Integral projection models perform better for small demographic data sets than matrix population models: a case study of two perennial herbs

1. Matrix population models are widely used to describe population dynamics, conduct population viability analyses and derive management recommendations for plant populations. For endangered or invasive species, management decisions are often based on small demographic data sets. Hence, there is a need for population models which accurately assess population performance from such small data sets.
2. We used demographic data on two perennial herbs with different life histories to compare the accuracy and precision of the traditional matrix population model and the recently developed integral projection model (IPM) in relation to the amount of data.
3. For large data sets both matrix models and IPMs produced identical estimates of population growth rate (λ). However, for small data sets containing fewer than 300 individuals, IPMs often produced smaller bias and variance for λ than matrix models despite different matrix structures and sampling techniques used to construct the matrix population models.
4. Synthesis and applications . Our results suggest that the smaller bias and variance of λ estimates make IPMs preferable to matrix population models for small demographic data sets with a few hundred individuals. These results are likely to be applicable to a wide range of herbaceous, perennial plant species where demographic fate can be modelled as a function of a continuous state variable such as size. We recommend the use of IPMs to assess population performance and management strategies particularly for endangered or invasive perennial herbs where little demographic data are available.     

3种高山植物的物候和种群分布格局在融雪梯度上的变化

在青藏高原东部的一个高山雪床,沿着融雪梯度分别设置早融、中间和晚融3个融雪部位,然后测定川西小黄菊（Pyrethrum tatsienense）、长叶火绒草（Leontopodium longifolium）和圆穗蓼（Polygonum macrophyllum）在3个融雪部位上的物候差异以及种群分布格局的变化。结果表明：从早融到晚融的梯度上,3个物种的物候期都不同程度地有所推迟。其中,开始生长的时间推迟12~14 d,始花期推迟6~8 d,盛花期推迟6 d左右,但同一种植物在不同的融雪部位上的衰老枯黄期趋于一致,这标志着在晚融部位同一植物的生长期要缩短。在种群层次上,长叶火绒草和圆穗蓼的分布格局随着融雪的推迟都发生了一定的变化,基本上表现为从早融部位的集群分布到中间或晚融部位的随机分布。川西小黄菊在各个融雪部位上都表现为集群分布,但集群的强度随融雪的推迟逐渐减弱。     

The effects of phenological mismatches on demography

Climate change is altering the phenology of species across the world, but what are the consequences of these phenological changes for the demography and population dynamics of species? Time-sensitive relationships, such as migration, breeding and predation, may be disrupted or altered, which may in turn alter the rates of reproduction and survival, leading some populations to decline and others to increase in abundance. However, finding evidence for disrupted relationships, or lack thereof, and their demographic effects, is difficult because the necessary detailed observational data are rare. Moreover, we do not know how sensitive species will generally be to phenological mismatches when they occur. Existing long-term studies provide preliminary data for analysing the phenology and demography of species in several locations. In many instances, though, observational protocols may need to be optimized to characterize timing-based multi-trophic interactions. As a basis for future research, we outline some of the key questions and approaches to improving our understanding of the relationships among phenology, demography and climate in a multi-trophic context. There are many challenges associated with this line of research, not the least of which is the need for detailed, long-term data on many organisms in a single system. However, we identify key questions that can be addressed with data that already exist and propose approaches that could guide future research.     

Predicting impacts of Arctic climate change: Past lessons and future challenges

General circulation models predict increases in temperature and precipitation in the Arctic as the result of increases in atmospheric carbon dioxide concentrations. Arctic ecosystems are strongly constrained by temperature, and may be expected to be markedly influenced by climate change. Perturbation experiments have been used to predict how Arctic ecosystems will respond to global climatic change, but these have often simulated individual perturbations (e.g. temperature alone) and have largely been confined to the short Arctic summer. The importance of interactions between global change variables (e.g. CO₂, temperature, precipitation) has rarely been examined, and much experimentation has been short-term. Similarly, very little experimentation has occurred in the winter when General circulation models predict the largest changes in climate will take place. Recent studies have clearly demonstrated that Arctic ecosystems are not dormant during the winter and thus much greater emphasis on experimentation during this period is essential to improve our understanding of how these ecosystems will respond to global change. This, combined with more long-term experimentation, direct observation of natural vegetation change (e.g. at the tundra/taiga boundary) and improvements in model predictions is necessary if we are to understand the future nature and extent of Arctic ecosystems in a changing climate.     

