Developmental rate isomorphy in insects and mites

When the proportion of total developmental time spent in a particular developmental stage does not change with temperature, an organism shows "rate isomorphy." This is the case only if the lower developmental threshold is the same for all developmental stages. In this study, the incidence of rate isomorphy in seven species of mites and 342 species from 11 insect orders (some represented by several populations) was determined. Whether a species shows rate isomorphy or not was determined over a range of temperatures where the relationship between the rate of development and temperature is linear. Proportion of total developmental time spent in a particular stage was plotted against temperature and the existence of rate isomorphy inferred from a zero change in proportion. Rate isomorphy was detected in 243 (57%) of 426 populations. In the rest of the cases, rate isomorphy was violated by deviations in the proportion of time spent in a stage by an average of 0.2% (range 4.5E-06% to 2.8%) at the mean of the range of temperatures of all the data sets (11 degrees C). The violations occurred most frequently at the extremes of the linear phase, which is attributed to methodical biases, mortality at low temperatures, or too coarse an estimate of developmental time at high temperatures. Similarly, a meta-analysis also revealed an overall prevalence of rate isomorphy. Consequently, in insect and mite species, all the developmental stages appear to have the same population-specific lower developmental threshold. The existence of rate isomorphy could be of great practical importance, for example, in the timing of life-history events and in determining preadult thermal requirements. There are also indications that it may act as a phylogenetic constraint.     

Linking life‐history theory and metabolic theory explains the offspring size‐temperature relationship

Temperature often affects maternal investment in offspring. Across and within species, mothers in colder environments generally produce larger offspring than mothers in warmer environments, but the underlying drivers of this relationship remain unresolved. We formally evaluated the ubiquity of the temperature–offspring size relationship and found strong support for a negative relationship across a wide variety of ectotherms. We then tested an explanation for this relationship that formally links life‐history and metabolic theories. We estimated the costs of development across temperatures using a series of laboratory experiments on model organisms, and a meta‐analysis across 72 species of ectotherms spanning five phyla. We found that both metabolic and developmental rates increase with temperature, but developmental rate is more temperature sensitive than metabolic rate, such that the overall costs of development decrease with temperature. Hence, within a species’ natural temperature range, development at relatively cooler temperatures requires mothers to produce larger, better provisioned offspring.     

How to compare the lower developmental thresholds

According to the hypothesis of isomorphy rate, all the lower developmental thresholds of different developmental stages of an insect are equal. However, there is lack of a formal statistical method for testing whether there is a significant difference among the lower developmental thresholds on the basis of the traditional linear model describing developmental rate as a linear function of temperature. For comparing the lower developmental thresholds of different developmental stages, a new method based on the Chow test is proposed in the current study. Another feasible way based on the linear model proposed by Ikemoto and Takai is also proposed. The lower developmental thresholds can be compared by the analysis of covariance on this linear model. The current study can be used to test the hypothesis of isomorphy rate. When comparing the lower developmental thresholds of different geographical populations for one insect species, the two methods proposed here are also applicable.     

Effect of temperature on development and survival of the parasitoid Pnigalio pectinicornis (Hymenoptera: Eulophidae) reared on Phyllocnistis citrella (Lepidoptera: Gracillariidae)

Laboratory studies were conducted to assess the effect of temperature on the development and survival of the indigenous parasitoid Pnigalio pectinicornis L. on the citrus leaf miner Phyllocnistis citrella Stainton as host, fed on leaves of Citrus sinensis L. Osbck cultivar Washington navel and Citrus reticulata Blanco cultivar Clementine. Experiments were conducted at five constant temperatures ranging from 15 to 32.5 degrees C, with 60 +/- 10% RH and a photoperiod of 14:10 (L:D) h. The relationship between the developmental rate and temperature was determined using both linear and nonlinear (Lactin's formula) models. Developmental time of immature stages tended to be shorter as the temperature increased the range from 15 to 30 degrees C. Mortality was greater at the temperatures extreme tested. Both linear and nonlinear models provided a reliable fit of developmental rates versus temperature for all immature stages. Developmental thresholds that were estimated by the linear model for eggs were higher than those estimated by the nonlinear model. However, higher values of the low developmental threshold for larva and pupa stage of P. pectinicornis were estimated by the Lactin-2 model than that by the linear model. The potential of these models to predict the phenology of this parasitoid and its biological characteristics found in this study are discussed for its proper use as a biological control agent.     

