A roadmap for ladybird conservation and recovery.

Ladybirds (Coleoptera: Coccinellidae) provide services that are critical to food production, and they fulfill an ecological role as a food source for predators. The richness, abundance, and distribution of ladybirds, however, are compromised by many anthropogenic threats. Meanwhile, a lack of knowledge of the conservation status of most species and the factors driving their population dynamics hinders the development and implementation of conservation strategies for ladybirds. We conducted a review of the literature on the ecology, diversity, and conservation of ladybirds to identify their key ecological threats. Ladybird populations are most affected by climate factors, landscape composition, and biological invasions. We suggest mitigating actions for ladybird conservation and recovery. Short-term actions include citizen science programs and education, protective measures for habitat recovery and threatened species, prevention of the introduction of non-native species, and the maintenance and restoration of natural areas and landscape heterogeneity. Mid-term actions involve the analysis of data from monitoring programs and insect collections to disentangle the effect of different threats to ladybird populations, understand habitat use by taxa on which there is limited knowledge, and quantify temporal trends of abundance, diversity, and biomass along a management-intensity gradient. Long-term actions include the development of a worldwide monitoring program based on standardized sampling to fill data gaps, increase explanatory power, streamline analyses, and facilitate global collaborations.

Las catarinas (Coleoptera: Coccinellidae) proporcionan servicios que son críticos para la producción de alimento, y juegan un papel ecológico como fuente de alimento para depredadores. Sin embargo, la riqueza, abundancia y distribución de catarinas están en peligro debido a muchas amenazas antropogénicas. La carencia de conocimiento sobre el estatus de conservación de la mayoría de las especies y los factores que inciden en su dinámica poblacional dificulta el desarrollo e implementación de estrategias de conservación para las catarinas. Realizamos una revisión de la literatura sobre la ecología, diversidad y conservación de catarinas para identificar sus amenazas ecológicas clave. Las poblaciones de catarinas fueron afectadas mayormente por factores climáticos, composición del paisaje e invasiones biológicas. Proponemos acciones de mitigación para la conservación y recuperación de catarinas. Acciones a corto plazo incluyen programas de ciencia y educación ciudadana, medidas de protección para la recuperación de hábitat y de especies amenazadas, prevención de la introducción de especies no nativas y el mantenimiento y restauración de áreas naturales y la heterogeneidad del paisaje. Acciones a mediano plazo implican el análisis de datos obtenidos de programas de monitoreo y colecciones de insectos para desenmarañar el efecto de las diferentes amenazas a las poblaciones de catarinas, comprender el uso del hábitat por taxa de los que se tiene conocimiento limitado y cuantifica las tendencias temporales de la abundancia, diversidad y biomasa a lo largo de un gradiente de intensidad de manejo. Acciones a largo plazo incluyen el desarrollo de un programa de monitoreo a nivel mundial basado en muestreos estandarizados para subsanar la falta de datos, incrementar el poder explicativo, optimizar los análisis y facilitar colaboraciones globales.

Effect of Male Body Size on Female Reproduction in Pyrrhocoris apterus (L.) (Heteroptera, Pyrrhocoridae).

Females and males of the abundant heteropteran species Pyrrhocoris apterus (L.) mate with the larger of the available partners. The male benefits from copulation with a large female, which is more productive than a small female. However, the benefit to females from copulation with a large male has not yet been investigated. Overwintered sexually immature adults were transferred from the field to indoors for a long day at 25 °C and subsequently kept as female-male pairs with different partner size ratios. The female lays eggs in several successive clutches. We investigated the influence of male size on the number and weight of eggs in individual clutches and the time elapsed from the transfer to their deposition. These characteristics of reproduction were first regressed on female size, and residuals of this regression were then regressed on male size. The positive effect of increasing male size on the characteristics of female reproduction manifested as a significant value of the latter regressions. The expected positive effect of increasing male body size on female reproductive characteristics was not detected. Several reasons for this deficiency are discussed.

Which Seed Properties Determine the Preferences of Carabid Beetle Seed Predators?

