Life after establishment: factors structuring the success of a mountain invader away from disturbed roadsides

Climate change and increased anthropogenic activity may both alter the current ranges of non-native plant species in mountainous areas, and could result in increased success of such species at higher elevations in the future. However, the course that management should take is often unclear due to a lack of information about the dynamics of how successful mountain invaders spread away from roadsides. The goals of this study were to determine if patterns of growth of a successful mountain invader, Linaria dalmatica (L.) Mill., (as measured by species cover) were: (1) similar to those of establishment (as measured by probability of occurrence), and (2) structured by the extant plant community. Study sites were established throughout the current elevation range of L. dalmatica in the Greater Yellowstone Ecosystem, and cover of the species was measured along with several vegetative community characteristics. Elevation influenced probability of occurrence (i.e., chance of establishment) for L. dalmatica, but not cover (which represents growth after establishment). L. dalmatica cover was negatively associated with several vegetative community characteristics which did not appear to be influenced by the presence of L. dalmatica. These results suggest that L. dalmatica establishment may be limited by climate, but that spread of established populations away from roadsides is most influenced by properties of the vegetative community. They further suggest that the resident vegetative communities structure the abundance of this invader, and that to limit spread of this species in mountainous areas, disturbance to the existing vegetative communities should be minimized.     

Temperature and host-plant effects on development and population growth of Mecinus janthinus (Coleoptera: Curculionidae), a biological control agent for invasive Linaria spp.

Mecinus janthinus Germar is a European stem-mining weevil that has been established in North America as a biological control agent against the invasive European weeds Linaria vulgaris P. Mill. and Linaria dalmatica (L.) P. Mill. (Scrophulariaceae). Establishment success and impact of the weevil have varied widely among sites. We investigated the hypothesis that some of this variation may be due to a lack of sufficient time for M. janthinus to develop to the adult (overwintering) stage in less favorable climates. Development time of M. janthinus was measured in L. vulgaris and L. dalmatica at four constant temperatures, and logistic regression was used to derive a model for the effect of temperature on development. Development rates were simulated using historic climate data for a site in central Alberta (where establishment was marginal on L. vulgaris) and one in southern British Columbia (where outbreaks occurred, resulting in heavy damage to L. dalmatica). The model showed that, on average, the British Columbia site had 50 more days available for the weevil to lay eggs that could reach the adult stage in time for overwintering than did the Alberta site. This may explain the more rapid population buildup at the British Columbia site. This model could be used to predict the climatic suitability of other areas for establishment of M. janthinus. An unexplained result was the very low survival rate of eggs laid in L. dalmatica under the same experimental conditions.     

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.     

Assessing seed and microsite limitation on population dynamics of a gypsophyte through experimental soil crust disturbance and seed addition

Understanding the factors limiting population growth is crucial for species management and conservation. We assessed the effects of seed and microsite limitation, along with climate variables, on Helianthemum squamatum, a gypsum soil specialist, in two sites in central Spain. We evaluated the effects of experimental seed addition and soil crust disturbance on H. squamatum vital rates (survival, growth and reproduction) across four years. We used this information to build integral projection models (IPMs) for each combination of management (seed addition or soil disturbance), site and year. We examined differences in population growth rate (λ) due to management using life table response experiments. Soil crust disturbance increased survival of mid to large size individuals and germination. Contributions to λ of positive individual growth (progression) and negative individual growth (retrogression) due to managements varied among years and sites. Soil crust disturbance increased λ in the site with the highest plant density, and seed addition had a moderate positive effect on λ in the site with lowest plant density. Population growth rate (λ) decreased by half in the driest year. Differences in management effects between sites may represent a shift from seed to microsite limitation at increasing densities. This shift underscores the importance of considering what factors limit population growth when selecting a management strategy.     

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.     

