Comparative demography of two giant caulescent rosettes (Espeletia timotensis and E. spicata) from the high tropical Andes

Using field data from previous studies we built matrix models for two populations of giant rosettes, Espeletia timotensis Cuatrec. and E. spicata Sch. Bip. Wedd., from the Andes Cordillera in Mérida, Venezuela. We analysed the models and calculated population growth rate (λ), sensitivities, elasticities and the sensitivity of the elasticities to changes in the vital rates. The analysis showed that the two species behave alike in general demographic terms. In both models, population growth rate is positive and sensitivities of λ to changes in vital rates decrease markedly in this order: plant establishment, progression of juvenile–adult, germination and survival. The relative contributions of vital rates to λ (elasticities) are very similar to those of other woody plant species: a higher contribution of survival and a very low contribution of fecundity. Transition from seedling to juvenile is most important and the younger established stages (juveniles and young adults) play a predominant demographic role in both populations. Seed banks and older adults are playing a relatively minor role in the dynamics of both populations. However, they may be important in relation to unpredictable, favourable or detrimental events. Perturbation analysis of elasticities showed that increasing the rate of plant establishment will decrease the relative importance of stasis. We conclude that both species are demographically very close, and similar to other long‐lived woody plant species. However, the two species differ in the role of the seed bank, which seems more important in the demography of E. spicata than in E. timotensis.     

Better alone? A demographic case study of the hemiparasite Castilleja tenuiflora (Orobanchaceae): A first approximation

Castilleja tenuiflora is a facultative root hemiparasitic plant that has colonized a disturbed lava field in central Mexico. To determine the effects of hemiparasitism on the population dynamics of the parasite, we identified a set of potential hosts and quantified their effects on the vital rates of C. tenuiflora during 2016–2018. Connections between the roots of the hemiparasite and the hosts were confirmed with a scanning electron microscope. Annual matrices considering two conditions (with and without potential hosts) were built based on vital rates for each year, and annual stochastic finite rate growth rates (λ_s) were calculated. Plants produced more reproductive structures with hosts than without hosts. A Life Table Response Experiment (LTRE) was performed to compare the contributions of vital rates between conditions. We identified 19 species of potential hosts for this generalist hemiparasite. Stochastic lambda with hosts λ_s = 1.02 (CI = 0.9999, 1.1) tended to be higher than without them λ_s = 0.9503 (CI = 0.9055, 0.9981). The highest elasticity values correspond to survival. LTRE indicated that the most important parameters are survival and fecundity; the total contribution of fecundity (0.0192) to the difference in growth was three times lower than that of survival (0.0603). Piqueria trinervia was the most abundant host, and C. tenuiflora had a higher lambda with it than with other species. Individuals can grow alone, but hosts can have a positive effect on the vital parameters of C. tenuiflora and on λ.     

Pair-wise analyses of the effects of demographic processes, vital rates, and life stages on the spatiotemporal variation in the population dynamics of the riparian tree Aesculus turbinata Blume

Population growth rates (λ) of the riparian tree Aesculus turbinata varied from 0.9988 to 1.0524 spatiotemporally. We conducted a series of pair-wise demographic and matrix analyses, including randomization tests, three types of life table response experiments (LTREs), analysis of variance and χ² tests, to test which life stages had the greatest effect on this variation in λ. Randomization tests detected significant variations in λ between plots affected or not by typhoons in three habitats and between periods with high and low recruitment in one habitat. Mixed-level LTREs identified that the demographic processes and life stages that had the strongest effect on the actual variation in λ were: (1) progressions of small and intermediate juveniles and (2) founding process from seeds to 1-year-old seedlings. These juvenile stages had medium sensitivities and variances that explained high upper-level LTRE contributions. Lower-level LTREs showed that the vital rates contributing the most were the growth rates of these juvenile stages. These findings demonstrate that progression from one stage to the next, growth rates of 1-year-old seedlings, and stunted aging juveniles are the most important stages in the population dynamics of this long-lived primary tree species. Transition matrix elements with high elasticities had little effect on the variation in λ, indicating that high-elasticity vital rates do not necessarily drive variation in population growth. As compared with the results of randomization tests, significant differences in vital rates examined using ANOVA or χ² tests showed that typhoon disturbance had the greatest effect on the demographic parameters of individual trees.     