Ground beetles are important invertebrate seed predators in temperate agro-ecosystems. However, there is a lack of information regarding which seed properties are important to carabids when they select seeds for consumption. Therefore, seed properties, such as size, shape, morphological defence, and chemical composition, were measured, and in addition to seed taxonomy and ecology, these data were used to explain carabid preferences. Carabid preferences were assessed using a multi-choice experiment with 28 species of weed seeds presented to 37 species of Carabidae. Multiple regression on distance matrices (MRM) was used to determine the importance of particular sets of seed properties for carabids. The analysis was conducted for the full set of carabids (37 species) as well as for subsets of species belonging to the tribes of Harpalini or Zabrini. For the complete set of species, seed dimensions, seed mass, taxonomy, plant strategy, and seed coat properties significantly explained carabid preferences (proportion of explained variance, R2 = 0.465). The model for Harpalini fit the data comparably well (R2 = 0.477), and seed dimensions, seed mass and seed coat properties were significant. In comparison to that for Harpalini, the model for Zabrini had much lower explanatory power (R2 = 0.248), and the properties that significantly affected the preferences were seed dimensions, seed mass, taxonomy, plant strategy, and seed coat properties. This result suggests that the seed traits that carabids respond to may be specific to taxonomic and likely relate to the degree of specialisation for seeds. This study contributes to understanding the mechanisms that determine the preferences of carabid beetles for seeds.

Behavioural thermoregulation hastens spring mating activity in Pyrrhocoris apterus (Heteroptera: Pyrrhocoridae).

Post-diapause, overwintered adults of the true bug Pyrrhocoris apterus L. (Heteroptera, Pyrrhocoridae) form conspicuous aggregations at warm spots in early spring. Using a combination of laboratory experiments and field observations, we assessed the influence of this behaviour on the seasonal timing of reproduction. In the laboratory, post-diapause pairs mated after an accumulation of 80° days (dd), and females started to lay eggs after an additional 80 dd (considering 10.3 °C as a lower temperature threshold for post-diapause development). In the field, however, females mated as early as late March (based on data from four seasons), which corresponded to the accumulation of <15 dd (based on summing the temperatures recorded at a meteorology station). Such a 'discrepancy' between laboratory and field data is explained by thermoregulation. Aggregated adults bask in sunny spots, thereby increasing their body temperatures to ~25 °C, which is 7-16 °C above the temperature of the surrounding ground. This thermal excess speeds up their post-diapause development and enables early mating and oviposition. As a result, behavioural thermoregulation hastens the seasonal start of reproduction in P. apterus by > 1 month.

Differences in the Phenology of Harmonia axyridis (Coleoptera: Coccinellidae) and Native Coccinellids in Central Europe.

Harmonia axyridis (Pallas), an invasive non-native species in central Europe, can outcompete other aphidophagous species. The distribution and abundance of H. axyridis vary depending on different host plants, and its effects on native coccinellid communities may change accordingly. The distribution and abundance of coccinellids in central Europe (50°N, 14°E) were investigated from 2010 to 2016. Coccinellids were counted at regular intervals on cereals (Avena, Hordeum, and Triticum), herbaceous plants (Matricaria and Urtica) and trees (Acer, Betula, and Tilia). Additionally, the occurrence over time of each species on these plants was assessed and used as an index of persistence. Across all years, the adults and larvae of H. axyridis were the dominant species of coccinellid on trees. However, H. axyridis was less abundant on herbaceous plants and cereals than on trees. Populations of native coccinellids and H. axyridis co-occurred on trees and persisted for the same length of time, while native coccinellids persisted longer than H. axyridis on herbaceous plants and cereals. Compared to 1976-1986, in the 2010s, the abundance of native species decreased on all plants by 50-70%. The presence of H. axyridis could be considered as a factor driving changes in the assemblages of native coccinellids.

Aphids (Homoptera: Aphididae) on Winter Wheat: Predicting Maximum Abundance of Metopolophium dirhodum.

