Using statistics to design and estimate vital rates in matrix population models for a perennial herb

Matrix population models are widely used to assess population status and to inform management decisions. Despite existing theories for building such models, model construction is often partially based on expert opinion. So far, model structure has received relatively little attention, although it may affect estimates of population dynamics. Here, we assessed the consequences of two published matrix structures (a 4 × 4 matrix based on expert opinion and a 10 × 10 matrix based on statistical modeling) for estimates of vital rates and stochastic population dynamics of the long-lived herb Astragalus scaphoides. We explored the ways in which choice of model structure alters the accuracy (i.e., mean) and precision (i.e., variance) of predicted population dynamics. We found that model structure had a negligible effect on the accuracy and precision of vital rates and stochastic stage distribution. However, the 10 × 10 matrix produced lower estimates of stochastic population growth rates than the 4 × 4 matrix, and more accurately predicted the observed trends in population abundance for three out of four study populations. Moreover, estimates of realized variation in population growth rate due to fluctuations in population stage structure over time were occasionally sensitive to matrix structure, suggesting differential roles of transient dynamics. Our study indicates that statistical modeling for choosing categories in matrix models might be preferable over expert opinion to accurately predict population trends and can provide a more objective way for model construction when the biological knowledge of the species is limited.     

Linking environmental and demographic data to predict future population viability of a perennial herb

Recent advances in stochastic demography provide tools to examine the importance of random and periodic variation in vital rates for population dynamics. In this study, we explore with simulations the effect of disturbance regime on population dynamics and viability. We collected 7 years of demographic data in three populations of the perennial herb Primula farinosa, and used these data to examine how variation in vital rates affected population viability parameters (stochastic growth rate, λ_S), and how vital rates were related to weather conditions. Elasticity analysis indicated that the stochastic growth rate was very sensitive to changes in regeneration, quantified as the production, survival, and germination of seeds. In one of the study years, all seedlings and mature plants in the demography plots died. This extinction coincided with the driest summer during the study period. Simulations suggested that a future increase in the frequency of high-mortality years due to climate change would result in reduced population growth rate, and an increased importance of survival in the seed bank for population viability. The results illustrate how the limited demographic data typically available for many natural systems can be used in simulation models to assess how environmental change will affect population viability.     

Recruitment rates exhibit high elasticity and high temporal variation in populations of a short-lived perennial herb

Empirical studies for different life histories have shown an inverse relationship between elasticity (i.e. the proportional contribution to population growth rate) and temporal variation in vital rates. It is accepted that this relationship indicates the effect of selective pressures in reducing variation in those life‐history traits with a major impact on fitness. In this paper, we sought to determine whether changes in environmental conditions affect the relationship between elasticity of vital rates and their temporal variation, and whether vital rates with simultaneously large elasticity and temporal variation might represent a characteristic life‐history strategy. We used demographic data on 13 populations of the short‐lived Hypericum cumulicola over 5–6 years, in three time‐since‐fire classes. For each population of each time‐since‐fire, we computed the mean matrix over years and its respective elasticity matrix, and the coefficients of variation in matrix entries over study years as an estimate of temporal variability. We found that mean elasticity negatively significantly correlated with temporal variation in vital rates in populations (overall eight out of 13) included in each time‐since‐fire. However, seedling recruitment exhibited both high elasticity and high temporal variation in almost all study populations. These results indicated that (1) the general relationship between elasticity and temporal variation in vital rates was not modified by environmental changes due to time‐since‐fire, and (2) high elasticity and high temporal variation in seedling recruitment in H. cumulicola is a particular trait of the species' life history. After seed survival in the soil seed bank, seedling recruitment represents the most important life‐history trait influencing H. cumulicola population growth rate (and fitness). The high temporal variability in seedling recruitment suggests that this trait is determined by environmental cues, leading to an increase in population size and subsequent replenishment of the seed bank in favorable years.     

Size-dependent gender modification in a hermaphroditic perennial herb

The size-advantage model predicts that hermaphroditic organisms adjust sex allocation depending on their resource status. We investigated the relationship between size and sex allocation in the co-sexual perennial herbs Trillium erectum and Trillium grandiflorum at two sites in southern Ontario, Canada by measuring pollen and ovule production and biomass allocation at flowering and fruiting. In both species, there was a strong relationship between size and gender; larger plants allocated proportionately more biomass to female reproduction and produced fewer pollen grains relative to ovules than smaller plants. Variation in gender was better explained by size than age, although age and size were correlated. While the relationship between size and gender was similar between species, T. erectum allocated proportionately more to female reproduction than T. grandiflorum, independent of size. In the absence of pollen limitation, there was no evidence of secondary adjustment of gender at fruiting. The results are discussed in the context of models predicting size-dependent gender modification in animal-pollinated plants. Evidence about the pollination and seed dispersal biology of Trillium spp. suggests that the relative effects of local mate and resource competition may be important in driving size-dependent sex allocation in these species.     