Variation in vital-rate sensitivity between populations of Texas horned lizards

Demographic studies of imperiled populations can aid managers in planning conservation actions. However, applicability of findings for a single population across a species’ range is sometimes questionable. We conducted long-term studies (8 and 9 years, respectively) of 2 populations of the lizard Phrynosoma cornutum separated by 1000 km within the historical distribution of the species. The sites were a 15-ha urban wildlife reserve on Tinker Air Force Base (TAFB) in central Oklahoma and a 6000-ha wildland site in southern Texas, the Chaparral Wildlife Management Area (CWMA). We predicted a trade-off between the effect of adult survival and fecundity on population growth rate (λ), leading to population-specific contributions of individual vital rates to λ and individualized strategies for conservation and management of this taxon. The CWMA population had lower adult survival and higher fecundity than TAFB. As predicted, there was a trade-off in the effects of adult survival and fecundity on λ between the two sites; fecundity affected λ more at CWMA than at TAFB. However, adult survival had the smallest effect on λ in both populations. We found that recruitment in P. cornutum most affected λ at both sites, with hatchling survival having the strongest influence on λ. Management strategies focusing on hatchling survival would strongly benefit both populations. As a consequence, within the constraint of the need to more accurately estimate hatchling survival, managers across the range of species such as P. cornutum could adopt similar management priorities with respect to stage classes, despite intra-population differences in population vital rates.     

Native species richness buffers invader impact in undisturbed but not disturbed grassland assemblages

Many systems are prone to both exotic plant invasion and frequent natural disturbances. Native species richness can buffer the effects of invasion or disturbance when imposed in isolation, but it is largely unknown whether richness provides substantial resistance against invader impact in the face of disturbance. We experimentally examined how disturbance (drought/burning) influenced the impact of three exotic invaders (Centaurea stoebe, Linaria dalmatica, or Potentilla recta) on native abundance across a gradient of species richness, using previously constructed grassland assemblages. We found that invaders had higher cover in experimentally disturbed plots than in undisturbed plots across all levels of native species richness. Although exotic species varied in cover, all three invaders had significant impacts on native cover in disturbed plots. Regardless of disturbance, however, invader cover diminished with increasing richness. Invader impacts on native cover also diminished at higher richness levels, but only in undisturbed plots. In disturbed plots, invaders strongly impacted native cover across all richness levels, as disturbance favoured invaders over native species. By examining these ecological processes concurrently, we found that disturbance exacerbated invader impacts on native abundance. Although diversity provided a buffering effect against invader impact without disturbance, the combination of invasion and disturbance markedly depressed native abundance, even in high richness assemblages.     

Linking vital rates to invasiveness of a perennial herb

Invaders generally show better individual performance than non-invaders and, therefore, vital rates (survival, growth, fecundity) could potentially be used to predict species invasiveness outside their native range. Comparative studies have usually correlated vital rates with the invasiveness status of species, while few studies have investigated them in relation to population growth rate. Here, I examined the influence of five vital rates (plant establishment, survival, growth, flowering probability, seed production) and their variability (across geographic regions, habitat types, population sizes and population densities) on population growth rate (λ) using data from 37 populations of an invasive, iteroparous herb (Lupinus polyphyllus) in a part of its invaded range in Finland. Variation in vital rates was often related to habitat type and population density. The performance of the populations varied from declining to rapidly increasing independently of habitat type, population size or population density, but differed between regions. The population growth rate increased linearly with plant establishment, and with the survival and growth of vegetative individuals, while the survival of flowering individuals and annual seed production were not related to λ. The vital rates responsible for rapid population growth varied among populations. These findings highlight the importance of both regional and local conditions to plant population dynamics, demonstrating that individual vital rates do not necessarily correlate with λ. Therefore, to understand the role of individual vital rates in a species ability to invade, it is necessary to quantify their effect on population growth rate.     

Microbially mediated CH4 consumption and N2O emission is affected by elevated CO2, soil water content, and composition of semi-arid grassland species

Elevated CO₂ affects plant productivity, but also water availability and plant species composition in semi-arid grasslands, thereby potentially causing complex effects on CH₄ consumption and N₂O emission. We studied the effects of atmospheric CO₂ concentration (400 vs 780 μL L^?1), water content (15 vs 20% gravimetric soil moisture), and composition of semi-arid grassland species (perennial grasses Bouteloua gracilis, Hesperostipa comata, and Pascopyrum smithii; sub-shrub Artemisia frigida; invasive forb Linaria dalmatica grown in monoculture and all five species together) on CH₄ consumption and N₂O emission in a full factorial greenhouse experiment. We used a unique method where we measured microbial effects on CH₄ consumption and N₂O emission in isolation from effects of gas diffusivity. Microbially mediated CH₄ consumption was significantly higher under elevated CO₂ (by 20%), but was not affected by soil water content or plant species composition. Microbially mediated N₂O emission was not significantly affected by elevated CO₂, but was significantly higher with high water content (by 67%) and differed significantly among species. Treatment effects on CH₄ consumption and N₂O emission often could not be explained simply by differences in soil moisture, suggesting that treatment-induced changes in other soil and microbial properties played a role in causing these effects.     

