, characteristic ofsemi-natural grasslands in Scandinavia, were examined in south-eastern Sweden.The perennial forbs were studied for several years in two populations each,located in habitats characteristic for the species in the study area.Demographic transition probabilities varied significantly between populationsand among years for the species. Transition matrix modelling yielded-values (population growth rates) that ranged from 0.89 to 1.14 forA. eupatoria and from 0.94 to 1.04 for G.rivale. The elasticity analysis showed that stasis in the adultstageclasses contributed most to . The life-table response experimentanalysis produced similar results as the elasticity analysis, where stasistogether with progression in the adult stage classes made a large contributionto the observed spatial variation in . Simulations of expected time toextinction were in the order of centuries for the study populations. Seedlingrecruitment was enhanced by seed addition and small scale disturbance inpopulations. For the intermediately abundant and more patchily distributedA. eupatoria, a regional survey of local populations wasconducted in the study area which revealed that most populations wererelativelysmall (< 100 individuals) and restricted to grassland fragments in roadverges. Furthermore, an attempt was made to estimate fruit dispersal from localpopulations.     

Field evaluation of doubled haploid plants in the Apiaceae: dill (Anethum graveolens L.), caraway (Carum carvi L.), and fennel (Foeniculum vulgare Mill.)

The Apiaceae family includes vegetables, as well as herb and spice crops. Compared to major crops, there have been few breeding or genetic improvement programs for any of the Apiaceae, especially the herb and spice species. Haploidy technology can be used to develop uniform, true-breeding lines, as well as to accelerate breeding programs. Field trials of dill (Anethum graveolens L.), caraway (Carum carvi L.), and fennel (Foeniculum vulgare Mill.) doubled haploid (DH) lines were conducted over 2–5 cropping seasons. Several of the DH dill lines had desirable agronomic characteristics such as short uniform stature along with early maturity that would be useful for crop improvement. Seed yields and the essential oil content of the seed harvested from the best performing DH dill lines were either equal to or higher than the parental line. A DH annual caraway line was identified that produced higher seed yields than the industry standard. The main constituents of the essential oil for the DH lines of both dill and caraway were similar to the parental lines. Fennel DH lines exhibited differences in height but were too late in maturity for seed production under prairie conditions. The results indicate that not only were we able to generate DH lines that could be used in a crop improvement program, but we developed DH lines that could be used directly as cultivars as these lines performed better than the industry standard (parental line).     

Effect of reproductive modes and environmental heterogeneity in the population dynamics of a geographically widespread clonal desert cactus

The dynamics of plant populations in arid environments are largely affected by the unpredictable environmental conditions and are fine-tuned by biotic factors, such as modes of recruitment. A single species must cope with both spatial and temporal heterogeneity that trigger pulses of sexual and clonal establishment throughout its distributional range. We studied two populations of the clonal, purple prickly pear cactus, Opuntia macrocentra, in order to contrast the factors responsible for the population dynamics of a common, widely distributed species. The study sites were located in protected areas that correspond to extreme latitudinal locations for this species within the Chihuahuan Desert. We studied both populations for four consecutive years and determined the demographic consequences of environmental variability and the mode of reproduction using matrix population models, life table response experiments (LTREs), and loop and perturbation analyses. Although both populations seemed fairly stable (population growth rate, λ∼1), different demographic parameters and different life cycle routes were responsible for this stability in each population. In the southernmost population (MBR) LTRE and loop and elasticity analyses showed that stasis is the demographic process with the highest contributions to λ, followed by sexual reproduction, and clonal propagation contributed the least. The northern population (CR) had both higher elasticities and larger contributions of stasis, followed by clonal propagation and sexual recruitment. Loop analysis also showed that individuals in CR have more paths to complete a life cycle than those in MBR. As a consequence, each population differed in life history traits (e.g., size class structure, size at sexual maturity, and reproductive value). Numerical perturbation analyses showed a small effect of the seed bank on the λ of both populations, while the transition from seeds to seedlings had an important effect mainly in the northern population. Clonal propagation (higher survival and higher contributions to vital rates) seems to be more important for maintaining populations over long time periods than sexual reproduction.     