Negative evidence of local adaptation to the establishment conditions in a perennial herb

The differential adaptation of populations of the same species to their local environmental conditions through divergent selection, known as local adaptation, is a key step in the process of diversification of species. Here, we explore the local adaptation of the perennial mountain herb Helleborus foetidus to variable environmental conditions of seedling emergence and establishment at two different spatial scales (habitats and regions) with special attention to the role of physical and chemical soil properties. The possibility of local adaptation was evaluated under the ??local versus foreign?? and the ??home versus away?? criteria. Reciprocal sowing experiments were carried out by cross-sowing seeds among habitats and regions, controlling for maternal effects by means of seed mass, and recording seedling emergence and survival. Several topsoil properties were measured linked to each sowing point. Only partial patterns of local adaptation were found, which were insufficient to eventually state the existence of local adaptation at any spatial scale or under any criteria assessed. Here, we discuss how soil properties and selection on seed size may be related to the non-achievement of local adaptation criteria. Negative evidence of local adaptation seems to be due to a congruency in the selective pressures exerted by the different soil environments on seedling emergence and survival.     

Climate change and longterm patch dynamics of a perennial herb

Many perennial herbs are patchily distributed across the landscape. How such patch systems change over time is poorly known. What are the rates of patch establishment, extinction and area growth, and to what degree are changes synchronous and exhibiting trends? Can, for example, climate change signals be discerned? These questions are of interest also concerning the long-term population dynamics of the insects utilizing patchy plant resources. All patches of Vincetoxicum hirundinaria, a long-lived, patchily distributed herb, were mapped within a 12 km² area in SE Sweden. Patch areas were measured in 1977 and then almost yearly from 1990 to 2008.None of the original patches went extinct, but eleven new patches were established during the 32-year study period. However, the new patches combined represented only 0.6% of the total patch area in 2008. Average patch size did not change up to the early 1990s, but thereafter most patches grew slowly and steadily in size. This increase coincided with a climate shift towards longer growing seasons with warmer, sunnier and wetter summers. Larger patches and patches where plants had more flowers per shoot expanded faster.The patch system is presently not in equilibrium and this is probably the result of the new climatic regime. The number of patches increases steadily, most patches increase synchronously in size, and there is a positive feed back of patch size on rate of increase. Still, rates of increase are low, and in a three decade perspective the distribution of plant resources at the landscape scale remains essentially unchanged.     

Leaf damage decreases fitness and constrains phenotypic plasticity to drought of a perennial herb

Mediterranean-type ecosystems are increasingly prone to drought stress. Herbivory might limit plant functional responses to water shortage. This may occur as a result of plant resource depletion or due to the fact that leaf damage and drought may elicit opposite phenotypic responses. We evaluated the impact of herbivory on plant fitness in the field, and the effects of leaf damage on phenotypic plasticity to reduced soil moisture in a greenhouse. The study species was Convolvulus demissus, a perennial herb endemic to central Chile, which has a Mediterranean-type climate. Controlled herbivory by chrysomelid beetles (natural herbivores) in the field had a negative impact on plant fitness, estimated as number of fruits. Whereas reduced soil moisture alone did not affect seedling survival, damaged seedlings (simulated herbivory) had greater mortality when growing under water shortage. The hypothesis that herbivory would constrain phenotypic plasticity was supported by significant statistical interactions between leaf damage and soil moisture, followed by inspections of reaction norms. This was verified both overall (all phenotypic traits taken together, MANOVA) and in four of the six traits evaluated (ANOVAs). When plants were damaged, the reaction norms in response to low soil moisture of water use efficiency, root:shoot ratio and xylem water potential showed reduced slopes. While undamaged plants increased root biomass in response to low moisture, the opposite trend was found for damaged plants. The simultaneous occurrence of herbivory and drought events might curtail recruitment in plant populations of central Chile and other Mediterranean-type ecosystems due to the inability of damaged seedlings to show functional responses to low soil moisture. This finding is of ecological significance in view of current and projected trends of increased aridity in these ecosystems.     