Remnant population dynamics in the facultative biennial Carum carvi in fragmented semi-natural grasslands

Transition matrix models were used to examine the population dynamics in the facultative biennial Carum carvi L. in semi-natural grasslands, specifically to assess what life cycle stages are important for population development and to evaluate the effects of environmental stochasticity on population persistence and, hence, the ability to develop remnant populations. The demographic studies were conducted over a 4-year period in three moderately grazed grasslands that differed in onset and duration of grazing. Experimental seed-sowing was also conducted in disturbed and undisturbed plots in the populations. Deterministic and stochastic models yielded overall negative population growth (λ < 1) for the populations. λ was sensitive to transitions in the most frequent vegetative stage classes. Elasticity analysis indicated that a large proportion of population growth could be ascribed to the stasis of individuals in the largest vegetative stage class. Life-table response experiment (LTRE) analyses showed also that progression to larger stage classes was important in explaining the between-population variation in λ. The expected time to extinction was on the order of several decades for the study populations. Seed-sowing indicated that seedling establishment was limited by both seed and micro-site availability. The populations of C. carvi seem to be able to persist for a rather long time in moderately grazed semi-natural grasslands, even in cases where populations are destined to become extinct. The results, thus, indicate that “biennials” are able to maintain remnant populations in managed semi-natural grasslands.     

Population dynamics and comparative demographics in sympatric populations of the round-leaved orchids Platanthera macrophylla and P. orbiculata

Orchids (Orchidaceae) are a family of flowering plants with a high proportion of threatened taxa making them an important focus of plant conservation. Orchid conservation efforts are most effective when informed by reliable demographic research. We utilized transition matrix models to examine the population dynamics and demography within sympatric populations of a species pair of terrestrial round-leaved orchids, Platanthera macrophylla and P. orbiculata. The models were parameterized from a large data set spanning 9 years from field observations of over 1,000 orchids. Life table response experiments (LTRE) were used to identify which life history transitions, and which vital rates within those transitions, most contributed to observed differences between the two species and most contributed to interannual variation within each species. Results from mean transition matrices projected finite rates of population growth that were not significantly different between the two species, with P. macrophylla near the replacement rate and P. orbiculata below it. LTRE revealed that the difference in population growth rates between the two species was mostly due to differences in fecundity (flowering adult to protocorm transition) driven by differences in fruit set and seed germination into protocorm, which were much greater for P. macrophylla. The two primary contributors to interannual variation in population growth rates for both orchids were adult survival and fruit set, respectively. These findings indicate that any environmental disturbances harming adult survival or fecundity will have a disproportionately negative effect on the orchid populations.     

Quantifying the links between land use and population growth rate in a declining farmland bird

Land use is likely to be a key driver of population dynamics of species inhabiting anthropogenic landscapes, such as farmlands. Understanding the relationships between land use and variation in population growth rates is therefore critical for the management of many farmland species. Using 24 years of data of a declining farmland bird in an integrated population model, we examined how spatiotemporal variation in land use (defined as habitats with “Short” and “Tall” ground vegetation during the breeding season) and habitat‐specific demographic parameters relates to variation in population growth taking into account individual movements between habitats. We also evaluated contributions to population growth using transient life table response experiments which gives information on contribution of past variation of parameters and real‐time elasticities which suggests future scenarios to change growth rates. LTRE analyses revealed a clear contribution of Short habitats to the annual variation in population growth rate that was mostly due to fledgling recruitment, whereas there was no evidence for a contribution of Tall habitats. Only 18% of the variation in population growth was explained by the modeled local demography, the remaining variation being explained by apparent immigration (i.e., the residual variation). We discuss potential biological and methodological reasons for high contributions of apparent immigration in open populations. In line with LTRE analysis, real‐time elasticity analysis revealed that demographic parameters linked to Short habitats had a stronger potential to influence population growth rate than those of Tall habitats. Most particularly, an increase of the proportion of Short sites occupied by Old breeders could have a distinct positive impact on population growth. High‐quality Short habitats such as grazed pastures have been declining in southern Sweden. Converting low‐quality to high‐quality habitats could therefore change the present negative population trend of this, and other species with similar habitat requirements.     