Warming and free-air CO2 enrichment alter demographics in four co-occurring grassland species

Species differ in their responses to global changes such as rising CO(2) and temperature, meaning that global changes are likely to change the structure of plant communities. Such alterations in community composition must be underlain by changes in the population dynamics of component species. Here, the impact of elevated CO(2) (550 micromol mol(-1)) and warming (+2 degrees C) on the population growth of four plant species important in Australian temperate grasslands is reported. Data collected from the Tasmanian free-air CO(2) enrichment (TasFACE) experiment between 2003 and 2006 were analysed using population matrix models. Population growth of Themeda triandra, a perennial C(4) grass, was largely unaffected by either factor but population growth of Austrodanthonia caespitosa, a perennial C(3) grass, was reduced substantially in elevated CO(2) plots. Warming and elevated CO(2) had antagonistic effects on population growth of two invasive weeds, Hypochaeris radicata and Leontodon taraxacoides, with warming causing population decline. Analysis of life cycle stages showed that seed production, seedling emergence and establishment were important factors in the responses of the species to global changes. These results show that the demographic approach is very useful in understanding the variable responses of plants to global changes and in elucidating the life cycle stages that are most responsive.     

Rapid response to habitat restoration by the perennial Primula veris as revealed by demographic monitoring

The demography of Primula veris, a typical species of the species-rich Mesobromion grasslands, was investigated at two contrasting habitats in Eastern Belgium. Both a forested site and a clear-cut parcel were part of formerly larger calcareous grassland areas. By monitoring the demographic response of the target species, this study attempts to clarify the differences in fecundity, growth and survival between a restored and degraded calcareous grassland site. A study of plant traits showed a decrease in number of flowers, inflorescence stalks and plant size under a closing tree canopy. We surveyed individuals in permanent plots between 1999 and 2001. At the forested site, a first sign of decline included lower proportions of flowering individuals and afterwards an increase in mortality was found. Alternatively, removing canopy resulted in an immediate flowering response and both increased growth and seedling recruitment the year after. At both sites, survival rates strongly depended on the initial state, location and year. Projection matrix analysis revealed large differences in modelled population growth rates between sites and years. Under closing canopy the species showed only slow population decline, but the decrease was larger in the last survey year when higher mortality affected the number of reproductive individuals. Transitions between stages with the highest impact on population growth rate were identified by elasticity analysis. If calcareous grassland is forested, survival of reproductive adults is predicted to be very important for conservation of P. veris populations. However, seedling recruitment needs to be raised to guarantee long-term persistence of the populations. On the other hand, clear-felling of a site for restoration of species-rich calcareous grasslands may result in a rapid recovery of certain species.     

高山林线形成机制及假说的探讨

高山林线作为森林向苔原过渡的敏感带,历来被生态学家视为生态脆弱区、外界干扰信号的放大器和全球变化重要的预警区.20世纪80年代开始,许多研究关注全球变化对高山林线的影响,这一阶段针对林线海拔高度、纬度位置之间的关系及对未来热环境的响应研究已成为众多学者研究的焦点之一.众多学者依据自身研究目的、不同树种出现上限的原因、局部上影响林木的因子间的差异,在区域上对高山林线的形成机制的提出了诸多假说.本文针对这些假说进行了综述和探讨,并就其中不足之处提出质疑,提出今后需要深入研究的几个方向.

Abstract：

As a sensitive transitional zone between forest and tundra, timberline has always been considered by ecologists to be an ecologically fragile zone, an amplifier of outside interference signals, and a global climate change early warning zone. Since the 1980s, many studies have been made on the effects of global climate change on alpine timberline, mainly addressed the re-lationships of the upper limits of timberline with altitude and latitude, and in particular, the re-sponses of timberline location to global warming. Several hypotheses were proposed to examine the timberline formation of various tree species affected by environmental factors. This paper sum-marized these hypotheses, and discussed some potential studies in the future.     

Warming, plant phenology and the spatial dimension of trophic mismatch for large herbivores

Temporal advancement of resource availability by warming in seasonal environments can reduce reproductive success of vertebrates if their own reproductive phenology does not also advance with warming. Indirect evidence from large-scale analyses suggests, however, that migratory vertebrates might compensate for this by tracking phenological variation across landscapes. Results from our two-year warming experiment combined with seven years of observations of plant phenology and offspring production by caribou (Rangifer tarandus) in Greenland, however, contradict evidence from large-scale analyses. At spatial scales relevant to the foraging horizon of individual herbivores, spatial variability in plant phenology was reduced--not increased--by both experimental and observed warming. Concurrently, offspring production by female caribou declined with reductions in spatial variability in plant phenology. By highlighting the spatial dimension of trophic mismatch, these results reveal heretofore unexpected adverse consequences of climatic warming for herbivore population ecology.     