In Central Europe, the most abundant aphid infesting the leaves of small grain cereals is Metopolophium dirhodum (Walker) (Homoptera: Aphididae). Annual variation in its seasonal dynamics was evaluated using a 25-yr series of standardized weekly censuses of winter wheat plots. M. dirhodum made up >50 % of the aphids on the foliage. Date of immigration (8 May-3 July), length of period of population increase (0-9 wk), and date of attaining maximum abundance (28 May-22 July) varied greatly. For the prediction, we regressed maximum numbers/tiller on numbers recorded in the first week after heading. The regression of maximum abundance on nonzero aphid counts revealed a critical number of ≥1.50 aphids/tiller, which if exceeded resulted in a harmful maximum abundance of ≥10 aphids/tiller at the peak. Zero aphid counts resulted in 10% of cases with a harmful maximum abundance. Using this regression for prediction will result in 18% of the recorded cases being false negatives and 9% false positives. Parallel annual variation in the average maximum numbers of M. dirhodum, Sitobion avenae (Fabricius) (Homoptera: Aphididae), and Rhopalosiphum padi (Linné) (Homoptera: Aphididae) indicated the following factors that affected their abundance: temperature in winter and host plant quality. The predictions apply only in areas where M. dirhodum is holocyclic and aphids do not overwinter in wheat stands.

Estimating Prey Consumption in Natural Populations of Harmonia axyridis (Coleoptera: Coccinellidae) Using Production of Feces.

Production of feces (PF) is a useful proxy indicating quantity of ingested food. Although influenced by many uncontrolled factors PF provides insight into food consumption under natural conditions. In Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) variation in PF was investigated in populations of various hostplants at localities of central Bohemia (50N, 14E), throughout the season of 2016. The adults collected from these hostplants were starved for 48 h at standard conditions, and dry mass of feces produced during this period was measured. Despite enormous differences in PF among individuals, significant variation over the season occurred in average PF of both males and females. PF increased with abundance of aphids and was significantly greater in females than males. Gravidity, as manifested through oviposition within 48 h after capture, was associated with increased PF, while hostplant and color morph did not affect variation in PF among individuals. From PF as measured in this study, it can be estimated that at sites hosting abundant aphid populations H. axyridis (as an adult male or female) may consume 19 (male) to 45 (female) aphids per day (assumed body length 1mm). In the absence of aphids, adults may consume one to nine individuals of alternative prey per day (body length 1-2 mm).

Spatial and temporal changes in the abundance and compostion of ladybird (Coleoptera: Coccinellidae) communities.

Because of their services to agriculture most ladybirds (Coleoptera: Coccinellidae) are intensively studied predators of mainly phytophagous pests. The study of the long-term variation in the composition of their communities was stimulated by recent dramatic changes in the abundance of some species. We review and evaluate possible effects of the main causes cited in the literature. Agricultural and habitat changes (particularly urbanization) affect coccinellid abundance, both negatively and positively. In the temperate zone dominant species occur most frequently associated with abundant prey populations on crops, weeds and planted stands of trees resulting from human activity. Invasive non-native species of coccinellids may endanger native species through intraguild predation or competition for resources, but their supposed serious negative effects on native species can differ considerably. Climatic change may influence coccinellid species in several ways, including indirect effects through lower trophic levels and desynchronisation of the phenologies of host plants, prey and coccinellid populations. In the near future we do not expect climate warming to have important effects on ladybird diversity globally, but local changes in the composition of coccinellid communities and abundance of particular species could occur.

Treating Prey With Glyphosate Does Not Alter the Demographic Parameters and Predation of the Harmonia axyridis (Coleoptera: Coccinellidae).

Glyphosate is an herbicide that is used worldwide with potential environmental risks to nontarget organisms. We applied an age-stage, two-sex life table approach to assess the sublethal effects of short-term oral exposure to a glyphosate-based herbicide on the life table parameters and biocontrol potential of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). Aphids (Metopolophium dirhodum (Walker) (Sternorrhyncha: Aphididae)) treated with herbicide (an isopropylamine-salt of glyphosate) at low recommended, maximum recommended, and double the maximum recommended concentration for agricultural situations, and untreated controls were offered to the fourth instar of H. axyridis for 24 h. Development, consumption, and fecundity were measured daily until death. We detected minor differences in the hatching rate and mean generation time, whereas the longevity, fecundity, net reproductive rate, intrinsic rate of increase, finite rate of increase, and consumption were unaffected across treatments. We conclude that biocontrol potential of H. axyridis was not affected by acute oral intoxication by a glyphosate-based herbicide during the larval stage for 24 h under the study design.