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.     

Demographic responses of Pinguicula ionantha to prescribed fire: a regression-design LTRE approach

This study describes the use of periodic matrix analysis and regression-design life table response experiments (LTRE) to investigate the effects of prescribed fire on demographic responses of Pinguicula ionantha, a federally listed plant endemic to the herb bog/savanna community in north Florida. Multi-state mark–recapture models with dead recoveries were used to estimate survival and transition probabilities for over 2,300 individuals in 12 populations of P. ionantha. These estimates were applied to parameterize matrix models used in further analyses. P. ionantha demographics were found to be strongly dependent on prescribed fire events. Periodic matrix models were used to evaluate season of burn (either growing or dormant season) for fire return intervals ranging from 1 to 20 years. Annual growing and biannual dormant season fires maximized population growth rates for this species. A regression design LTRE was used to evaluate the effect of number of days since last fire on population growth. Maximum population growth rates calculated using standard asymptotic analysis were realized shortly following a burn event (<2 years), and a regression design LTRE showed that short-term fire-mediated changes in vital rates translated into observed increases in population growth. The LTRE identified fecundity and individual growth as contributing most to increases in post-fire population growth. Our analyses found that the current four-year prescribed fire return intervals used at the study sites can be significantly shortened to increase the population growth rates of this rare species. Understanding the role of fire frequency and season in creating and maintaining appropriate habitat for this species may aid in the conservation of this and other rare herb bog/savanna inhabitants. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Stochastic demography and population dynamics in the red kangaroo Macropus rufus

1.  Many organisms inhabit strongly fluctuating environments but their demography and population dynamics are often analysed using deterministic models and elasticity analysis, where elasticity is defined as the proportional change in population growth rate caused by a proportional change in a vital rate. Deterministic analyses may not necessarily be informative because large variation in a vital rate with a small deterministic elasticity may affect the population growth rate more than a small change in a less variable vital rate having high deterministic elasticity.
2.  We analyse a stochastic environment model of the red kangaroo ( Macropus rufus ), a species inhabiting an environment characterized by unpredictable and highly variable rainfall, and calculate the elasticity of the stochastic growth rate with respect to the mean and variability in vital rates.
3.  Juvenile survival is the most variable vital rate but a proportional change in the mean adult survival rate has a much stronger effect on the stochastic growth rate.
4.  Even if changes in average rainfall have a larger impact on population growth rate, increased variability in rainfall may still be important also in long-lived species. The elasticity with respect to the standard deviation of rainfall is comparable to the mean elasticities of all vital rates but the survival in age class 3 because increased variation in rainfall affects both the mean and variability of vital rates.
5.  Red kangaroos are harvested and, under the current rainfall pattern, an annual harvest fraction of c . 20% would yield a stochastic growth rate about unity. However, if average rainfall drops by more than c . 10%, any level of harvesting may be unsustainable, emphasizing the need for integrating climate change predictions in population management and increase our understanding of how environmental stochasticity translates into population growth rate.     