Population dynamics of diploid and hexaploid populations of a perennial herb

BACKGROUND AND AIMS: Despite the recent enormous increase in the number of studies on polyploid species, no studies to date have explored the population dynamics of these taxa. It is thus not known whether the commonly reported differences in single life-history traits between taxa of different ploidy levels result in differences in population dynamics. METHODS: This study explores differences in single life-history traits and in the complete life cycle between populations of different ploidy levels and compares these differences with differences observed between different habitat types and years. Diploid and hexaploid populations of a perennial herb, Aster amellus, are used as the study system. Transition matrix models were used to describe the dynamics of the populations, and population growth rates, elasticity values and life-table response experiments were used to compare the dynamics between populations and years. KEY RESULTS: The results indicate that between-year variation in population dynamics is much larger than variation between different ploidy levels and different habitat conditions. Significant differences exist, however, in the structure of the transition matrices, indicating that the dynamics of the different ploidy levels are different. Strong differences in probability of extinction of local populations were also found, with hexaploid populations having higher probability than diploid populations, indicating strong potential differences in persistence of these populations. CONCLUSIONS: This is the first study on complete population dynamics of plants of different ploidy levels. This knowledge will help to understand the ability of new ploidy levels to spread into new areas and persist there, and the interactions of different ploidy levels in secondary contact zones. This knowledge will also contribute to understanding of interactions of different ploidy levels with other plant species or other interacting organisms such as pollinators or herbivores.     

Age-specific, density-dependent and environment-based mortality of a short-lived perennial herb

Density-independent and density-dependent processes affect plant mortality. Although less well understood, age-specific mortality can also play an important role in plant mortality. The goal of this study was to analyse several factors accounting for mortality in the Mediterranean short-lived perennial herb Lobularia maritima. We followed three cohorts of plants (from emergence to death) during 4 years in field conditions. We collected data on plant mortality of the effect of biotic agents (moth larvae and mycoplasma-like organisms, MLOs) and environmental variables. We also estimated density-dependent relationships affecting the fate of seedlings and adults. Results show that cohorts differed in their survival curves and ageing significantly increased mortality risk. Seedling mortality was density-dependent whereas adult mortality was not affected by density. MLO infection led to higher plant mortality whereas moth larvae attack did not affect plant mortality. In general, seedlings and adult plants experienced the highest mortality events in summer. We found, however, weak relationships between weather records and plant mortality. Age and size structures were not correlated. Overall, this study provides a comprehensive review of age-specific, density-dependent and density-independent factors that account for mortality of L. maritima plants throughout their life cycle in field conditions, highlighting the fact that age is an important factor in determining plant population dynamics.     

Simultaneous selection on vegetative and reproductive phenology in a perennial herb

The timing of different life‐history events is often correlated, and selection might only rarely be exerted independently on the timing of a single event. In plants, phenotypic selection has often been shown to favor earlier flowering. However, little is known about to what extent this selection acts directly versus indirectly via vegetative phenology, and if selection on the two traits is correlational. We estimated direct, indirect, and correlational phenotypic selection on vegetative and reproductive phenology over 3 years for flowering individuals of the perennial herb Lathyrus vernus. Direct selection favored earlier flowering and shorter timespans between leaf‐out and flowering in all years. However, early flowering was associated with early leaf‐out, and the direction of selection on leaf‐out day varied among years. As a result, selection on leaf‐out weakened selection for early flowering in one of the study years. We found no evidence of correlational selection. Our results highlight the importance of including temporally correlated traits when exploring selection on the phenology of seasonal events.     

Differential seed dispersal in Oxalis acetosella,a cleistogamous perennial herb

Explosive seed dispersal in the cleistogamous perennial forest herb Oxalis acetosella was studied during two growing seasons, to determine whether seeds derived from chasmogamous (CH) and cleistogamous (CL) flowers differ in dispersal distance. Seed dispersal distance, seed weight, and height of fruits were measured for both flower types, and the effects of phenology and year were also taken into account. The dispersal experiment was performed indoors, using plants transplanted from natural populations to pots. CL seeds were thrown significantly further than were CH seeds, though there was a considerable overlap in dispersal distances. There was also a significant positive relationship between seed weight and dispersal distance. No relationship was found between fruit height and dispersal distance. The results of this study contradict the common view that CL progeny should always be dispersed closer to the mother plant than CH progeny. The ecological implications of the dispersal difference are unclear, especially since it is uncertain whether CH seeds are generally outcrossed or not. Variation in dispersal distance in O. acetosella seems to be mainly dependent on a combination of reproductive mode and variation in seed weight.     

Development of microsatellite markers in a clonal perennial herb,Convallaria keiskei

We developed eight polymorphic microsatellite simple sequence repeat (SSR) loci from genomic DNA of a clonal perennial herb, Convallaria keiskei, using a dual‐suppression‐polymerase chain reaction (PCR) technique and an improved technique. These markers with four to 10 alleles per locus identified 29 genotypes in 82 samples collected from a population in Hokkaido, Japan. The observed and expected heterozygosities ranged from 0.241 to 0.862 and from 0.639 to 0.825, respectively. These SSR markers will be available to identify genets and evaluate genetic diversity of C. keiskei.     