Treatment by glyphosate-based herbicide alters life history parameters of the rose-grain aphid Metopolophium dirhodum.

Glyphosate is the number one herbicide in the world. We investigated the sub-lethal effects of this herbicide on the aphid Metopolophium dirhodum (Walker), using an age-stage, two-sex life table approach. Three concentrations of the herbicide (low - 33.5, medium - 66.9 and high - 133.8 mmol dm(-3) of active ingredient) and distilled water as the control were used. The LC50 of the IPA salt of glyphosate on M. dirhodum was equivalent to 174.9 mmol dm(-3) of the active ingredient (CI95: 153.0, 199.0). The population parameters were significantly negatively affected by herbicide application, and this negative effect was progressive with the increasing concentration of the herbicide. A difference of two orders of magnitude existed in the predicted population development of M. dirhodum between the high concentration of the herbicide and the control. This is the first study that comprehensively documents such a negative effect on the population of an herbivorous insect.

Invasive Insects Differ from Non-Invasive in Their Thermal Requirements.

We tested whether two basic thermal requirements for insect development, lower developmental thresholds, i.e. temperatures at which development ceases, and sums of effective temperatures, i.e. numbers of day degrees above the lower developmental thresholds necessary to complete development, differ among insect species that proved to be successful invaders in regions outside their native range and those that did not. Focusing on species traits underlying invasiveness that are related to temperature provides insights into the mechanisms of insect invasions. The screening of thermal requirements thus could improve risk-assessment schemes by incorporating these traits in predictions of potentially invasive insect species. We compared 100 pairs of taxonomically-related species originating from the same continent, one invasive and the other not reported as invasive. Invasive species have higher lower developmental thresholds than those never recorded outside their native ranges. Invasive species also have a lower sum of effective temperatures, though not significantly. However, the differences between invasive and non-invasive species in the two physiological measures were significantly inversely correlated. This result suggests that many species are currently prevented from invading by low temperatures in some parts of the world. Those species that will overcome current climatic constraints in regions outside their native distribution due to climate change could become even more serious future invaders than present-day species, due to their potentially faster development.

Plasticity of the thermal requirements of exotherms and adaptation to environmental conditions.

UNLABELLED: In exothermal organisms, temperature is an important determinant of the rate of ecophysiological processes, which monotonically increase between the minimum (t d min) and maximum (t d max) temperatures typical for each species. In insects, t d min and t d max are correlated and there is a approximately 20°C interval (thermal window W T = t d max - t d min) between them over which insects can develop. We assumed that other exotherms have similar thermal windows because the thermal kinetics of their physiological processes are similar. In this study, we determined the thermal requirements for germination in plants. Seeds of 125 species of Central European wild herbaceous and crop plants were germinated at nine constant temperatures between 5 and 37°C, and the time to germination of 50% of the seeds D and rate of germination R (=1/D) were determined for each temperature and the Lactin model used to determine t d min, t d max, and W T. The average width of the thermal windows for seeds was significantly wider (mean 24°C, 95% CI 22.7-24.2°C), varied more (between 14.5 and 37.5°C) and development occurred at lower temperatures than recorded for insects. The limiting temperatures for germination, t d min and t d max, were not coupled, so the width of the thermal window increased with both a decrease in t d min and/or increase in t d max. Variation in W T was not associated with taxonomic affiliation, adult longevity, or domestication of the different species, but tends to vary with seed size. Plants are poor at regulating their temperature and cannot move to a more suitable location and as a consequence have to cope with wider ranges in temperatures than insects and possibly do this by having wider thermal windows. SYNTHESIS: The study indicated specificity of W T in different exotherm taxa and/or their development stages.

Population dynamics of aphids on cereals: digging in the time-series data to reveal population regulation caused by temperature.