Urbanization promotes the loss of seasonal dynamics in the semi-natural grasslands of an East Asian megacity

The biodiversity of agricultural landscapes has been noticeably affected by rapid urbanization. Although many studies have examined species diversity per unit area (alpha diversity), knowledge about the patterns of species turnover (beta diversity) in urban areas remains limited. Furthermore, most beta diversity studies have focused on spatial heterogeneity; however, losses of temporal heterogeneity resulting from urbanization remain limited. In this study, we examined how urbanization is associated with decreases in the seasonal heterogeneity of species composition, which could be used as an indicator of the loss of seasonality by ecologists and policy makers aiming to conserve biodiversity. We investigated (1) changes in species richness based on seasonal averages (alpha diversity) and (2) the seasonal turnover of species composition (beta diversity) for flowering plants and butterflies along a rural-urban gradient in semi-natural grasslands. The response variables were alpha and beta diversity for flowering plants and butterflies, and the explanatory variables were urban areas within a 1-km radius of the center of each site. Increasing urban area caused both the seasonal alpha and beta diversity of flowering plants and butterflies to decline. These results supported the homogenization hypothesis for the seasonality of plants and butterflies in semi-natural grasslands of dominant urban areas in East Asia. Future studies should focus on investigating how urbanization is causing both declines in seasonality and changes in the spatial heterogeneity of species composition and associated biodiversity loss. Ecologists and policy makers should focus on developing strategies to halt the loss of temporal biological heterogeneity to maintain biodiversity.     

The influence of time since introduction on the population growth of introduced species and the consequences for management

Several processes likely act to change the demographic rates of introduced species over time, and this changing demography could influence the optimal management of invasive populations. Optimal management strategies should be derived based on the demography. However, we have a poor understanding of the degree to which the demography of introduced species changes following initial introduction. We used published matrix population models of introduced plant populations to test how population growth rate and elasticity change with time since introduction. We did not find a significant relationship between population growth rate and time since introduction. However, elasticity to stasis increased while elasticity to growth decreased with time since introduction. Broadly, as time since introduction progressed the elasticities of the introduced plant populations became more similar to those that have been reported for native species. These results suggest that the optimal management strategy should be derived incorporating elasticity through time, especially when the time scope of management is long or the available demographic data were obtained in the past.     

Low congruence between the diversity of Waxcap (Hygrocybe spp.) fungi and vascular plants in semi-natural grasslands

The diversity of fungi in semi-natural grasslands is poorly known, partly due to difficulties in species identification in the field but also because there are few specialists available. Diversity assessments of grassland fungi would be facilitated if a potential surrogate group for fungal diversity could be identified. The aim of this study was to assess whether plant diversity patterns in semi-natural grasslands are congruent with diversity patterns of Waxcap (Hygrocybe spp.) fungi. Waxcaps, together with several other groups of fungi (e.g. the genera Entoloma, Dermoloma, Camarophyllopsis, and the families Clavariaceae and Geoglossaceae) have habitat requirements similar to many plants typical for semi-natural grasslands and they are all threatened by ceased management and nutrient enrichment. Diversity data from 31 semi-natural grasslands in southern Sweden were used to examine if there is congruence in species richness, nestedness, β-diversity and species composition between plants and Hygrocybe fungi. Species richness of Hygrocybe was significantly positively correlated with plant richness, although the relationship was not strong (r²=0.14). Both plant and Hygrocybe species composition was significantly nested, i.e. species-poor sites contain a subset of species from species-rich sites, which suggests that rare species mostly occur in the species-rich sites. A species similarity analysis between the grassland sites showed that there is low overlap between species composition of plants and Hygrocybe, indicating that conservation decisions based solely on plants may not fulfil the requirements of the Hygrocybe species. The conclusion is that there is low congruence between plant and Hygrocybe species diversity, making plants a poor surrogate group for Hygrocybe fungi, and probably also for other grassland fungi.     