Plant Demography and Habitat: A Comparative Approach

Abstract Progress in plant demography will depend upon being able to synthesize a large body of data and this requires a means of comparing populations between sites and species. We have employed a comparative technique using elasticity analysis of stage-projection matrices to partition the contributions of fecundity (F), survival (L) and growth (G) to the finite rate of increase λ. Ordination of populations of 77 perennial herbs and trees in G-L-F space has shown that species segregate in this triangular space according to their life history and habitat. In the present paper we use the correspondence between demographic parameters and habitat revealed by this method to predict how succession and a variety of environmental factors such as grazing and fire are likely to alter the demography of populations and ultimately to change the composition of communities.     

Population modelling of Gelidium sesquipedale (Rhodophyta, Gelidiales)

A matrix population model of Gelidium sesquipedale, a commercial agarophyte from the Northeast Atlantic, was developed based on demographic data obtained during two years in a commercial stand of Cape Espichel, Portugal. G. sesquipedale individuals were classified into categories such as life cycle phase, spores, juveniles and adult frond size, because the species vital rates, fecundity, fertility, survival, growth and breakage depend on them. We also exemplify the use of a user-friendly modelling software, Stella, to develop a structured-population model. This is the first time this software has been used to model the demography of seaweed populations. The Stella model developed here behaved very similarly to the matrix model, because of its particular construction, which causes the forcing functions to be discrete rather than continuous. The relative importance of spore recruitment and vegetative growth of new fronds in both population growth and population structure was investigated. Elasticity analysis suggests that vegetative recruitment is the most important demographic parameter controlling population growth together with survival and transitions between juveniles (1–6 cm fronds) and class 1 fronds (6–9 cm fronds). On the other hand, sexual reproduction may, by itself, efficiently control the relative proportion of gametophytes and tetrasporophytes in the population, even though its contribution to recruitment is extremely small. A 40% difference in the growth rates of gametophyte and tetrasporophyte submatrices resulted from natural differences in spore recruitment rates. This revised version was published online in June 2006 with corrections to the Cover Date.     

全球变暖与陆地生态系统研究中的野外增温装置

由于化石燃料燃烧和森林砍伐等人类活动引起的地球大气层中温室气体(主要是二氧化碳)的富集已导致全球平均温度在20世纪升高了0.6 ℃,并将在本世纪继续上升1.4~5.8 ℃。这种地质历史上前所未有的全球变暖将对陆地植物和生态系统产生深远影响,并通过全球碳循环的改变反馈于全球气候变化。作为全球变化生态学的主要研究方法之一,生态系统增温实验能够为生态模型提供参数估计和模型验证。然而由于在世界各地使用的增温装置不同,使得各个生态系统之间的结果比较和整合难以实施,增加了模型预测的不确定性。该文通过比较几种常见的野外增温装置在模拟全球变暖情形时的优缺点,指出利用不同增温装置进行全球变暖研究中应注意的一些问题;同时探讨了全球变暖控制实验研究中的一些关键性的科学问题。     

The contributions of age and sex to variation in common tern population growth rate

1. The decomposition of population growth rate into contributions from different demographic rates has many applications, ranging from evolutionary biology to conservation and management. Demographic rates with low variance may be pivotal for population persistence, but variable rates can have a dramatic influence on population growth rate. 2. In this study, the mean and variance in population growth rate (lambda) is decomposed into contributions from different ages and demographic rates using prospective and retrospective matrix analyses for male and female components of an increasing common tern (Sterna hirundo) population. 3. Three main results emerged: (1) subadult return was highly influential in prospective and retrospective analyses; (2) different age-classes made different contributions to variation in lambda: older age classes consistently produced offspring whereas young adults performed well only in high quality years; and (3) demographic rate covariation explained a significant proportion of variation in both sexes. A large contribution to lambda did not imply a large contribution to its variation. 4. This decomposition strengthens the argument that the relationship between variation in demographic rates and variation in lambda is complex. Understanding this relationship and its consequences for population persistence and evolutionary change demands closer examination of the lives, and deaths, of the individuals within populations within species.