Climate change, humidity, and mortality in the United States

This paper estimates the effects of humidity and temperature on mortality rates in the United States (c. 1973-2002) in order to provide an insight into the potential health impacts of climate change. I find that humidity, like temperature, is an important determinant of mortality. Coupled with Hadley CM3 climate-change predictions, I project that mortality rates are likely to change little on the aggregate for the United States. However, distributional impacts matter: mortality rates are likely to decline in cold and dry areas, but increase in hot and humid areas. Further, accounting for humidity has important implications for evaluating these distributional effects.     

Thermal histories of brooding lobsters, <Emphasis Type="Italic">Homarus americanus</Emphasis>, in the Gulf of Maine

Although it is widely accepted that migration by ovigerous lobsters (Homarus americanus Milne Edwards) optimizes thermal conditions for embryonic development, temperatures experienced by freely moving lobsters have never been measured. The precise thermal histories of 30 ovigerous lobsters at large in the Gulf of Maine were recorded to compare thermal conditions experienced during a brooding season. Analysis of both the temperature and movement data revealed a clear difference between lobsters smaller and larger than the size when 50% of individuals are mature (carapace length 93 mm). Although small and large lobsters ultimately experienced a similar number of degree-days above 3.4°C (952.8 for small and 983.6 for large) from 25 September 2002 until 27 July 2003, large lobsters and their embryos experienced less extreme and less variable temperatures. They were exposed to more gradual cooling in the fall and more gradual warming in the spring. These data, which are the first to document the seasonal temperatures experienced by ovigerous lobsters, suggest that migrations do not necessarily increase the number of degree-days experienced by developing embryos, but do reduce the variation in their thermal regime.     

Seasonal activity and thermoregulation in Meriones unguiculatus: A gerbil's choice

Summary Mongolian gerbils were observed over seasonal changes in day length and temperature in an outdoor enclosure containing an extensive burrow system. Animals could choose between the stable environment of their burrow and the daily and seasonal flunctuations aboveground. Surgically implanted radio transmitters provided body temperature and location of animals.Gerbils avoided temperature extremes by emerging from their burrow when ambient temperature fell below burrow temperature in hot weather and rose above burrow temperature in cold weather. They delayed emergence in the winter until several hours after sunrise and emerged before or after sunset in summer (Fig. 1).Circadian changes in core-body temperature corresponded to increases and decreases in behavioral activity. Emergence from the burrow was predictable by an increase in body temperature, as a warm-up period preceded emergence (Fig. 3).We conclude that gerbils, when subjected to extreme temperatures in the wild, probably time their activity to correspond to preferred ambient temperatures and thus shift from diurnal activity in the winter to nocrurnal activity in the summer (Fig. 2). This desert rodent's exceptional thermoregulatory capacity adapts it for synchronizing its daily activity to temperature.     

Impact of minimum winter temperatures on the population dynamics of Dendroctonus frontalis.

Predicting population dynamics is a fundamental problem in applied ecology. Temperature is a potential driver of short-term population dynamics, and temperature data are widely available, but we generally lack validated models to predict dynamics based upon temperatures. A generalized approach involves estimating the temperatures experienced by a population, characterizing the demographic consequences of physiological responses to temperature, and testing for predicted effects on abundance. We employed this approach to test whether minimum winter temperatures are a meaningful driver of pestilence from Dendroctonus frontalis (the southern pine beetle) across the southeastern United States. A distance-weighted interpolation model provided good, spatially explicit, predictions of minimum winter air temperatures (a putative driver of beetle survival). A Newtonian heat transfer model with empirical cooling constants indicated that beetles within host trees are buffered from the lowest air temperatures by approximately 1-4 degrees C (depending on tree diameter and duration of cold bout). The life stage structure of beetles in the most northerly outbreak in recent times (New Jersey) were dominated by prepupae, which were more cold tolerant (by >3 degrees C) than other life stages. Analyses of beetle abundance data from 1987 to 2005 showed that minimum winter air temperature only explained 1.5% of the variance in interannual growth rates of beetle populations, indicating that it is but a weak driver of population dynamics in the southeastern United States as a whole. However, average population growth rate matched theoretical predictions of a process-based model of winter mortality from low temperatures; apparently our knowledge of population effects from winter temperatures is satisfactory, and may help to predict dynamics of northern populations, even while adding little to population predictions in southern forests. Recent episodes of D. frontalis outbreaks in northern forests may have been allowed by a warming trend from 1960 to 2004 of 3.3 degrees C in minimum winter air temperatures in the southeastern United States. Studies that combine climatic analyses, physiological experiments, and spatially replicated time series of population abundance can improve population predictions, contribute to a synthesis of population and physiological ecology, and aid in assessing the ecological consequences of climatic trends.     