Short-term population dynamics of Helianthemum caput-felis,a perennial Mediterranean coastal plant: a key element for an effective conservation programme

Plant population dynamics is fundamental to identify which parts of the life cycle are important when designing management actions. In this study, the short-term population dynamics of Helianthemum caput-felis (Cistaceae), a perennial plant distributed throughout the western Mediterranean Basin, was investigated by monitoring 98 permanent plots randomly placed along the overall distribution range for three years (2013–2015). Demographic surveys were analysed by Integral Projection Models (IPMs), and the analysis of population growth rates, elasticities, and life table response experiment (LTRE) analyses were performed. Under deterministic conditions, the H. caput-felis population showed a slight increase in the first censuses, but a general population decline followed. This trend was probably due to the shrinkage of plants in the last year, which correlated negatively with reproductive traits. Despite this decline, the population was at equilibrium under the simulated environmental stochasticity. The population dynamics of H. caput-felis followed the general pattern typical for long-lived Mediterranean plants: populations with growth rates closer to the equilibrium in the long term, determinant role of the survival-growth transitions for the population growth rate, slow growth and stasis, longevity of established individuals, shrinkage of medium-large plants, low recruitment rate, high generation times and mean long lifespans. The results of this study, which highlight how conservation measures can be focused on protecting and increasing the number of plants of medium-large size, provide important information for the planning management conservation actions for H. caput-felis and for several Mediterranean plants that show a similar life-history strategy.     

Habitat-Specific Population Growth of a Farmland Bird

Background

To assess population persistence of species living in heterogeneous landscapes, the effects of habitat on reproduction and survival have to be investigated.

Methodology/Principal Findings

We used a matrix population model to estimate habitat-specific population growth rates for a population of northern wheatears Oenanthe oenanthe breeding in farmland consisting of a mosaic of distinct habitat (land use) types. Based on extensive long-term data on reproduction and survival, habitats characterised by tall field layers (spring- and autumn-sown crop fields, ungrazed grasslands) displayed negative stochastic population growth rates (log λ_s: −0.332, −0.429, −0.168, respectively), that were markedly lower than growth rates of habitats characterised by permanently short field layers (pastures grazed by cattle or horses, and farmyards, log λ_s: −0.056, +0.081, −0.059). Although habitats differed with respect to reproductive performance, differences in habitat-specific population growth were largely due to differences in adult and first-year survival rates, as shown by a life table response experiment (LTRE).

Conclusions/Significance

Our results show that estimation of survival rates is important for realistic assessments of habitat quality. Results also indicate that grazed grasslands and farmyards may act as source habitats, whereas crop fields and ungrazed grasslands with tall field layers may act as sink habitats. We suggest that the strong decline of northern wheatears in Swedish farmland may be linked to the corresponding observed loss of high quality breeding habitat, i.e. grazed semi-natural grasslands.     

Sensitivity of the population growth rate to demographic variability within and between phases of the disturbance cycle

For species in disturbance-prone ecosystems, vital rates (survival, growth and reproduction) often vary both between and within phases of the cycle of disturbance and recovery; some of this variation is imposed by the environment, but some may represent adaptation of the life history to disturbance. Anthropogenic changes may amplify or impede these patterns of variation, and may have positive or negative effects on population growth. Using stochastic population projection matrix models, we develop stochastic elasticities (proportional derivatives of the long-run population growth rate) to gauge the population effects of three types of change in demographic variability (changes in within- and between-disturbance-phase variability and phase-specific changes). Computing these elasticities for five species of disturbance-influenced perennial plants, we pinpoint demographic rates that may reveal adaptation to disturbance, and we demonstrate that species may differ in their responses to different types of changes in demographic variability driven by climate change.     

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.     

Differential effects of abandonment on the demography of the grassland perennial Succisa pratensis