On the dynamics of a population subject to slowly changing vital rates

Approximate closed-form expressions are given for the birth and death rates, the age-specific growth rates, and the age distribution of a population subject to slowly changing vital rates. The slower these rates have changed in the recent past, the better the approximations will be. These results, which are reminiscent of stable population theory, are tested by projecting the female population of the United States subject to realistically changing levels of fertility. The closed-form expressions yield quite accurate results and show that the dynamics of a population subject to slowly changing vital rates are to a large extent independent of the particular patterns of fertility. As in stable population theory, the asymptotic behavior of such a population is essentially determined by current survival rates and recent values of the intrinsic growth rate.     

Reduced maternal fecundity of the high Andean perennial herb

Assessments of the effects of invertebrate herbivores on high-altitude plants have seldom taken into account both mutualistic and antagonistic interactions. To evaluate the effect of herbivores (antagonists) and pollinators (mutualists) on the female reproductive success of the high-Andean perennial herb Alstroemeria umbellata, we separately and simultaneously excluded aphids (herbivores), and bees and bumblebees (pollinators) in a 2×2 factorial design. In flowers with pollinators excluded, aphids did not reduce seed set per flower (i.e., a direct effect). However, in flowers exposed to pollinators, aphids reduced seed set by 1.7 times (i.e., a pollinator- mediated indirect effect). Likewise, both types of animals exerted non-additive effects on maternal fecundity. These results suggest a modulating role for herbivores on the selection pressures exerted by pollinators on A. umbellata.     

Phenotypic variability along a climatic gradient in a perennial afrotropical rainforest understorey herb

Plants evolved in response to climatic conditions, which shaped their geographic distribution, functional traits and genetic composition. In the face of climatic changes, plants have to react by either genetic adaptation, phenotypic plasticity or geographic range shift. Their reaction potential depends on their phenotypic and genetic variability which can be evaluated through regional scale trait estimates, however, little is known here about tropical African plants. To start filling this gap of knowledge, the aim of this study was to estimate the phenotypic variability in a widespread perennial herb from the understorey of tropical African rainforests: Sarcophrynium prionogonium (Marantaceae). We surveyed 211 individuals from eight populations distributed across four sites in Cameroon covering largely the climatic range of the study species. Fourteen key functional traits were measured monthly for 18 months (2013–2014). Individuals of the study species persisted under a wide range of environmental conditions and there was considerable intraspecific variability within and across populations. Still, plant vegetative growth decreased with dryness. Productivity was positively related to a combination of high temperatures and precipitation and under these favourable conditions strongly shaped by light. Seasonal patterns of flower and fruit development were strongly associated with seasonal rainfall. Thus, the predicted increased dryness in tropical Africa might be disadvantageous for the study species. In the past, plants reacted to such aridification tendencies (e.g. during the Pleistocene glacial cycles) by retracting to moist refugia. The current climatic changes, however, being much faster and larger might provide new challenges.     

The effects of large herbivores on the landscape dynamics of a perennial herb

Background and Aims

Models assessing the prospects of plant species at the landscape level often focus primarily on the relationship between species dynamics and landscape structure. However, the short-term prospects of species with slow responses to landscape changes depend on the factors affecting local population dynamics. In this study it is hypothesized that large herbivores may be a major factor affecting the short-term prospects of slow-responding species in the European landscape, because large herbivores have increased in number in this region in recent decades and can strongly influence local population dynamics.

Methods

The impact of browsing by large herbivores was simulated on the landscape-level dynamics of the dry grassland perennial polycarpic herb Scorzonera hispanica. A dynamic, spatially explicit model was used that incorporated information on the location of patches suitable for S. hispanica, local population dynamics (matrices including the impact of large herbivores), initial population sizes and dispersal rate of the species. Simulations were performed relating to the prospects of S. hispanica over the next 30 years under different rates of herbivory (browsing intensity) and varying frequencies of population destruction (e.g. by human activity).

Key Results

Although a high rate of herbivory was detected in most populations of S. hispanica, current landscape-level dynamics of S. hispanica were approximately in equilibrium. A decline or increase of over 20 % in the herbivory rate promoted rapid expansion or decline of S. hispanica, respectively. This effect was much stronger in the presence of population destruction.

Conclusions

Browsing by large herbivores can have a dramatic effect on the landscape dynamics of plant species. Changes in the density of large herbivores and the probability of population destruction should be incorporated into models predicting species abundance and distribution.