Growth and mortality rates of the fan mussel <Emphasis Type="Italic">Pinna nobilis</Emphasis> in Lake Vouliagmeni (Korinthiakos Gulf,Greece): a generalized additive modelling approach

The temporal patterns and the effect of shell size and depth on growth and mortality rates of the endangered fan mussel Pinna nobilis were investigated in the marine Lake Vouliagmeni (Korinthiakos Gulf, Greece). A total of 160 individuals were tagged and monitored monthly for a period of 17 months. At each visit, the size of the tagged individuals (shell width, w) was measured in situ and recorded. Any mortality event was also recorded and attributed to natural causes or (illegal) fishing. Growth and mortality rates were modeled with generalized additive models, which are non-parametric flexible models that free the researcher from the limiting concept of a strict parametric shape. The use of GAMs allowed the exploration of shapes of growth and mortality response curves in relation to predictor variables and allowed the fitting of statistical models that better agree with ecological theory. Growth rates had a seasonal pattern, with an extended period of very slow growth between late autumn and early spring, i.e., during the cold season, another short period of slow growth during August (when water temperatures reached their maximum values exceeding 29°C), and a peak in growth rates during late spring–early summer, probably related to an optimum combination of temperature and food availability. Growth rates varied with shell size, with a peak at w ∼4.5 cm, followed by a sharp decline to an approximately constant level, with sizes ranging from 9 to 15 cm, and a further decline with larger sizes down to almost zero for w > 20 cm. Growth rates did not vary substantially with depth. Although P. nobilis is a protected species in the EU and its fishing is strictly prohibited, fishing mortality was very high in Lake Vouliagmeni (much greater than natural mortality), especially during the hot season when the lake was crowded by summer visitors. The fan mussels were poached exclusively by free-diving and due to the high turbidity of the lake’s water, fishing mortality was higher in shallow areas (and mostly for large individuals) and was practically zero at depths >9 m. Due to fishing mortality, a size segregation of P. nobilis was observed in the lake: large individuals were restricted to deeper areas, while young and small individuals were more abundant in shallow areas where there was preferential recruitment. Natural mortality was strikingly size dependent and P. nobilis suffered high natural mortality during the first year of life; the probability of death by natural causes quickly diminished as the fan mussels grew in size. No depth-related differences in natural mortality were found.     

<Emphasis Type="Italic">Oithona similis</Emphasis> in a high latitude ecosystem: abundance,distribution and temperature limitation of fecundity rates in a sac spawning copepod

In this study we report the abundance, fecundity and an index of the relative survival of Oithona similis (nauplii to copepodites) across a large latitudinal and temperature range within the Southern Ocean. The abundance of O. similis was strongly related to temperature and to depth-integrated (0–100 m) chlorophyll a, abundance increasing with increasing temperature (and therefore decreasing latitude) and Chl a. In situ egg production rates and fecundity per female were also significantly and positively related to temperature and Chl a. Egg hatch times rapidly lengthen as temperature decreases and in sac spawning species the next batch of eggs cannot be produced until the previous clutch hatch. Consequently, we predict that in O. similis, fecundity must decline rapidly at low temperatures, especially below 5°C. A model comparing maximum rates of fecundity with in situ data suggested production rates were strongly food limited across our study region. However, the relationship of in situ and maximum rates to temperature were similar, confirming the importance of temperature. Further, as time taken to develop from egg to adult also rapidly extends with declining temperature, it is increasingly unlikely that O. similis will be able to maintain its population against mortality. Our findings have broad implications for the cold temperature (and hence geographic) limits of O. similis, but also for the distribution of other sac spawning copepods and planktonic species generally.     