Abandonment of traditional land-use practices can have strong effects on the abundance of species occurring in agricultural landscapes. However, the precise mechanisms by which individual performance and population dynamics are affected are still poorly understood. To assess how abandonment affects population dynamics of Succisa pratensis we used data from a 4-year field study in both abandoned and traditionally grazed areas in moist and mesic habitats to parameterize integral projection models. Abandoned populations had a lower long-term stochastic population growth rate (λ _S = 0.90) than traditionally managed populations (λ _S = 1.08), while λ _S did not differ between habitat types. The effect of abandonment differed significantly between years and had opposed effects on different vital rates. Individuals in abandoned populations experienced higher mortality rates and lower seedling establishment, but had higher growth rates and produced more flower heads per plant. Population viability analyses, based on a population survey of the whole study area in combination with our demographic models, showed that 32 % of the populations face a high risk of extinction (>80 %) within 20 years. These results suggest that immediate changes in management are needed to avoid extinctions and further declines in population sizes. Stochastic elasticity analyses and stochastic life table response experiments indicated that management strategies would be most effective if they increase survival of small plants as well as seedling establishment, while maintaining a high seed production. This may be achieved by varying the grazing intensity between years or excluding grazers when plants are flowering.     

Climate change and grassland management interactively influence the population dynamics of Bromus erectus (Poaceae)

Climate and land management are important environmental drivers that affect the structure of terrestrial plant communities worldwide. Demographic studies allow a mechanistic understanding of the pathways in which environmental factors change population size. Climate and land management might interactively influence vital rates and growth rates of populations, however, demographic studies have not heretofore considered both factors in combination. We used the Global Change Experimental Facility as a platform to study the effect of climate (ambient climate conditions vs. future climate conditions) and land management (sheep grazing vs. mowing) on the demography of the common grass, Bromus erectus growing in semi-natural grassland communities. Using an integral projection model, we found positive population growth rates for B. erectus under all treatment combinations, and an interactive effect of climate and land management. Under ambient climate conditions, population growth of B. erectus was higher in mowed than grazed grassland plots, while population growth rates were similar across both management types under future climatic conditions. This interaction was primarily due to between-treatment changes in seedling recruitment, a vital rate to which the population growth rate is particularly sensitive. The interaction found in this study highlights the importance of considering multiple environmental drivers in demographic studies, to better predict future plant population dynamics and ultimately changes in community structure.     

Biennial cycling caused by demographic delays in a fire-adapted annual plant

We explored models explaining population cycling in the annual Warea carteri. We modeled the life cycle of W. carteri and compared projected trajectories to independently observed trajectories (up to 16 years) of plants in 74 patches in three populations. We built matrix models with an annual time step for two populations, including four stages, (recently produced seeds, seeds in the seed bank, seedlings, and adults) and five vital rates, summarized in seven transitions. Fluctuations of both observed and modeled populations were evaluated using power spectra, autocorrelation, amplitude, and damping. Observed populations had two point cycling. Observed amplitude was higher in frequently burned populations, reached its maximum 1 year after fire, and then dampened. Asymptotic transition and vital rate elasticities showed that seedling survival was the most important factor for long-term population growth, but transient elasticities showed that recruitment from the seed bank was important during the first years post-fire. Deterministic modeling and elasticity analyses indicated that delayed germination (for 1 year) may explain biennial population cycling. Stochastic models created similar cycling with slower damping than deterministic models, but still had lower amplitudes (especially 1–3 years post-fire) than observed populations. The biennial cycle in W. carteri is likely caused by the delay in seed germination, which creates two overlapping cohorts of plants, much like a strict biennial. Fire initiates the cycle by killing aboveground individuals and promoting seedling recruitment in the first post-fire year.     

The generality of management recommendations across populations of an invasive perennial herb

Demographic models are widely used to produce management recommendations for different species. For invasive plants, current management recommendations to control local population growth are often based on data from a limited number of populations per species, and the assumption of stable population structure (asymptotic dynamics). However, spatial variation in population dynamics and deviation from a stable structure may affect these recommendations, calling into question their generality across populations of an invasive species. Here, I focused on intraspecific variation in population dynamics and investigated management recommendations generated by demographic models across 37 populations of a short-lived, invasive perennial herb (Lupinus polyphyllus). Models that relied on the proportional perturbations of vital rates (asymptotic elasticities) indicated an essential role for plant survival in long-term population dynamics. The rank order of elasticities for different vital rates (survival, growth, retrogression, fecundity) varied little among the 37 study populations regardless of population status (increasing or declining asymptotically). Summed elasticities for fecundity increased, while summed elasticities for survival decreased with increasing long-term population growth rate. Transient dynamics differed from asymptotic dynamics, but were qualitatively similar among populations, that is, depending on the initial size structure, populations tended to either increase or decline in density more rapidly than predicted by asymptotic growth rate. These findings indicate that although populations are likely to exhibit transient dynamics, management recommendations based on asymptotic elasticities for vital rates might be to some extent generalised across established populations of a given short-lived invasive plant species.     

