Population decline in California spotted owls near their southern range boundary

Species worldwide have begun to shift their range boundaries in response to climate change and other anthropogenic causes, with population declines at the trailing edge of a species' range often foreshadowing future changes in core parts of the range. Therefore, we analyzed a 30-year (1991–2019) data set for the California spotted owl (Strix occidentalis occidentalis) near its southern range boundary in southern California, USA, that included the largest regional population (San Bernardino Mountains) to estimate trends in territory occupancy and reproduction. We then assessed how these demographic rates were affected by habitat, wildfire, fuel treatments, and climate. Mean occupancy declined from 0.82 to 0.39 during our study, whereas reproductive output showed no temporal trends ( young/occupied territory). Territory extinction (extirpation) rates were relatively low in territories with more large trees (≥50 cm dbh), and colonization increased strongly with large tree density for low-elevation territories within the shrub-woodland ecotype but not for higher-elevation territories within mixed-conifer forest. High-severity wildfire had an adverse effect on occupancy: territory extinction rates steadily increased with the amount of high-severity fire within an owl territory during the previous 10 years, while colonization declined to nearly zero when ≥40% of a territory burned at high-severity during the previous 10 years. The effects of high-severity fire were unlikely to be confounded with post-fire fuel treatments, which primarily consisted of the removal, burning, or scattering of brush and small trees and snags (<40.6 cm dbh) and affected much smaller areas than high-severity fire. Of the 40 territories that received fuel treatments within 10 years of a fire, only 3 of them had post-fire fuel treatments that affected >5% of the territory, whereas average area burned at high severity for all 40 territories was 17%. Fuel treatments intended to modify fire behavior and reduce the likelihood of large, high-severity fires led to increases in territory extinction and colonization such that their net effect on occupancy was minimal. Our simulations of occupancy dynamics indicated that high-severity fire accounted for 9.6% of the observed decline in occupancy, whereas fuel treatments effectively accounted for none of the decline. Spotted owl reproductive output was lower at territories where fuel treatments occurred, but low- to moderate-severity fire resulted in much larger, population-level reductions in reproductive output (141 fewer young) from 2006–2019 than treatments (19 fewer young). Thus, the benefits of fuel treatments that reduce fire occurrence and severity appear to outweigh potential short-term costs to spotted owls and their habitat. Because high-severity fire only explained a modest amount of the long-term occupancy decline and much of the decline occurred in the 1990s before large fires occurred, additional factors are likely adversely affecting the owl population and merit further study. Nevertheless, the large observed population decline, limited evidence of owl dispersal among mountain ranges in the southern California metapopulation, and negative effects of increasingly large and severe fire suggest that California spotted owls at their southern range boundary are vulnerable to extirpation. In an era of climate change, owls in the core part of the range will likely become increasingly susceptible to warmer temperatures and increased severe fire activity in the future. Thus, the restoration of historical, low-severity fire regimes through fuels management while maintaining large trees is important to improving owl persistence.     

Cell communication by periodic cyclic-AMP pulses.

At the surface of aggregating cells of the slime mould, Dictyostelium discoideum, two different sites interacting with extracellular cAMP are detectable: binding sites and cycl-nucleotide phosphodiesterase. Both sites are developmentally regulated. An adequate stimulus for the chemoreceptor system in D. discoideum is the change of cAMP concentration in time, rather than concentration per se: long-term binding of cAMP causes only short-term response. The system is, consequently, adapted to the recognition of pulses rather than to steady-state concentrations of cAMP. The ce,lls are, nevertheless, able to sense stationary spatial gradients and to respond to them by chemotactic orientation. The possibility is discussed that they do so by transforming spatial concentration changes into temporal ones, using extending pseudopods as sensors. The cAMP recognition system is part of a molecular network involved in the generation of spatio-temporal patterns of cellular activities. This system controls the periodic formation of chemotactic signals and their propagation from cell to cell. The phosphodiesterase limits the duration of the cAMP pulses and thus sharply separates the periods of signalling; the binding sites at the cell surface are supposed to be the chemoreceptors. The control of cellular activities via cAMP receptors can be studied with biochemical techniques with cell suspensions in which spatial inhomogeneities are suppressed by intense stirring, whereas the temporal aspect of the spatiotemporal pattern is preserved. Under these conditions it can be shown that the extracellular cAMP concentration changes periodically, and that the phase of the cellular oscillator can be shifted by external pulses of cAMP. It can also be shown that small cAMP pulses induce a high output of cAMP, which demonstrates signal amplification, a function necessary for a cellular relay system.     