Experimental effect of cold,La Niña temperatures on the survival of reef fishes from Gorgona Island (eastern Pacific Ocean)

The eastern tropical Pacific (ETP) reefs are affected at irregular times by extremely cold temperatures that occur principally during La Niña events. The effects of these low temperatures on the survival of reef fishes were experimentally assessed by determining the critical thermal minimum (CTM) of 15 reef fish species from Gorgona Island (ETP), and comparing these CTMs with the records of temperature during past La Niña events. Among species, mean CTMs ranged from 10.8°C to 16.3°C, which were lower than the coldest temperature recorded during the last La Niña event (18°C during La Niña 1998-1999). However, the observed ranges of CTM for two species (Thalassoma lucassanum and Eucinostomus gracilis) extended above 18°C. These results suggest that most of the reef fishes we studied are physiologically tolerant to the cold temperatures encountered during La Niña, though decreases in at least two populations may be expected as a result of the mortality of less tolerant individuals. Although tolerant to cold temperatures, reef fish populations may still experience negative changes during La Niña, because other determinants in population maintenance (e.g. reproduction and recruitment) are more temperature sensitive. The effects of other cold phenomena on reef fish survival are also discussed herein.     

Background and temperature effects on Uca panacea color change

Organismal coloration is used for communication, camouflage, and thermoregulation. These functions of body coloration may impose conflicting demands upon color-changing organisms. Here, we examined interacting thermoregulatory and camouflage color change responses when fiddler crabs Uca panacea were subject to simultaneously changing temperatures (10, 25, 40 °C) and backgrounds (black, white). Crab coloration lightened on a white background and at high temperatures and darkened on a black background and at low temperatures, reflecting the camouflage and thermoregulatory functions of color change. Synergistic background and temperature treatments (i.e., hot/white or cold/black) induced strong color change responses. When temperature and background were in conflict (i.e., hot/black or cold/white), responses to background coloration constrained thermoregulatory color change such that carapace coloration did not change. Such conflicts are likely to be common in nature, especially in highly heterogeneous environments. Throughout the experiment, males remained lighter than females and showed a greater response to the hot/white treatment, driven by a stronger response to high temperatures. These differences may reflect the physiological, morphological, and behavioral differences associated with sexual selection and sexual dimorphism in this species.     

Life-history trade-offs influence disease in changing climates: strategies of an amphibian pathogen

Life-history trade-offs allow many animals to maintain reproductive fitness across a range of climatic conditions. When used by parasites and pathogens, these strategies may influence patterns of disease in changing climates. The chytrid fungus, Batrachochytrium dendrobatidis, is linked to global declines of amphibian populations. Short-term growth in culture is maximal at 17 degrees-25 degrees C. This has been used in an argument that global warming, which increases the time that amphibians spend at these temperatures in cloud-covered montane environments, has led to extinctions. Here we show that the amphibian chytrid responds to decreasing temperatures with trade-offs that increase fecundity as maturation rate slows and increase infectivity as growth decreases. At 17 degrees-25 degrees C, infectious zoospores encyst (settle and develop a cell wall) and develop into the zoospore-producing stage (zoosporangium) faster, while at 7 degrees-10 degrees C, greater numbers of zoospores are produced per zoosporangium; these remain infectious for a longer period of time. We modeled the population growth of B. dendrobatidis through time at various temperatures using delayed differential equations and observational data for four parameters: developmental rate of thalli, fecundity, rate of zoospore encystment, and rate of zoospore survival. From the models, it is clear that life-history trade-offs allow B. dendrobatidis to maintain a relatively high long-term growth rate at low temperatures, so that it maintains high fitness across a range of temperatures. When a seven-day cold shock is simulated, the outcome is intermediate between the two constant temperature regimes, and in culture, a sudden drop in temperature induces zoospore release. These trade-offs can be ecologically important for a variety of organisms with complex life histories, including pathogenic microorganisms. The effect of temperature on amphibian mortality will depend on the interaction between fungal growth and host immune function and will be modified by host ecology, behavior, and life history. These results demonstrate that B. dendrobatidis populations can grow at high rates across a broad range of environmental temperatures and help to explain why it is so successful in cold montane environments.     