Are thermal constants constant? A test using two species of ladybird

There is a controversy about whether the thermal constants, lower developmental threshold, rate of development and corresponding degree days required for development, change when a species is reared under different developmental conditions. We present a more precise way of measuring these constants using the linear relationship between the rate of development and temperature. First we use the equation proposed by Ikemoto and Takai (2000) to determine the linear phase of development and then a generalised linear model having a different variance at low and high temperatures, specific for each condition, to estimate the parameters of the linear relationship. Using this method, we show that providing the difference in food quality is sufficiently great, an aphidophagous ladybird develops significantly faster and starts developing at a significantly lower temperature on a good than on a poor quality diet. Adaptive significance of the thermal constants not remaining constant is discussed in terms of a trade-off between growth and rate of development, when temperature and food quality varies.     

Experimental taphonomy of embryo preservation in a Cenozoic brooding bivalve

To distinguish between alternative explanations for the presence of synchronous broods in the Miocene-Pliocene bivalve, Transenriella species. we performed in situ burial experiments of the Recent species T. corfusa . All Recent Transennella species are asynchronous brooders; a single brood contains all or most developmental stages. Specimens from Miocene—Pliocene deposits of California suggest that some members of this taxon were synchronous brooders, i.e., all the embryos of a brood develop simultaneously with only one developmental stage represented at any time. The presence of synchronous Transennella broods in the Miocene—Pliocene could indicate that an evolutionary change in mode of reproduction has occurred in this genus. Alternatively, asynchronous brooding in this taxon may he conservative and preferential preservation of later stages of development, or seasonal variation in reproduction, could result in a taphonomic overprint. Our burial experiments indicate that the earliest stages of development are almost entirely lost; however. there is enough preservation of the later stages of development to distinguish the two modes of reproduction. Additionally. we discovered a single fossil specimen with an asynchronous brood. Based primarily on this specimen, and observations from the burial experiments, we conclude that the fossil synchronous broods are an artifact of preservation and asynchronous brooding in Transenella is conservative.     

Testing the rate isomorphy hypothesis using five statistical methods

Abstract Organisms are said to be in developmental rate isomorphy when the proportions of developmental stage durations are unaffected by temperature. Comprehensive stage‐specific developmental data were generated on the cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), at eight temperatures ranging from 16°C to 30°C (in 2°C increments) and five analytical methods were used to test the rate isomorphy hypothesis, including: (i) direct comparison of lower developmental thresholds with standard errors based on the traditional linear equation describing developmental rate as the linear function of temperature; (ii) analysis of covariance to compare the lower developmental thresholds of different stages based on the Ikemoto‐Takai linear equation; (iii) testing the significance of the slope item in the regression line of versus temperature, where p is the ratio of the developmental duration of a particular developmental stage to the entire pre‐imaginal developmental duration for one insect or mite species; (iv) analysis of variance to test for significant differences between the ratios of developmental stage durations to that of pre‐imaginal development; and (v) checking whether there is an element less than a given level of significance in the p‐value matrix of rotating regression line. The results revealed no significant difference among the lower developmental thresholds or among the aforementioned ratios, and thus convincingly confirmed the rate isomorphy hypothesis.     