Habitat fragmentation and plant populations: is what we know demographically irrelevant?

Abstract: Habitat fragmentation is considered a leading cause of plant extinction, and matrix models provide a powerful set of tools with which to identifying mechanisms that influence population declines. We surveyed the ecological literature to determine what components of plant demography have been studied in fragmented habitats, and determined the elasticity values of the vital rates influenced by these components. We found that there is a major disparity between the ecological processes and stages of life history with large demographic impacts and the focus of empirical research on plants in fragmented habitats. While the growth and survivorship of large, established individuals have the highest elasticity values, the focus of empirical research has been on components of reproduction and seedling dynamics. We argue that elucidating the demographic mechanisms underlying population declines in fragmented habitats, and developing strategies for mitigating these declines, will be challenging without a greater focus on understanding how fragmentation alters adult plant growth and survivorship.     

Demography of the demosponge Amphimedon compressa: Evaluation of the importance of sexual versus asexual recruitment to its population dynamics

We used size-based population matrix models to describe the demography of the demosponge Amphimedon compressa. The relative importance of growth, survivorship, and recruitment to population growth (λ) was assessed by performing elasticity and life table response experiment (LTRE) analyses. We also evaluated the relative contribution of sexual and asexual recruitment to λ by analyzing four different scenarios: (1) the combined impact of sexual and asexual recruitment, (2) the impact of only sexual recruits, (3) the impact of only asexual fragmentation, and (4) the impact of no recruitment. Size-based transition matrices were parameterized with field data collected at two sites in the southwest coast of Puerto Rico: Media Luna West (MLW), a reef exposed to high water movement and at Las Pelotas (LP), a reef experiencing low water motion. Estimated λ of 0.8940 and 0.7973 at MLW and LP respectively suggest that both populations are declining. Elasticity analysis indicated that survivorship may be the most important contributor to λ at both sites. However, λ at MLW was influenced more by the survivorship of small individuals whereas survivorship of the large size-class contributed the most to λ at LP. LTRE analysis indicates that the difference in λ between sites was mostly due to difference in survivorship of small sponges. At both sites, λ decreased considerably when sexual recruitment was excluded from the model whereas the absence of asexual recruitment barely changed λ. Therefore, it is suggested that sexual recruitment plays a major role (in comparison to asexual fragmentation) in the population dynamics of this sponge at the studied sites. We conclude that spatial variability in water motion plays an important role in population dynamics of A. compressa by influencing survivorship patterns, including the relative contribution of asexual and sexual recruitment to population growth.     

Population dynamics of the epiphytic bromeliad Tillandsia butzii in cloud forest

Epiphytes are a major component of tropical montane cloud forests. Over-exploitation and forest loss and degradation affect remnant populations. In this study, we analysed the population dynamics of the epiphytic bromeliad Tillandsia butzii over a 2-y period in a tropical montane cloud forest fragment in southern Mexico. Matrix analysis revealed that the T. butzii population is likely to be stable at the study site. On average the λ value did not differ significantly from unity: λ (95% confidence interval) = 0.978 (0.936–1.001). λ was highly influenced by stasis, to a lesser extent by growth and only slightly by fecundity. Overall, adult plant stasis and phalanx growth habit played a fundamental role in population maintenance. T. butzii tolerance to xeric conditions may contribute to population stability in the studied region.     