近60年新疆库尔勒市的气候变化特点分析

根据库尔勒市气象局和研究区安装的自动气象站的气象资料,采用滑动平均、距平分析、线性回归等方法,分析库尔勒市近60年的气候变化的基本特征,结果表明：（1）库尔勒市平均温度有4暖3冷的冷暖变化阶段,该地区平均气温呈上升趋势,特别是20世纪90年代以后气温明显增加;近10年的气温变化较大,比前50年的平均值上升了1℃;平均气温存在着明显的线性升温趋势。（2）库尔勒市年降水量变化趋势不是很明显,有2多4少的干湿变化阶段。近10年的降水量与前50年的降水量相比较偏少了2．4mm;年降水量不存在明显的线性变化趋势。（3）库尔勒市平均极端最高最低气温呈上升趋势,60年来该地区最高气温上升率与南疆最高气温上升率一致。近10年的极端最高最低气温与前50年的分别上升0．36℃和1．2℃,最低气温的上升对年平均气温的升高贡献最大。（4）通过比较库尔勒市气象站提供的气象数据和自动气象站提供的数据可以看出,该区域一定程度上受到城市热岛效应和人为干扰,因此需要在城市周边的某些地区安装气象仪器,以提高气象数据的精确性。     

Firms' and states’ responses to laxer environmental standards

On June 1, 2017, President Trump announced the United States' withdrawal from the Paris agreement on climate change. Despite this decision, American firms continued investing in low-carbon technologies and some states committed to tougher environmental standards. To understand this apparent paradox, this paper studies how a weakening of environmental standards affects the behavior of profit-maximizing firms. It finds that a relaxation of emission standards (i) may increase firms’ incentives to adopt clean technologies, but not to pollute less; (ii) may negatively affect industry profitability if it is perceived as temporary; and, when this is the case, (iii) the unilateral adoption of stricter standards by large states may increase the expected profitability of every firm.     

The main and added effects of heat on mortality in 33 Chinese cities from 2007 to 2013

● The main and added effect from heat co-existed in China. ● Both of the main and added effect could increase the mortality risk of population. ● Females, the elderly, the less educated and inland residents were more vulnerable. Increases in ambient temperatures and the frequency of extreme heat events constitute important burdens on global public health. However, evidence on their effects on public health is limited and inconclusive in China. In this study, data on daily deaths recorded in 33 Chinese cities from 2007 to 2013 was used to evaluate the effect of heat on mortality in China. In addition to the definition of a heatwave established by the China Meteorological Administration, we combined four city-specific relative thresholds (90th, 92.5th, 95th, and 97.5th percentiles) of the daily mean temperature during the study period and three durations of ≥ 2, ≥ 3, and ≥ 4 days, from which 13 heatwave definitions were developed. Then, we estimated the main and added effects of heat at the city level using a quasi-Poisson generalized additive model combined with a distributed lag nonlinear model. Next, the estimates for the effects were pooled at the national level using a multivariable meta-analysis. Subgroup analysis was performed according to sex, age, educational attainment, and spatially stratified heterogeneity. The results showed that the mortality risk increased from 22.3% to 37.1% due to the effects of the different heatwave definitions. The added effects were much lower, with the highest increase of 3.9% (95% CI: 1.7%–6.1%) in mortality risk. Females, the elderly, populations with low educational levels, and populations living inland in China were found to be the most vulnerable to the detrimental effects of heat. These findings have important implications for the improvement of early warning systems for heatwaves.     

Effects of temperature and precipitation on grassland bird nesting success as mediated by patch size

Grassland birds are declining faster than any other bird guild across North America. Shrinking ranges and population declines are attributed to widespread habitat loss and increasingly fragmented landscapes of agriculture and other land uses that are misaligned with grassland bird conservation. Concurrent with habitat loss and degradation, temperate grasslands have been disproportionally affected by climate change relative to most other terrestrial biomes. Distributions of grassland birds often correlate with gradients in climate, but few researchers have explored the consequences of weather on the demography of grassland birds inhabiting a range of grassland fragments. To do so, we modeled the effects of temperature and precipitation on nesting success rates of 12 grassland bird species inhabiting a range of grassland patches across North America (21,000 nests from 81 individual studies). Higher amounts of precipitation in the preceding year were associated with higher nesting success, but wetter conditions during the active breeding season reduced nesting success. Extremely cold or hot conditions during the early breeding season were associated with lower rates of nesting success. The direct and indirect influence of temperature and precipitation on nesting success was moderated by grassland patch size. The positive effects of precipitation in the preceding year on nesting success were strongest in relatively small grassland patches and had little effect in large patches. Conversely, warm temperatures reduced nesting success in small grassland patches but increased nesting success in large patches. Mechanisms underlying these differences may be patch‐size‐induced variation in microclimates and predator activity. Although the exact cause is unclear, large grassland patches, the most common metric of grassland conservation, appears to moderate the effects of weather on grassland‐bird demography and could be an effective component of climate‐change adaptation.     