Aphid populations show periodic fluctuations and many causes are attributed to their dynamic. We investigated the regulation by temperature of the aphid populations composed of Metopolophium dirhodum, Sitobion avenae, and Rhopalosiphum padi on winter wheat using a 24 years long time series data. We computed the sum of daily temperatures above 5 °C, the threshold temperature for aphid development, and the sum of daily temperatures within the [0(threshold for wheat development),5] °C interval. Applying Generalised Additive Model framework we tested influences of temperature history expressed via degree days before the start of the aphid immigration on the length of their occurrence. We aimed to estimate the magnitude and direction of this influence, and how far to the past before the start of the aphid season the temperature effect goes and then identify processes responsible for the effect. We fitted four models that differed in the way of correcting for abundance in the previous year and in specification of temperature effects. Abundance in the previous year did not affect the length of period of aphid population growth on wheat. The temperature effect on the period length increased up to 123 days before the start of the current season, i.e. when wheat completed vernalization. Increased sum of daily temperatures above 5 °C and the sum of daily temperatures within the [0,5] °C interval both shortened the length of period of aphid population growth. Stronger effect of the latter suggests that wheat can escape from aphid attacks if during winter temperatures range from 0 to 5 °C. The temperature influence was not homogeneous in time. The strongest effect of past temperature was about 50 to 80 and 90 to 110 days before the beginning of the current aphid season indicating important role of termination of aphid egg dormancy and egg hatching.

The invasion history, distribution and colour pattern forms of the harlequin ladybird beetle Harmonia axyridis (Pall.) (Coleoptera, Coccinellidae) in Slovakia, Central Europe.

The harlequin ladybird beetle Harmonia axyridis (Coleoptera, Coccinellidae) has invaded and established in Slovakia. Following unintentional introduction in 2008, the spread of the alien coccinellid was very fast. By the end of 2009, it was recorded across the whole country, and by the end of 2012 it was widely distributed and common in various habitats, particularly gardens, orchards and urban areas, where it was most frequent on trees. The rate of eastward spread was approximately 200 km year(-1), similar to the overall rate of spread in Europe. Between 2008 and 2012, the coccinellid was recorded in a total of 153 localities, in altitudes ranging from 98 to 1,250 m. Most records of this species were made in lowlands, hilly areas and valleys separating mountain ridges. However, it was only rarely documented in areas above 700 m a.s.l. The non-melanic colour form (f. succinea) was dominant along a longitudinal transect including eight urban areas across Slovakia, with the frequency of melanic forms (f. spectabilis and f. conspicua together) between 6.3 and 19.2% and a median equal to 10.5%. The invasion history and distribution of H. axyridis in Slovakia are discussed with regard to the time sequence of records, rate of spread, altitudinal distribution, anthropogenic dispersal, effective recording, proportion of melanic forms and other relevant aspects associated with the spread of this successful invader.

Molecular approach to describing a seed-based food web: the post-dispersal granivore community of an invasive plant.

UNLABELLED: Communities of post-dispersal granivores can shape the density and dispersion of exotic plants and invasive weeds, yet plant ecologists have a limited perception of the relative trophic linkages between a seed species and members of its granivore community. Dandelion seeds marked with Rabbit IgG were disseminated into replicated plots in the recipient habitat (South Dakota) and the native range (Czech Republic). Arthropods were collected in pitfall traps, and their guts were searched for the protein marker using enzyme-linked immunosorbent assay (ELISA). Seed dishes were placed in each plot, and dandelion seed removal rates were measured. The entire experiment was repeated five times over the dandelion flowering period. Gut analysis revealed that approximately 22% of specimens tested positive for the seed marker. A more diverse granivore community had trophic linkages to seeds than has been previously realized under field conditions. This community included taxa such as isopods, millipedes, weevils, rove beetles, and caterpillars, in addition to the traditionally recognized ants, crickets, and carabid beetles. Rarefaction and Chao analysis estimated approximately 16 and 27 species in the granivore communities of the Czech Republic and South Dakota, respectively. SYNTHESIS: Generalist granivore communities are diverse and polyphagous, and are clearly important as a form of biotic resistance to invasive and weedy plants. These granivore communities can be managed to limit population growth of these pests.

Temperature effects on pitfall catches of epigeal arthropods: a model and method for bias correction.