Effect of temperature on early development of white and lake sturgeon,Acipenser transmontanus and A. fulvescens

Synopsis The effect of constant incubation temperatures (between 10°C and 26°C) on the developmental rates was found to fit a similar exponential relationship in both the lake and white sturgeon embryos and larvae. Although the lake sturgeon had an overall slower rate of development than the white sturgeon, no statistically significant difference was detected in the slopes of the exponential equations describing the effect of temperature on developmental rate. The effect of these incubation temperatures on embryonic survival also did not differ between these two species. Both species exhibited optimal survival between 14–17° C and incipient mortalities occurred at 20°C. Temperatures above 20°C were lethal for white sturgeon embryos. No effect of low incubation temperature on survival was evident from this study. A comparison of these North American species with Eurasian acipenserids suggests that all the sturgeon that have been examined exhibit a similar influence of incubation temperature on developmental rate.     

Effect of temperature on the duration of egg, nauplius and copepodite development of some freshwater benthic Copepoda

SUMMARY. The development times of the egg, nauplius and copepodite stages of seven species of freshwater benthic copepods ( Paracyclops fimbriatus, Canthocamptus staphylinus, Attheyella crassa, Moraria brevipes, M. mrazeki, Bryocamptus echinatus and Paracamptus schmeili ) were determined at constant temperatures ranging from 1.5 to 24°C using detritus as food. Several regression models used earlier to describe the temperature dependence of development were tested on this material.
Species differed both in the rapidity of development and in the magnitude of the effect of temperature. Development times usually decreased with increasing temperature, but in M. brevipes temperatures exceeding 16°C retarded postembryonic development. P. fimbriatus developed at a rate comparable to that reported earlier for littoral cyclopoids. All the harpacticoid species developed more slowly than planktonic, littoral or benthic copepods previously studied, but faster than subterranean species. The effect of temperature was most marked in species reproducing in warm water.
The relationship between rate of development and temperature was not linear. None of the curvilinear temperature functions tested was statistically adequate in all cases. However, three of the models gave a reasonable fit to all egg development data, and one of these, the semilogarithmic quadratic equation, ln D = In a + T In b + T² In c , also satisfactorily described postembryonic development. This model accommodates the retardation of development at higher temperatures, and can therefore be recommended for further testing.     

Temperature-dependent development of Chilocorus bipustulatus (Coleoptera: Coccinellidae)

The effect of temperature on development and survival of Chilocorus bipustulatus L. (Coleoptera: Coccinellidae), a predator of many scale insects, was studied under laboratory conditions. The duration of development of egg, first, second, third, and fourth larval instars, pupa, and preovioposition period at seven constant temperatures (15, 17.5, 20, 25, 30, 32.5, and 35°C) was measured. Development time decreased significantly with increasing temperature within the range 15-30°C. Survival was higher at medium temperatures (17.5-30(ο)C) in comparison with that at more extreme temperature regimens (15 and >30(ο)C). Egg and first larval instars were the stages where C. bipustulatus suffered the highest mortality levels at all temperatures. The highest survival was recorded when experimental individuals were older than the third larval instar. Thermal requirements of development (developmental thresholds, thermal constant, optimum temperature) of C. bipustulatus were estimated with application of linear and one nonlinear models (Logan I). Upper and lower developmental thresholds ranged between 35.2-37.9 and 11.1-13.0°C, respectively. The optimum temperature for development (where maximum rate of development occurs) was estimated at between 33.6 and 34.7°C. The thermal constant for total development was estimated 474.7 degree-days.     

温度对以截形叶螨为猎物的斯氏钝绥螨生长发育的影响

测定了斯氏钝绥螨Amblyseius swirskii（Athias—Henriot）在14、17、20、23、26、29、32、35℃和RH=85％的条件下,以截形叶螨为猎物各螨态的发育历期,分析发育速率与温度的关系。结果表明,14℃不适宜斯氏钝绥螨的生长发育,存活率仅17．5％;在17～35℃之间其生长发育正常,存活率均100％,各螨态的发育历期与温度均呈负相关关系,发育历期随着温度的升高明显缩短,未成熟期总发育历期从21．36d缩至3．97d。采用线性日度模型拟合温度与发育速率的关系表明,该模型能较好地描述其发育速率在适宜温度范围内随温度升高而加快的现象。斯氏钝绥螨不同发育阶段的发育起点温度和有效积温不同,未成熟期的发育起点温度和有效积温分别为11．92℃和94．69日·度。     