Relationships between effects of environmental temperature and seston on growth and mortality of Mytilus edulis in a temperate northern estuary

The growth and mortality of experimentally-rafted Mytilus edulis L. of known age at 7 locations in a northern estuary (Damariscotta River, Maine, USA) were related to environmental temperatures and to the presence or absence of various potential food sources. All particles were regarded as potential food substrates. Growth decreased appreciably at sites where water temperatures exceeded 20°C, but mussels survived a wide range of elevated temperature exposures, ranging from 0 to 149 degree-days in excess of 20°C. The maximum temperature was 25°C. Mortalities of mussels at all sites but did affect the extent of mortality, which increased abruptly in late summer, when water temperatures were declining. Differences in degree-days of exposure to elevated temperatures did not influence the timing of mortality, which occurred synchronously at all times, but did affect the extent of mortality, which ranged from 35 to 90%. The period of high mortality was preceded by a rapid decline in phytoplankton standing crop. Total particle concentrations decreased during this period, but the shift toward larger particles suggests that there was little, if any, decrease in total volume of material in the seston, at least through August. It is suggested that living phytoplankton provided the critical energy source for these mussels, and that the mortalities were caused by rapidly reduced ration at a time of metabolic stress. The role of temperature and the possible role of the gametogenic cycle are discussed. In addition, the feeding efficiency of M. edulis may have decreased during this period due to an increase in mean particle diameter. Chlorophyll was divided into a nannoplankton fraction (<-20 m) and a larger fraction (>20 m). A smaller size criterion for this distinction is proposed for future studies. The importance of nannoplankton to the primary production of this estuary and the role of nannoplankton and nannoplankton-sized particles in the diet of mussels in nature are discussed.     

How temperature and habitat quality affect parasitoid lifetime reproductive success—A simulation study

Adult parasitoid females lay their eggs in or on host insects. Most species are incapable of de novo lipogenesis as adults, and lipids accumulated during the larval stage are allocated either to egg production or to adult survival. Lipid consumption increases with distance covered by the parasitoids and thus with the distance between available hosts within a habitat. Temperature should affect parasitoid fitness because it changes the constraint imposed by a limited reserve of lipids and because it influences behaviour. Climate change involves both an increase in average temperature and an increased frequency of extreme weather such as heat waves. We investigated how the predicted increase of temperature will affect parasitoid fitness and how this depends on habitat parameters (spatial distribution of hosts and lipid cost of habitat exploitation). We studied optimal behaviour and calculated fitness at different temperatures and in different habitats using a stochastic dynamic programming model for pro-ovigenic parasitoids (which mature all their eggs before becoming adult). We show that an increase in temperature decreases fitness of parasitoids adapted to lower temperatures. This decrease in fitness depends on habitat quality. In field conditions (assuming small costs of intra-patch foraging), the loss of fitness should be larger in habitats with high inter-patch distance and in habitats with a more aggregated distribution of hosts. The foraging behaviour of parasitoids is also affected; at higher temperature we show that intra-patch foraging becomes less efficient, and patch residence times are longer.     

Episodic death across species of desert shrubs

Extreme events shape population and community trajectories. We report episodic mortality across common species of thousands of long-lived perennials individually tagged and monitored for 20 years in the Colorado Desert of California following severe regional drought. Demographic records from 1984 to 2004 show 15 years of virtual stasis in populations of adult shrubs and cacti, punctuated by a 55-100% die-off of six of the seven most common perennial species. In this episode, adults that experienced reduced growth in a lesser drought during 1984-1989 failed to survive the drought of 2002. The significance of this event is potentially profound because population dynamics of long-lived plants can be far more strongly affected by deaths of adults, which in deserts potentially live for centuries, than by seedling births or deaths. Differential mortality and rates of recovery during and after extreme climatic events quite likely determine the species composition of plant and associated animal communities for at least decades. The die-off recorded in this closely monitored community provides a unique window into the mechanics of this process of species decline and replacement.     