Population cycles in freshwater fish

Year-class strength in fish populations frequently follows an erratic pattern. However, predictable cyclic fluctuations in recruitment have been reported in a number of instances. In fish in which a single age-class is responsible for all or most of the production of eggs, a cycle may be set up with a period equal to the time taken to reach maturity. Moreover, density-dependence may act through fecundity or via survival of eggs or fish subject to interactions with predators or competitors. This paper reviews examples of cyclic variation in year-class strength and discusses the range of underlying causes.     

Population cycles can maintain foraging polymorphism

The maintenance of genetic variability in morphological traits that affect fitness is poorly understood. We present a simple Mendelian model of genetic traits affecting foraging efficiency in grazing ungulates, based on a trade-off between rates of energy extraction at low versus high levels of plant abundance. The model suggests that variation in foraging efficiency could be maintained via lottery competition arising as a direct consequence of dynamically unstable interactions between consumers and their food resources. Lottery competition is a plausible mechanism for explaining wide variability in foraging efficiency, such as that documented in unstable Soay sheep populations on the St Kilda archipelago.     

Water pulses and biogeochemical cycles in arid and semiarid ecosystems

The episodic nature of water availability in arid and semiarid ecosystems has significant consequences on belowground carbon and nutrient cycling. Pulsed water events directly control belowground processes through soil wet-dry cycles. Rapid soil microbial response to incident moisture availability often results in almost instantaneous C and N mineralization, followed by shifts in C/N of microbially available substrate, and an offset in the balance between nutrient immobilization and mineralization. Nitrogen inputs from biological soil crusts are also highly sensitive to pulsed rain events, and nitrogen losses, particularly gaseous losses due to denitrification and nitrate leaching, are tightly linked to pulses of water availability. The magnitude of the effect of water pulses on carbon and nutrient pools, however, depends on the distribution of resource availability and soil organisms, both of which are strongly affected by the spatial and temporal heterogeneity of vegetation cover, topographic position and soil texture. The inverse texture hypothesis for net primary production in water-limited ecosystems suggests that coarse-textured soils have higher NPP than fine-textured soils in very arid zones due to reduced evaporative losses, while NPP is greater in fine-textured soils in higher rainfall ecosystems due to increased water-holding capacity. With respect to belowground processes, fine-textured soils tend to have higher water-holding capacity and labile C and N pools than coarse-textured soils, and often show a much greater flush of N mineralization. The result of the interaction of texture and pulsed rainfall events suggests a corollary hypothesis for nutrient turnover in arid and semiarid ecosystems with a linear increase of N mineralization in coarse-textured soils, but a saturating response for fine-textured soils due to the importance of soil C and N pools. Seasonal distribution of water pulses can lead to the accumulation of mineral N in the dry season, decoupling resource supply and microbial and plant demand, and resulting in increased losses via other pathways and reduction in overall soil nutrient pools. The asynchrony of resource availability, particularly nitrogen versus water due to pulsed water events, may be central to understanding the consequences for ecosystem nutrient retention and long-term effects on carbon and nutrient pools. Finally, global change effects due to changes in the nature and size of pulsed water events and increased asynchrony of water availability and growing season will likely have impacts on biogeochemical cycling in water-limited ecosystems.     

Population cycles in microtines: The senescence hypothesis

Summary The cause of population cycles in microtines (voles and lemmings) remains an enigma. I propose a new solution to this problem based on a crucial feature of microtine biology, shifts in age structure, that has been ignored until now. Empirical evidence indicates that age structure must shift markedly towards older animals during declines because of three characteristics of the previous peak year: a shortened breeding season, total replacement of the breeding population from peak to decline and density-dependent social inhibition of maturation of young. Declines become inevitable as populations composed of older animals survive and reproduce poorly because of the effects of senescence, possibly interacting with the experiences of peak density and I present both theoretical and empirical evidence for this hypothesis. Although a variety of physiological systems deteriorate with aging, I focus on a crucial one — the inability of older animals to effectively maintain homeostasis in the face of environmental challenges because of a progressive deterioration in the endocrine feedback mechanisms involved in the hippocampal—hypothalamic—pituitary—adrenal axis. Microtine populations will not exhibit cycles where age structure shifts are prevented owing to extrinsic factors such as intense predation. Six testable predictions are made that can falsify this hypothesis.     