Spatial variation in population dynamics of the sand-burrowing amphipod Haustorioides japonicus

Densities of sandy beach organisms along coastlines often vary considerably even within small local scales. In order to understand the demographic basis of density variations in shore animals, we examined the population dynamics of the amphipod Haustorioides japonicus Kamihira, which is an ideal subject because it lacks a planktonic dispersal stage in its life history. Quantitative sampling at three adjacent sandy beaches along the Sea of Japan coast revealed that the spring density, i.e., the density of overwintered individuals, was similar among the three beaches. However, the amount of recruitment of newly hatched juveniles from late spring to summer greatly differed among the beaches, resulting in large spatial variations in the summer density. Cohort analyses revealed that at all beaches, young-of-the-year (YOY) individuals reproduced and then died, and that individuals which recruited in late summer overwintered. Moreover, growth, fecundity, and survival schedules of overwintered and YOY individuals were very similar among beaches. These observations suggest that the difference in recruitment success, i.e., the survival of newly hatched juveniles, is the primary cause of the inter-population variation in the summer density. It is likely that variations in salinity and sandy sediment affect the recruitment success of newly hatched individuals. This study indicates that biological productivity may differ substantially even among adjacent sandy beaches, with important implications for beach management.     

Elasticity analysis as an important tool in evolutionary and population ecology

Elasticity analysis estimates the proportional change in the population growth rate for a proportional change in a vital rate (i.e. survival, growth or reproduction). It can be used to pinpoint those parts of an organism’s life history that should be the focus of management effort, or those that contribute most to fitness. Recent theoretical work has emphasized some limitations of the technique, has overcome other problems, and has shown that it is robust to some violations of its underlying assumptions. Thus, although care is needed, elasticity analysis is a simple first step in answering important questions in evolutionary and population ecology.     

The relative importance of sexual and clonal reproduction for population growth in the perennial herb Fragaria vesca

The relative importance of sexual and clonal reproduction for population growth in clonal plants is highly variable. Clonal reproduction is often more important than sexual reproduction but there is considerable interspecific variation and the importance of the two reproductive modes can change with environmental conditions. We carried out a demographic study on the woodland strawberry (Fragaria vesca), a widespread clonal herb, at 12 sites in Switzerland during 2 years. Study sites were selected in two different habitats, i.e., forest and forest edge. We used periodic matrix models to estimate annual population growth rates and carried out prospective analyses to identify life cycle components that influence population growth rates most. Retrospective analyses were applied to study how the two different habitats affected population dynamics. Furthermore, we tested whether trade-offs between sexual and clonal reproduction occurred. There were large differences in annual population growth rates between sites and large within-site differences between years. Results of the prospective analyses clearly indicate that clonal reproduction is the dominant reproductive pathway whereas sexual reproduction is rather insignificant for population growth. Compared to forest habitats, forest edge habitats had higher population growth rates in the first year but smaller growth rates in the second year. We attribute these differing habitat effects to different water availabilities during consecutive years. No trade-offs between sexual and clonal reproduction were found. In conclusion, population growth of F. vesca relies heavily on clonal reproduction. Furthermore, reproduction and survival rates of F. vesca depend highly on spatio-temporal variation of environmental conditions.     

破碎化生境中粗柄独尾草种群大小对繁殖特性的影响

景观破碎化是目前存在的一种普遍现象,导致物种生存的原有生境总面积减小,产生许多大小不一的斑块种群。不同斑块生境中植物种群的大小直接反映植物对环境的响应特征和适应对策,与植物繁殖有关的开花结实特性、种子形态及萌发特性的研究将有助于深入了解植物在不同生境中的生存策略。以粗柄独尾草(Eremurus inderiensis)为研究对象,以野外调查为基础,对比研究不同斑块粗柄独尾草种群的种群大小和生殖成功,揭示不同生境中粗柄独尾草的生态适应性。结果表明:(1)不同斑块生境中,种群大小差异较大,且较小的种群幼苗比例也较小,种群大小与种群幼苗比例呈显著相关;(2)较小的种群自然座果率、结籽率较低,种群间差异显著;(3)随着生境破碎化程度增加,种子千粒重,种子长、宽等形态特征呈减小趋势,但种翅占比增大;(4)在粗柄独尾草最适萌发温度下,随着生境破碎化程度增加,种子萌发率有下降趋势。针对人类干扰程度强的衰退型粗柄独尾草种群,亟需减少人为干扰,依据不同生境中的干扰因素及种群生存现状,制定科学与切实可行的保护和恢复策略。