Larval development of the crabCancer magister in temperature regimes simulating outer-coast and inland-water habitats

Postlarval megalopae of the Dungeness crabCancer magister inhabiting offshore coastal waters along the west toast of North America are larger and settle earlier than do those occupying the inland waters of the Puget Sound basin (Washington, USA, and British Columbia, Canada). The Puget Sound habitat is characterized by low initial temperatures that steadily increase during the course of zoeal development, while offshore temperatures are more moderate and stable. Larvae were raised in the laboratory from hatching to megalopa in three temperature treatments: (1) constant 10 °C (CO), (2) a regime that temporally simulated temperatures found off the central California coast during the larval season (CA); and (3) a regime simulating temperatures found within the Puget Sound basin (PS). Zoeal duration was 44% longer in the PS treatment than in the other two, at least partially accounting for observed differentes between outer-toast and inland-water settlement times. Although differentes were measured in megalopal weights and carapace lengths among treatments, results do not explain differences in megalopal size observed between outer-toast and inland-water individuals. Survival to megalopa was highest in the PS treatment, with the difference due to significantly lower mortality than in CO and CA treatments during the terminal zoeal stage. Daily instantaneous mortality rates were lower in the PS than in CO or CA treatments, indicating Chat extended larval duration will not necessarily result in reduced settlement success.     

Marine reserves reduce risk of climate-driven phase shift by reinstating size- and habitat-specific trophic interactions

Spatial closures in the marine environment are widely accepted as effective conservation and fisheries management tools. Given increasing human-derived stressors acting on marine ecosystems, the need for such effective action is urgently clear. Here we explore mechanisms underlying the utility of marine reserves to reinstate trophic dynamics and to increase resilience of kelp beds against climate-driven phase shift to sea urchin barrens on the rapidly warming Tasmanian east coast. Tethering and tagging experiments were used to examine size- and shelter-specific survival of the range-extending sea urchin Centrostephanus rodgersii (Diadematidae) translocated to reefs inside and outside no-take Tasmanian marine reserves. Results show that survival rates of C. rodgersii exposed on flat reef substratum by tethering were approximately seven times (small urchins 10.1 times; large urchins 6.1 times) lower on protected reef within marine reserve boundaries (high abundance of large predatory-capable lobsters) compared to fished reef (large predatory lobsters absent). When able to seek crevice shelter, tag-resighting models estimated that mortality rates of C. rodgersii were lower overall but remained 3.3 times (small urchins 2.1 times; large urchins 6.4 times) higher in the presence of large lobsters inside marine reserves, with higher survival of small urchins owing to greater access to crevices relative to large urchins. Indeed, shelter was 6.3 times and 3.1 times more important to survival of small and large urchins, respectively, on reserved relative to fished reef. Experimental results corroborate with surveys throughout the range extension region, showing greater occurrence of overgrazing on high-relief rocky habitats where shelter for C. rodgersii is readily available. This shows that ecosystem impacts mediated by range extension of such habitat-modifying organisms will be heterogeneous in space, and that marine systems with a more natural complement of large and thus functional predators, as achievable within no-take reserves, will minimize local risk of phase shifts by reinstating size and habitat-specific predator-prey dynamics eroded by fishing. Importantly, our findings also highlight the crucial need to account for the influence of size dynamics and habitat complexity on rates of key predator-prey interactions when managing expectations of ecosystem-level responses within marine reserve boundaries.     

Life-history analysis in “physiological” compared with “sidereal” time: An example with an amphipod (Gammarus lawrencianus) in a varying environment

The multivoltine, estuarine amphipodGammarus lawrencianus has four generations per year in an environment where temperatures range seasonally from –1° to 25°C. Temperature-response curves for rates of brood production and development were determined by laboratory experiments and field observation. The life history and population dynamics were observed over a full annual cycle (1981) for a field population located at Rocky Run, Porter's Lake, Nova Scotia, Canada. On a natural (i.e., sidereal) time scale, the generations appear to have very different life histories: the two summer generations have short lives, rapid development and mature at small size (classicr-selection), whereas the overwintering generations have relatively low rates of mortality, slow development and mature at large size (classicK-selection). This pattern (larger size at maturity at lower temperatures) is widespread in aquatic poikilotherms. Similar life-history differences are evident among cohorts of the summer generations that mature at different temperatures. When time is expressed on a physiological scale that removes the effect of temperature on embryonic development and reproductive rate, the variation within and among generations is greatly reduced. In particular, an apparent alternation betweenr- andK-selection largely disappears. Because the generations are temporally isolated, it might be surmised that natural selection acting on the summer generations might antagonize the effects of natural selection acting on the fall and winter generations. However, the scaling of the rates of development, maturation, growth, reproduction and mortality on the physiological time scale derived from the temperature dependence of development and reproductive rate gives a very different and more homogeneous pattern.