The nature of the dependence of the rate of development on temperature determines the greatest efficiency of metabolism at optimal temperatures

Metabolism intensity and development rate have different pattern of dependence on temperature. Oxygen intake and several other metabolic processes bear an exponential relationship to temperature. The pattern of this relationship is similar in different poikilothermal species. On the contrary, the relationship between the rate of development and temperature is species-specific and more complex. Hence, the curves obtained by plotting oxygen intake per developmental stage against temperature resemble parabola and the minimum values of oxygen intake correspond to optimal temperatures. Such a correspondence is almost solely determined by parameters of the relationship between the rate of development and temperature. Therefore, the efficiency of metabolism does not determine the range of optimal temperatures. It is suggested that adaptive alterations of this range during evolution proceed via changes of the parameters of relationship between the rate of development and temperature dependence. Relatively small number of mutations is required to produce such changes.     

Development, survivorship, and reproduction of Helicoverpa armigera (Lepidoptera: Noctuidae) under constant and alternating temperatures

Laboratory studies were conducted to assess the effect of temperature on the survival, development, fecundity, and longevity of Helicoverpa armigera (Hübner) at 11 constant temperatures ranging from 12.5 to 40 degrees C, as well as at five alternating temperature regimes (25-10, 30-15, 32.5-17.5, 35-20, and 35-27.5 degrees C) and under a photoperiod of 16:8 (L:D) h. H. armigera reared at constant temperatures did not develop from egg to adult (emergence) outside the temperature range of 17.5-32.5 degrees C. The alternating conditions expanded this range from 10 to 35 degrees C. The lowest developmental thresholds of the immature stages were estimated by a linear model and ranged from 10.17 (pupal stage) to 11.95 degrees C (egg stage) at constant temperature regimes and from 1.1 to 5.5 degrees C, respectively at alternating temperatures. The values of developmental thresholds estimated using the nonlinear (Lactin-2) model were lower than those estimated by the linear model for constant and alternating temperature regimes except for larval and pupal stages at constant temperatures. Mean adult longevity fluctuated from 34.4 d at 15 degrees C to 7.6 d at 35 degrees C. Females reared under all alternating temperature regimes laid more eggs than females reared at any, except the 25 degrees C, constant temperature treatment. The intrinsic rate of increase was highest at 27.5 degrees C, at both the constant and the corresponding alternating temperature regimes (0.147 and 0.139, respectively). Extreme temperatures had a negative effect on life table parameters.     

Temperature and diet effects on immature development of predatory mite Typhlodromus athenas Swirski and Ragusa (Acari: Phyotseiidae)

Temperature and food quality both can influence growth rates and developmental time of herbivorous insects and mites. Typhlodromus athenas Swirski and Ragusa is an indigenous mite in the Mediterranean region and data on its temperature dependent development are lacking. In the current study, temperature-dependent development and survival of T. athenas immature stages were evaluated on eggs and all stages of Tetranychus urticae Koch, as well as on almond (Prunus amygdalis Batsch) pollen, under seven constant temperatures ranging from 15 to 35°C, 65% RH, and a photoperiod of 16:8 (L:D) h. On both diets survival was considerably high at all temperatures. The longest developmental period of immature stages was recorded at 15°C, whereas the shortest was at 30°C. Female immatures on almond pollen had shorter developmental time compared to that on twospotted spider mites. Food had a significant effect on female total developmental time at temperatures lower than 25°C. The lower developmental thresholds, estimated by a linear model, for egg-to-adult of females and males fed on pollen were 8.60 and 8.77°C, respectively, whereas on T. urticae they were 10.15 and 10.62°C, respectively. Higher values of tmin for total development were estimated by a nonlinear model (Lactin-2), and ranged from 10.21°C for both females and males on almond, to 11.07 for females and 10.78°C for males on prey. Moreover, this model estimated optimal and lethal temperatures accurately. The results of this study indicate that T. athenas appears better adapted to higher temperatures that occur in the Mediterranean region and may be a useful biological control agent.     