Response of population size to changing vital rates in random environments

Population size and population growth rate respond to changes in vital rates like survival and fertility. In deterministic environments change in population growth rate alone determines change in population size. In random environments, population size at any time t is a random variable so that change in population size obeys a probability distribution. We analytically show that, in a density-independent population, the proportional change in population size with respect to a small proportional change in a vital rate has an asymptotic normal distribution. Its mean grows linearly at a rate equal to the elasticity of the long-term stochastic growth rate λ _S while the standard deviation scales as $\sqrt t$ . Consequently, a vital rate with a larger elasticity of λ _S may produce a larger mean change in population size compared to one with a smaller elasticity of λ _S. But a given percentage change in population size may be more likely when the vital rate with smaller elasticity is perturbed. Hence, the response of population size to perturbation of a vital rate depends not only on the elasticity of the population growth rate but also on the variance in change in population size. Our results provide a formula to calculate the probability that population size changes by a given percentage that works well even for short time periods.     

Twenty‐five years after: post‐introduction association of Mecinus janthinus s.l. with invasive host toadflaxes Linaria vulgaris and Linaria dalmatica in North America

Linaria vulgaris, common or yellow toadflax, and Linaria dalmatica, Dalmatian toadflax (Plantaginaceae), are Eurasian perennial forbs invasive throughout temperate North America. These Linaria species have been the targets of classical biological control programmes in Canada and the USA since the 1960s. The first effective toadflax biological control agent, the stem‐mining weevil Mecinus janthinus (Coleoptera: Curculionidae) was introduced from Europe in the 1990s. This weevil has become established on L. dalmatica and L. vulgaris in both countries, although it has shown greater success in controlling the former toadflax species. Genetic and ecological studies of native range M. janthinus populations revealed that weevils previously identified as a single species in fact include two cryptic species, now recognised as M. janthinus, associated with yellow toadflax, and the recently confirmed species Mecinus janthiniformis, associated with Dalmatian toadflax. The results of a comprehensive study characterising haplotype identities, distributions and frequencies within M. janthinus s.l. native range source populations were compared to those populations currently established in the USA and Canada. The presence of both Mecinus species in North America was confirmed, and revealed with a few exceptions a high and consistent level of host fidelity throughout the adopted and native ranges. Genetic analysis based on mitochondrial cytochrome oxidase subunit II gene (mtCOII) defined the origin and records the subsequent North American establishment, by haplotype, of the European founder populations of M. janthinus (northern Switzerland and southern Germany) and M. janthiniformis (southern Macedonia), and provided population genetic indices for the studied populations. This analysis together with existing North American shipment receipt, release and rearing records elucidates probable redistribution routes and sources of both weevil species from initially released and established adopted range populations.     

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.     

Top-down and bottom-up controls on Dalmatian toadflax (<Emphasis Type="Italic">Linaria dalmatica</Emphasis>) performance along the Colorado Front Range,USA

Plant performance is influenced by both top-down (e.g., herbivores) and bottom-up (e.g., soil nutrients) controls. Research investigating the collective effects of such factors may provide important insight into the success and management of invasive plants. Through a combination of observational and experimental field studies, we examined top-down and bottom-up effects on the growth and reproduction of an invasive plant, Linaria dalmatica. First, we assessed attack levels and impacts of an introduced biocontrol agent, the stem-mining weevil Mecinus janthinus, on L. dalmatica plants across multiple years and sites. Then, we conducted a manipulative experiment to examine the effects of weevil attack, soil nitrogen availability, and interspecific competition on L. dalmatica. We found substantial variations in weevil attack within populations as well as across sites and years. Observational and experimental data showed that increased weevil attack was associated with a reduction in plant biomass and seed production, but only at the highest levels of attack. Nitrogen addition had a strong positive effect on plant performance, with a two-fold increase in biomass and seed production. Clipping neighboring vegetation resulted in no significant effects on L. dalmatica performance, suggesting that plants remained resource limited or continued to experienced belowground competitive effects. Overall, our research indicates that M. janthinus can exert top-down effects on L. dalmatica; however, weevil densities and attack rates observed in this study have not reached sufficient levels to yield effective control. Moreover, bottom-up controls, in particular, soil nitrogen availability, may have a large influence on the success and spread of this invasive plant.     