Circannual cycles in golden-mantled ground squirrels: fall and spring cold pulses

Summary Endogenous circannual rhythms of male golden-mantled ground squirrels (Spermophilus lateralis) exposed to low ambient temperature (6 °C) at various times were compared to the rhythms of a control group kept in a warm room (21 °C) throughout. A 20-week pulse of cold in the spring delayed the subsequent peak body weight and molt offset. However, in the second year after the spring cold pulse, a delay in the rhythm was evident only using peak weight as a phase marker. A 20-week cold pulse in the fall resulted in an earlier peak body weight and earlier onset of pigmented scrotum and descended testes, but there was no evidence from subsequent years that rhythms had been phase advanced. A third of the animals kept continuously at 6 °C remained with pigmented scrotum, descended testes and low body weight for more than a year. Circannual periodicities of animals that continued to display rhythms in the cold room were not significantly longer than those of controls in the warm room. The results suggest that low temperature in the fall can alter the expression of circannual rhythms without much affecting their phasing, while low temperature in the spring produces phase delays which last longer but have not been proved to be persistent.Abbreviation PRC phase response curve     

Semiparametric regression for periodic longitudinal hormone data from multiple menstrual cycles

We consider semiparametric regression for periodic longitudinal data. Parametric fixed effects are used to model the covariate effects and a periodic nonparametric smooth function is used to model the time effect. The within-subject correlation is modeled using subject-specific random effects and a random stochastic process with a periodic variance function. We use maximum penalized likelihood to estimate the regression coefficients and the periodic nonparametric time function, whose estimator is shown to be a periodic cubic smoothing spline. We use restricted maximum likelihood to simultaneously estimate the smoothing parameter and the variance components. We show that all model parameters can be easily obtained by fitting a linear mixed model. A common problem in the analysis of longitudinal data is to compare the time profiles of two groups, e.g., between treatment and placebo. We develop a scaled chi-squared test for the equality of two nonparametric time functions. The proposed model and the test are illustrated by analyzing hormone data collected during two consecutive menstrual cycles and their performance is evaluated through simulations.     

Globally attracting attenuant versus resonant cycles in periodic compensatory Leslie models

We use a periodically forced density-dependent compensatory Leslie model to study the combined effects of environmental fluctuations and age-structure on pioneer populations. In constant environments, the models have globally attracting positive fixed points. However, with the advent of periodic forcing, the models have globally attracting cycles. We derive conditions under which the cycle is attenuant, resonant, and neither attenuant nor resonant. These results show that the response of age-structured populations to environmental fluctuations is a complex function of the compensatory mechanisms at different life-history stages, the fertile age classes and the period of the environment.     

Population cycles caused by overcompensating density-dependence in an annual plant

Summary We analyse demographic data from a seven-year study of the annual crucifer Erophila verna which showed two-year cycles of abundance in the field. The unusual behaviour of this population is shown to be the consequence of overcompensating density-dependence. Very local differences in germination succes between plots can account for observed differences in their population dynamics over the long term.     

Population cycles caused by selection by density dependent competitive interactions

Several animal species have cyclic population dynamics with phase-related cycles in life history traits such as body mass, reproductive rate, and pre-reproductive period. Although many mechanisms have been proposed there is no agreement on the cause of these cycles, and no population equation that deduces both the abundance and the life history cycles from basic ecological constraints has been formulated. Here I deduce a population dynamic equation from the selection pressure of density dependent competitive interactions in order to explain the cyclic dynamics in abundance and life history traits. The model can explain cycles by evolutionary changes in the genotype or by plastic responses in the phenotype. It treats the population dynamic growth rate as an initial condition, and its density independent fundament is Fisher’s (1930, The Genetical Theory of Natural Selection, Oxford: Clarendon) fundamental theorem of natural selection that predicts a hyper-geometrical increase in abundance. The predicted periods coincide with the cyclic dynamics of Lepidoptera, and the Calder hypothesis, which suggests that the period of population cycles is proportional to the 1/4 power of body mass, follows from first principles of the proposed density dependent ecology.     

Transmission cycles, host range, evolution and emergence of arboviral disease

Many pandemics have been attributed to the ability of some RNA viruses to change their host range to include humans. Here, we review the mechanisms of disease emergence that are related to the host-range specificity of selected mosquito-borne alphaviruses and flaviviruses. We discuss viruses of medical importance, including Venezuelan equine and Japanese encephalitis viruses, dengue viruses and West Nile viruses.     