Effect of temperature on the development of the aquatic stages of Anopheles gambiae sensu stricto (Diptera: Culicidae)

Global warming may affect the future pattern of many arthropod-borne diseases, yet the relationship between temperature and development has been poorly described for many key vectors. Here the development of the aquatic stages of Africa's principal malaria vector, Anopheles gambiae s.s. Giles, is described at different temperatures. Development time from egg to adult was measured under laboratory conditions at constant temperatures between 10 and 40 degrees C. Rate of development from one immature stage to the next increased at higher temperatures to a peak around 28 degrees C and then declined. Adult development rate was greatest between 28 and 32 degrees C, although adult emergence was highest between 22 and 26 degrees C. No adults emerged below 18 degrees C or above 34 degrees C. Non-linear models were used to describe the relationship between developmental rate and temperature, which could be used for developing process-based models of malaria transmission. The utility of these findings is demonstrated by showing that a map where the climate is suitable for the development of aquatic stages of A. gambiae s.s. corresponded closely with the best map of malaria risk currently available for Africa.     

Evidence for lower plasticity in CTMAX at warmer developmental temperatures

Understanding the capacity for different species to reduce their susceptibility to climate change via phenotypic plasticity is essential for accurately predicting species extinction risk. The climatic variability hypothesis suggests that spatial and temporal variation in climatic variables should select for more plastic phenotypes. However, empirical support for this hypothesis is limited. Here, we examine the capacity for ten Drosophila species to increase their critical thermal maxima (CT_MAX) through developmental acclimation and/or adult heat hardening. Using four fluctuating developmental temperature regimes, ranging from 13 to 33 °C, we find that most species can increase their CT_MAX via developmental acclimation and adult hardening, but found no relationship between climatic variables and absolute measures of plasticity. However, when plasticity was dissected across developmental temperatures, a positive association between plasticity and one measure of climatic variability (temperature seasonality) was found when development took place between 26 and 28 °C, whereas a negative relationship was found when development took place between 20 and 23 °C. In addition, a decline in CT_MAX and egg‐to‐adult viability, a proxy for fitness, was observed in tropical species at the warmer developmental temperatures (26–28 °C); this suggests that tropical species may be at even greater risk from climate change than currently predicted. The combined effects of developmental acclimation and adult hardening on CT_MAX were small, contributing to a <0.60 °C shift in CT_MAX. Although small shifts in CT_MAX may increase population persistence in the shorter term, the degree to which they can contribute to meaningful responses in the long term is unclear.     

A comparison of different thermal performance functions describing temperature-dependent development rates

The impact of temperature on developmental duration of insects has been long kept a high profile in the studies of insect pests. The relationship between developmental rate, which is the reciprocal of developmental duration, is generally represented by a straight line over a range of moderate temperature; over two ranges of extreme temperature (i.e., low temperatures and high temperatures), the relationship cannot be accurately reflected by a straight line (Campbell et al., 1974). For describing the effect of constant temperature on developmental rate over the full range of temperature, some non-linear models were proposed. To analyze the effect of temperature on ectothermic performance, twelve non-linear functions, including Gaussian, Logan1, Logan2, Performance, Wang–Lan–Ding, Sharpe–Schoolfield, Ratkowsky, Brière1, Brière2, Weibull, modified Gaussian and exponentially modified Gaussian functions, are compared using the coefficient of determination, adjusted coefficient of determination, Akaike information criterion (AIC), Bayesian information criterion (BIC), corrected Akaike information criterion (AICC) and a new method best on a weighted average of the five listed indicators. These models were compared using the development rate data of two species of insects at the egg stage. We found that the Performance, Brière1 and Brière2 functions are all very suitable for explaining temperature-dependent development rates. The three functions both belong to the asymmetrical skew thermal performance curves, and show better goodness-of-fit than the symmetrical Gaussian function. The Performance function might be the best function, because it can reflect the linearity between temperatures and developmental rates below the optimal developmental temperature.     