Variation in the demographics of a rare central Oregon endemic, Astragalus peckii Piper (Fabaceae), with fluctuating levels of herbivory

Understanding variation in plant vital rates (survival, growth, and reproduction) and population demographic parameters for rare plant taxa facilitates effective management for long-term persistence. We evaluated demographics of the rare plant Astragalus peckii (Fabaceae), a state-listed Threatened plant in Oregon, USA, with particular emphasis on how a microlepidopteran herbivore, Sparganothis tunicana, impacted vital rates and population growth. Stage-based transition matrix models were used to compute population growth rate (λ) and elasticity from 2006 to 2009 at two populations: Bull Flat, which was located in the main population center; and Chiloquin, a naturally isolated population. Population growth at Bull Flat was stable to slightly declining (λ = 0.96, 95 % CI 0.91–1.00) whereas at Chiloquin, the isolated population, population growth was increasing (λ = 1.20, 95 % CI 1.15–1.24). Microlepidopteran herbivory was associated with different plant responses in each population. At Bull Flat, plant survival was lower with greater herbivore presence. At Chiloquin, reproduction was reduced in plants when herbivores were active earlier in the growing season. Despite these effects on plant vital rates, we found lower population growth only during one transition period at Bull Flat when we compared matrices with and without herbivory. In addition to herbivory, we also address the potential role precipitation plays as a contributor to site differences and temporal variation within sites. Overall, we illustrate how two populations can have different responses to the same disturbance factor and highlight implications for management of different populations across the landscape.     

Population dynamics of the non-native freshwater gastropod,Cipangopaludina chinensis (Viviparidae): a capture-mark-recapture study

Capture-mark-recapture (CMR) is commonly used in conservation biology, but rarely used to study non-native species in freshwater habitats. The power of CMR lies in the ability to go beyond simple density estimates and to quantify invasion dynamics and vital population parameters. I applied CMR to a population of the non-native Chinese mystery snail (Cipangopaludina chinensis, Viviparidae) in a 1.46 ha pond on Long Island, NY to estimate population size and survival probability in the waterbody and to uncover potential mechanisms for enormous differences in introduction success within and between waterbodies (observed densities range <1–40 individuals m^?2). The C. chinensis population increased from approximately 150 to nearly 970 individuals from 2010 to 2012. Daily capture probabilities were low (<0.2) for snails of all sizes. Daily survival probabilities were size-dependent (almost 1.0 for snails larger than 30 mm shell length, and decreasing below that threshold), suggesting size-dependent mortality. This study highlights the ease of applying CMR to C. chinensis and its potential for other non-native species. Traditional survey methods such as density estimates with transects or quadrats cannot document increasing population sizes or size-specific mortality factors, which are essential for understanding introduction success and dynamics.     

Climate‐driven vital rates do not always mean climate‐driven population

Current climatic changes have increased the need to forecast population responses to climate variability. A common approach to address this question is through models that project current population state using the functional relationship between demographic rates and climatic variables. We argue that this approach can lead to erroneous conclusions when interpopulation dispersal is not considered. We found that immigration can release the population from climate‐driven trajectories even when local vital rates are climate dependent. We illustrated this using individual‐based data on a trans‐equatorial migratory seabird, the Scopoli's shearwater Calonectris diomedea, in which the variation of vital rates has been associated with large‐scale climatic indices. We compared the population annual growth rate λ_i, estimated using local climate‐driven parameters with ρ_i, a population growth rate directly estimated from individual information and that accounts for immigration. While λ_i varied as a function of climatic variables, reflecting the climate‐dependent parameters, ρ_i did not, indicating that dispersal decouples the relationship between population growth and climate variables from that between climatic variables and vital rates. Our results suggest caution when assessing demographic effects of climatic variability especially in open populations for very mobile organisms such as fish, marine mammals, bats, or birds. When a population model cannot be validated or it is not detailed enough, ignoring immigration might lead to misleading climate‐driven projections.     