Carabids and other epigeal arthropods make important contributions to biodiversity, food webs and biocontrol of invertebrate pests and weeds. Pitfall trapping is widely used for sampling carabid populations, but this technique yields biased estimates of abundance ('activity-density') because individual activity - which is affected by climatic factors - affects the rate of catch. To date, the impact of temperature on pitfall catches, while suspected to be large, has not been quantified, and no method is available to account for it. This lack of knowledge and the unavailability of a method for bias correction affect the confidence that can be placed on results of ecological field studies based on pitfall data.Here, we develop a simple model for the effect of temperature, assuming a constant proportional change in the rate of catch per °C change in temperature, r, consistent with an exponential Q10 response to temperature. We fit this model to 38 time series of pitfall catches and accompanying temperature records from the literature, using first differences and other detrending methods to account for seasonality. We use meta-analysis to assess consistency of the estimated parameter r among studies.The mean rate of increase in total catch across data sets was 0·0863 ± 0·0058 per °C of maximum temperature and 0·0497 ± 0·0107 per °C of minimum temperature. Multiple regression analyses of 19 data sets showed that temperature is the key climatic variable affecting total catch. Relationships between temperature and catch were also identified at species level. Correction for temperature bias had substantial effects on seasonal trends of carabid catches.Synthesis and Applications. The effect of temperature on pitfall catches is shown here to be substantial and worthy of consideration when interpreting results of pitfall trapping. The exponential model can be used both for effect estimation and for bias correction of observed data. Correcting for temperature-related trapping bias is straightforward and enables population estimates to be more comparable. It may thus improve data interpretation in ecological, conservation and monitoring studies, and assist in better management and conservation of habitats and ecosystem services. Nevertheless, field ecologists should remain vigilant for other sources of bias.

Efficiency of host utilisation by coleopteran parasitoid.

In insect larvae, optimising food utilisation with respect to available meals and time is essential for achieving maximum adult body size, which is a relevant proxy of fitness. We studied the efficiency of food conversion, body size, mortality, and development time in a solitary idiobiont ectoparasitoid, Brachinus explodens (Coleoptera: Carabidae), reared in the laboratory on the pupae of another carabid genus, Amara. The efficiency of conversion index (ECI - ratio of ingested to assimilated food) was, on average, 54.1±1.1% (n=76), with a minimum of 26.9% and a maximum of 81.6%. The rate of increase in biomass gained (W(gained)) with biomass of the host was constant in females, but it decreased in males over the range of host body mass. Females, therefore, grew heavier from hosts of the same mass compared to males. Body length increased with the host mass and was correlated with W(gained) identically for both sexes. Mortality was unaffected by the host mass, but it significantly increased below 20°C. In contrast, the development time of the feeding phase of the larva increased with the host mass at 20.3 and 23.7°C, but it remained unaffected at 26.9°C and in all three temperatures considering pupal development. W(gained) increased with development time up to ca. 8 days of larval feeding at 23.7°C. To our knowledge, our data are the first on food utilisation in solitary idiobiont coleopteran ectoparasitoids, and they present the highest values of ECI in insects.

Developmental database for phenology models: related insect and mite species have similar thermal requirements.

Two values of thermal requirements, the lower developmental threshold (LDT), that is, the temperature at which development ceases, and the sum of effective temperatures, that is, day degrees above the LDT control the development of ectotherms and are used in phenology models to predict time at which the development of individual stages of a species will be completed. To assist in the rapid development of phenology models, we merged a previously published database of thermal requirements for insects, gathered by online search in CAB Abstracts, with independently collected data for insects and mites from original studies. The merged database comprises developmental times at various constant temperatures on 1,054 insect and mite species, many of them in several populations, mostly pests and their natural enemies, from all over the world. We show that closely related species share similar thermal requirements and therefore, for a species with unknown thermal requirements, the value of LDT and sum of effective temperatures of its most related species from the database can be used.

A general rule for the dependence of developmental rate on temperature in ectothermic animals.

In animals that do not regulate their body temperature by the production of heat, the proportion of the total developmental time spent in a particular developmental stage does not change with temperature. In the quasi-linear region of the relationship between developmental rate and temperature, all of the developmental stages appear to have the same species-specific lower developmental threshold. This trait, which is called developmental isomorphy, constrains developmental adaptations of ectotherms to their environments and facilitates the precise timing of life-history events.

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.