温度对番茄潜叶蛾生长发育和繁殖的影响

【目的】番茄潜叶蛾Tuta absoluta是一种对番茄具有毁灭性危害的世界性入侵害虫,本研究旨在探索温度对入侵番茄潜叶蛾种群生长发育和繁殖的影响,为预测番茄潜叶蛾的分布区域、田间发生动态提供基础。【方法】在室内测定了番茄叶片上番茄潜叶蛾在15, 20, 25, 30和35℃ 5个恒定温度条件下各虫态的发育历期和存活率、繁殖力和种群增长参数,并应用不同模型分析发育速率、内禀增长率和净生殖率与温度的关系,估计发育起点温度、发育极限温度、发育最适温度、有效积温和年发生代数。【结果】在恒温15~30℃范围内,番茄潜叶蛾各虫态的发育历期随温度升高而逐渐缩短。25℃下幼虫期存活率、成虫前期存活率、雌虫总产卵量、净生殖率、内禀增长率和周限增长率均最大。在35℃下,卵的存活率骤降至11%,孵化的幼虫无法正常发育。卵期、幼虫期、蛹期、成虫前期、全世代的有效积温分别为104.17, 232.59, 129.87, 434.78和526.32日·度,该虫在新疆伊宁县和察布查尔锡伯自治县的理论发生代数为4~5代。基于发育速率与温度关系的模型与基于种群增长参数与温度关系的模型所计算的积温需求参数不同,基于内禀增长率求得的番茄潜叶蛾的发育起点温度、发育极限高温和发育最适温度分别为12.46, 30.40和27.36℃。【结论】入侵我国新疆地区的番茄潜叶蛾适温范围广泛,在我国大部分地区具有极高的扩散风险。     

Physiological control of water flux in conifers

Summary Pre-hatching developmental times for prosobranch gastropods are greatly influenced by temperature and taxonomic affinity. If the data used here (including all available data from the Muricacea) are a representative sample, then reasonably accurate estimates of developmental time can be obtained for most prosobranchs knowing only temperature and taxon. Times are also significantly affected by egg or hatching size. Correlations between developmental time and hatching form are probably accounted for by egg size. Prehatching periods are little, if at all, longer for metamorphosed hatchlings than for swimming hatchlings; in any event, differences are small relative to typical free swimming periods. Therefore, the planktonic period is a substantial addition to the total pre-juvenile period. Many embryos die before hatching. More would survive if development were faster; development is, therefore, prolonged at a measurable selective cost. Factors promoting extended developmental periods should be evaluated with these costs in mind. For example, providing much of the yolk as nurse-eggs may allow a species to have a large hatching size and at the same time a relatively brief developmental time.     

Influence of temperature on developmental rate,wing length,and larval head capsule size of pestiferous midge Chironomus crassicaudatus (Diptera: Chironomidae)

Larvae of Chironomus crassicaudatus Malloch were reared individually at nine constant temperatures from 12.5 to 32.5 degrees C (2.5 degrees C increments) for 120 d. Duration of immature stages (egg, four instars, and pupa), head capsule width of fourth instars, and wing length were recorded. Some adults emerged at all temperatures, except at 12.5 degrees C where individuals developed to fourth instars during the experiment. Sharpe and DeMichele's four-parameter model with high-temperature inhibition described the temperature-dependent developmental rates. The slowest development was observed at 15 degrees C, with developmental rate peaking between 25 and 27.5 degrees C. Developmental rate increased rapidly with increasing temperature up to 20 degrees C, slowed between 20 and 27.5 degrees C, and decreased at temperatures >27.5 degrees C. No developmental inhibition at high temperatures was observed in eggs. The most apparent high-temperature inhibition of development was recorded in fourth instars, which comprised the largest proportion of developmental time. Males developed faster than females, but females had wider larval head capsules and longer wings than males. Adult size was negatively related with temperature in both sexes, but this relationship was steeper in males than in females. Larval size peaked at 20 degrees C, whereas the head capsule width was reduced at temperatures higher and lower than 20 degrees C.