Thermal tolerance affects mutualist attendance in an ant–plant protection mutualism

The influence of native fauna on non-native plant population growth, size, and distribution is not well documented. Previous studies have shown that native insects associated with tall thistle (Cirsium altissimum) also feed on the leaves, stems, and flower heads of the Eurasian congener C. vulgare, thus limiting individual plant performance. In this study, we tested the effects of insect herbivores on the population growth rate of C. vulgare. We experimentally initiated invasions by adding seeds at four unoccupied grassland sites in eastern Nebraska, USA, and recorded plant establishment, survival, and reproduction. Cumulative foliage and floral herbivory reduced C. vulgare seedling density, and prevented almost any reproduction by C. vulgare in half the sites. The matrix model we constructed showed that this herbivory resulted in a reduction of the asymptotic population growth rate (λ), from an 88 % annual increase to a 54 % annual decline. These results provide strong support for the hypothesis that indigenous herbivores limit population invasion of this non-native plant species into otherwise suitable grassland habitat.     

Performance of Forest Bryophytes with Different Geographical Distributions Transplanted across a Topographically Heterogeneous Landscape

Most species distribution models assume a close link between climatic conditions and species distributions. Yet, we know little about the link between species'' geographical distributions and the sensitivity of performance to local environmental factors. We studied the performance of three bryophyte species transplanted at south- and north-facing slopes in a boreal forest landscape in Sweden. At the same sites, we measured both air and ground temperature. We hypothesized that the two southerly distributed species Eurhynchium angustirete and Herzogiella seligeri perform better on south-facing slopes and in warm conditions, and that the northerly distributed species Barbilophozia lycopodioides perform better on north-facing slopes and in relatively cool conditions. The northern, but not the two southern species, showed the predicted relationship with slope aspect. However, the performance of one of the two southern species was still enhanced by warm temperatures. An important reason for the inconsistent results can be that microclimatic gradients across landscapes are complex and influenced by many climate-forcing factors. Therefore, comparing only north- and south-facing slopes might not capture the complexity of microclimatic gradients. Population growth rates and potential distributions are the integrated results of all vital rates. Still, the study of selected vital rates constitutes an important first step to understand the relationship between population growth rates and geographical distributions and is essential to better predict how climate change influences species distributions.     

The effects of mulching and abandonment on the viability of the perennial grassland species Plantago lanceolata

Abandonment and fragmentation are the primary causes of decline for many light-demanding grassland species in the modern agricultural landscape. In order to maintain a flora rich in such species, design of management practices based upon knowledge of life-history consequences following management shifts is essential. In this study, we investigated the demography of the perennial grassland species Plantago lanceolata for 4 years in one mulched site and two abandoned sites to compare how the two treatments influenced population dynamics. Compared to abandonment, mulching had a small but positive effect on the population growth rate (λ) of P. lanceolata. In both the mulched and the abandoned sites, growth and stasis among the largest plants contributed most strongly to λ, while the contribution of fecundity was negligible in both treatments. λ of P. lanceolata varied more between years than between treatments, most notably because λ values below unity were observed in the abandoned sites in a 1-year period that included a severe winter. The population of the mulched site consisted of larger ramets and had higher clonal growth rates than the populations of the abandoned sites. This explains the apparently higher resilience to unfavorable climatic conditions in the mulched site. We conclude that management by mulching is better than abandonment for the long-term persistence of P. lanceolata. However, due to low recruitment also in the mulched site, even mulching may be insufficient for P. lanceolata to persist over long time periods.     

Population dynamics in canopy gaps: nonlinear response to variable light regimes by an understory plant

Gap-dependent species are typically understood to have higher population growth rates (λs) when they are exposed to higher light transmittance. I investigated the relationship between both diffuse light and direct light transmittance and λ for the gap-dependent plant Trollius laxus using 5 years of data from 20 subpopulations (11 in created, experimental canopy gaps; 9 in intact canopy control areas). There was a nonlinear (unimodal) relationship between diffuse light and λ for T. laxus under the wide range of light levels encountered at the gap and control subpopulations [4–58 % diffuse photosynthetic photon flux density (PPFD)]. There was no relationship between direct light and λ. However, in the gaps, where light levels were generally greater than 20 % PPFD, both diffuse light and direct light had strong negative linear relationships with λ. Therefore, under wide-ranging light regimes, plant populations may show complicated, nonlinear responses to gap formation. Furthermore, gap-dependent plant populations may even decline in the brightest gaps. These results demonstrate that future studies on forest plant population dynamics should strive to include populations from a wide variety of light regimes, and avoid broadly categorizing light regimes as simply “gap” or “non-gap.”