A bipedal compliant walking model generates periodic gait cycles with realistic swing dynamics

A simple spring mechanics model can capture the dynamics of the center of mass (CoM) during human walking, which is coordinated by multiple joints. This simple spring model, however, only describes the CoM during the stance phase, and the mechanics involved in the bipedality of the human gait are limited. In this study, a bipedal spring walking model was proposed to demonstrate the dynamics of bipedal walking, including swing dynamics followed by the step-to-step transition. The model consists of two springs with different stiffnesses and rest lengths representing the stance leg and swing leg. One end of each spring has a foot mass, and the other end is attached to the body mass. To induce a forward swing that matches the gait phase, a torsional hip joint spring was introduced at each leg. To reflect the active knee flexion for foot clearance, the rest length of the swing leg was set shorter than that of the stance leg, generating a discrete elastic restoring force. The number of model parameters was reduced by introducing dependencies among stiffness parameters. The proposed model generates periodic gaits with dynamics-driven step-to-step transitions and realistic swing dynamics. While preserving the mimicry of the CoM and ground reaction force (GRF) data at various gait speeds, the proposed model emulated the kinematics of the swing leg. This result implies that the dynamics of human walking generated by the actuations of multiple body segments is describable by a simple spring mechanics.     

Morpho-physiology and maize grain yield under periodic soil flooding in successive selection cycles

This study was carried out to evaluate maize plants of different recurrent selection cycles of the variety (Zea mays L.—Saracura-BRS-4154) regarded to genetic gains, morphophysiology, and grain yield, achieved over the selection cycles under intermittent flooding of the soil. This variety has the capacity to survive and produce in temporarily flooded soils and was developed by the National Maize and Sorghum Research Center (EMBRAPA). The experiment was conducted in greenhouse by using ten alternating selection cycles (Cycle 1–18) and BR 107 a variety known for its susceptibility to flooding. The flooding initiated at six-leaf stage by applying 50 mm of water three times a week. At flowering, the following parameters were evaluated: rate of leaf photosynthesis, stomatal conductance, intercellular CO₂ concentration, transpiration rate, photosynthetically active leaf area, root porosity, relative chlorophyll content, grain yield, harvest index, ear length, and interval between male and female flowering. Yield as a function of root porosity and photosynthesis were also evaluated. An index was created in this study, in order to help the discussion of the characteristics evaluated, it was called “Relative Tolerance Value—RTV”, only gaseous exchange measurements was not included in this index. By the way, it was observed throughout the selection cycles an increase in all gaseous exchange parameters, being the cycle 18, the one which presented the greatest averages. RTV for leaf area showed the greatest values for cycles 7 and 18, whereas root porosity, chlorophyll relative content, and harvest index, the greater RTV values were found in cycles 17 and 18. The largest grain yield RTV was observed in cycle 7, followed by cycles 13, 15, and 18. Flooding resulted in longer Anthesis-Silking Interval, especially for the first cycles. At flooding condition, grain yield was strongly related to root porosity (R ² = 0.66). These results showed that the selection cycles of “Saracura” maize improved some morphophysiologic characters, which favor their survival in flooded environments, also resulting in higher productivity.     

Population cycles produced by delayed density dependence in an annual plant

In contrast to insect and animal populations, little attention has been directed to the study of cycles in plant populations. It has been argued on theoretical grounds that plants present stable dynamics. Nevertheless, there are examples where plant populations appear to exhibit oscillatory dynamics, but the oscillatory signal is variable and comes from very short time series data. Using a combination of time series, models, and empirical results, we present evidence of population cycles for Descurania sophia in a 16-year field experiment. Endogenous and exogenous causal mechanisms were studied to identify processes underlying this temporal dynamic. Our results show a 4-year cycle produced by delayed density dependence. We suggest that high nutrient levels might be responsible for the observed dynamics of D. sophia. Our results suggest that although plant population dynamics may be stabilized by direct density dependence, delayed density dependence could destabilize dynamics.     

Population cycles and kin selection in Red Grouse Lagopus lagopus scoticus

Populations of Red Grouse Lagopus lagopus scoticus often show cyclic fluctuations in numbers. There is good evidence from previous work that, each year, densities are limited by the territorial behaviour of cocks. Therefore, changes in territorial behaviour and associated territory size might cause changes in density from one year to the next. Demographic measurements and a population experiment were consistent with a model which explained cycles as a consequence of changes in behaviour resulting from kin selection: differential behaviour between kin and non-kin. The demographic measurements presented here covered 16 years and two cyclic-type fluctuations, and the experiment lasted 10 years. Long-term studies are essential if lengthy processes are to be understood reliably.