Accounting for stochasticity in demographic compensation along the elevational range of an alpine plant

Demographic compensation arises when vital rates change in opposite directions across populations, buffering the variation in population growth rates, and is a mechanism often invoked to explain the stability of species geographic ranges. However, studies on demographic compensation have disregarded the effects of temporal variation in vital rates and their temporal correlations, despite theoretical evidence that stochastic dynamics can affect population persistence in temporally varying environments. We carried out a seven‐year‐long demographic study on the perennial plant Arabis alpina (L.) across six populations encompassing most of its elevational range. We discovered demographic compensation in the form of negative correlations between the means of plant vital rates, but also between their temporal coefficients of variation, correlations and elasticities. Even if their contribution to demographic compensation was small, this highlights a previously overlooked, but potentially important, role of stochastic processes in stabilising population dynamics at range margins.     

Comparative demographic analysis in contrasting environments of Magnolia dealbata: an endangered species from Mexico

The study of population regulation and demography in natural habitats is critical for the conservation of rare and endangered species. We address the impact of cattle exclusion on the demographics of Magnolia dealbata (Magnoliaceae), an endangered species according to IUCN and Federal Mexican laws. Sixteen permanent plots were established, eight of which were enclosed to livestock, in the mountain cloud forest in Coyopolan, Mexico, which has the largest population of M. deadlbata. Censuses of the plots were undertaken annually during three annual cycles to record seed number, recruitment, mortality, and growth (defined as length and diameter at breast height). The effects of two treatments (with and without livestock exclusion) on the dynamics of M. dealbata were investigated using transition matrix models and life table response experiment (LTRE) analysis. Contrary to expectations, there was no significant effect of cattle exclusion on population growth rates (λ). Furthermore, the λ, estimated from the mean transition matrix for both treatments was greater than one. The transitions with the highest elasticity values were similar between the two treatments, while reproductive stage contributed more to differences in population growth rate and were less variable than the non-reproductive stage. LTRE analysis showed that treatment differences had little effect on λ. Livestock exclusion appears to lead more to differences in the arrangement of the values of the transition matrix than to the rate of population growth for M. dealbata.     

The many growth rates and elasticities of populations in random environments

Despite considerable interest in the dynamics of populations subject to temporally varying environments, alternate population growth rates and their sensitivities remain incompletely understood. For a Markovian environment, we compare and contrast the meanings of the stochastic growth rate (lambdaS), the growth rate of average population (lambdaM), the growth rate for average transition rates (lambdaA), and the growth rate of an aggregate represented by a megamatrix (shown here to equal lambdaM). We distinguish these growth rates by the averages that define them. We illustrate our results using data on an understory shrub in a hurricane-disturbed landscape, employing a range of hurricane frequencies. We demonstrate important differences among growth rates: lambdaS lambdaM. We show that stochastic elasticity, ESij, and megamatrix elasticity, EMij, describe a complex perturbation of both means and variances of rates by the same proportion. Megamatrix elasticities respond slightly and stochastic elasticities respond strongly to changing the frequency of disturbance in the habitat (in our example, the frequency of hurricanes). The elasticity EAij of lambdaA does not predict changes in the other elasticities. Because ES, although commonly utilized, is difficult to interpret, we introduce elasticities with a more direct interpretation: ESmu for perturbations of means and ESsigma for variances. We argue that a fundamental tool for studying selection pressures in varying environments is the response of growth rate to vital rates in all habitat states.