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This is the first compilation of our research and data from the literature on the duration and thermal reaction norms for development in ants. Altogether, 97 regression lines characterizing the linear dependence of ant brood (egg, larval, prepupal and pupal) development on temperature were obtained for 33 species from 15 genera and three subfamilies. Significant positive correlations between the durations of different immature stages were revealed. We found differences between thermal reaction norms for development for various immature stages, some taxonomic groups, and southern and northern groups of species. All immature stages appeared to be shorter on average at 25°C in northern ants compared to southern species; this difference is insignificant only for eggs. Highly significant negative correlations were revealed between the temperature threshold for development (TTD) and the sum of degree-days (SDD) for all immature stages. The latitudinal trends in intraspecific variation of thermal constants appeared to be opposite to those we observed at the interspecific level. At the latter, the coefficient of thermal sensitivity of development (i.e. the coefficient of linear regression of development rate on temperature) and TTD tends to decrease and SDD to increase from the south to the north. In contrast, at the intraspecific level, development became more temperature-sensitive in northern populations, i.e. characterized by higher slopes of regression lines of development rate on temperature, and higher TTDs. The species of the genus Formica are characterized by the shortest and the most temperature-sensitive immature development among all the ant species studied.

The data are obtained on development time at six constant temperatures (12, 14, 16, 18, 20, 22°C) and thermal requirements for preimaginal development in a ground beetle Amara communis from Arkhangelsk (64°34′N) and St. Petersburg (59°53′N). The larval and pupal development times were found to be significantly shorter in the Arkhangelsk than in the St. Petersburg population under all temperatures. As a result, total preimaginal development appeared to be shorter by 6.2–6.6% in the Arkhangelsk population. The regression lines of the larval, pupal and total (egg-to-adult) development rate on temperature for the Arkhangelsk population run above and steeper than the respective lines for the St. Petersburg population. Both populations share the similar values of the thermal thresholds (7.2–8.2°C). This explains faster preimaginal development in the northern population under all temperatures above the threshold. Thus, the slope of the regression lines increases, i.e., the sum of degree-days decreases, whereas the thermal threshold for development exhibited no distinctive changes from south to north in this species. Adults from Arkhangelsk reared in the experiments appeared heavier on the average in comparison with those from St. Petersburg, especially at 18–22°C. Temperature did not significantly affect adult weight, except the fact that the beetles were slightly heavier at 20 and 22°C. Consequently, the well-known “temperature-size rule” is violated in this species. Relative growth rate in larvae of A. communis increased considerably with temperature rise from 14 to 22°C. It was significantly higher in the beetles from Arkhangelsk at 18–22°C. There were no differences in larval growth rate between the two populations at 14 and 16°C.

Sexual dimorphism in body mass was studied in prepupae, pupae and immature adults of the green dock leaf beetle Gastrophysa viridula under a constant temperature of 24°C and two photoperiods (12L:12D and 22L:2D). Females were significantly heavier compared to males at all the three developmental stages; just after emergence from pupae they weighed, on average, 11.4 mg, while males—10.3 mg. Sex differences in photoperiodic body mass plasticity were revealed: under short-day conditions pupal and adult mass in females was significantly higher compared to long day, but not in males. Such a response to short day (the factor inducing reproductive diapause in this species) is likely to promote accumulation of greater amount of nutrients by females which is necessary for earlier oviposition after overwintering.

Development times of preimaginal stages and immature adult body mass of the dock leaf beetle Gastrophysa viridula from Bryansk (53°N, 34°E) were determined at six constant temperatures ranging from 16 to 26°C and at two photoperiods, 12L: 12D or 22L: 2D. The lower developmental threshold (LDT) and sum of degreedays (SDD) were calculated for eggs, larvae, and pupae; the values for the total preimaginal development were 7.9°C and 274.8 °C×d, respectively. The thermal reaction norms for preimaginal development were not affected by photoperiod, although the adult body mass was on average greater under short-day conditions. Our results confirm the previously published data on G. viridula concerning diapause induction by shorter daylengths, greater body mass under lower temperatures, and narrowing of the thermal optimum range during preimaginal development. The thermal reaction norms for larval development were shown to be the same in individuals that died during the pupal stage, and in those that successfully emerged as adults. Our results are more precise than the previous data on this species due to a larger sample size, careful control of rearing conditions, and the use of an alternative method for calculating LDT and SDD. In particular, a more accurate estimation of LDTs allowed us to demonstrate that rate isomorphy was violated in this species, contrary to what had been stated by Honěk and co-authors (2003).

As a result of acclimation populations of long-lived ectotherms should display lowered ability to counter cold stress in warmer periods of active season, and increased resistance in colder ones. We tested this proposition by investigating dynamics of cold resistance in Myrmica ants during most of the active season in two types of habitats. Resistance of ants to knock-down by cold and their rate of recovery after chill coma were expected to be lower in summer. Cooled at a rate of 0.17 °C min−1, the ants showed lower capability to resist knock-down in summer, and a significant lowering in knock-down temperature in response to colder weather both in spring and autumn as confirmed by linear regression against air temperatures. In a more eurytopic species M. rubra the responses were significantly faster in meadow than in forest habitats. However, times of recovery of the ants after 10 min at −3 °C did not change in parallel to air temperatures. Whereas M. rubra from forest habitats took less time to recover in early summer and early autumn, in their conspecifics from meadow habitats the contrary was the case. Regardless of habitat, recoveries tended to be faster in other investigated species, of which M. ruginodis (a forest stenotopic) recovered faster in early summer than later. According to the knock-down data, in warmer months the ants are indeed less resistant to cold stress, whilst the recovery data do not always support the proposition. The contrasting seasonal dynamics of the two measures of low-temperature resistance in field-fresh Myrmica suggest that knock-down (chill coma onset) is a better index of thermal acclimation, whilst the rate of recovery from chill coma is more indicative of interspecific differences and, possibly, behavioural thermoregulation.

The main parameters (thermal constants) characterizing the effects of constant temperatures on the duration and rate of development in ectotherms are considered. The advantages and drawbacks of the linear and nonlinear models describing the dependence of the development rate on temperature are discussed. The main patterns of interspecific variation of thermal constants in insects are described. The development of new ideas in the study of intraspecific variation of thermal reaction norms for development in insects is examined. The discrepancies in the data and their interpretation in this field are shown to result mostly from using the traditional methods of determining thermal constants. According to this method, the development rates calculated as reciprocals of the mean development times at each experimental temperature are used for regression analysis. The standard errors calculated according to this method are usually too large. Instead, a fundamentally new method is employed by the authors, according to which the individual development rates calculated for each individual are used for regression analysis of the reciprocals of the mean development times. This method reduces the standard errors of thermal constants by about an order and reveals significant differences between populations and other groups of insects. New data on intraspecific variation of the thermal reaction norms for development in Myrmica ants, the linden bug Pyrrhocoris apterus, and Calliphora blowflies are briefly described and discussed. The possible adaptive significance of these forms of intraspecific variation is discussed. The prospects of research in this new field are outlined.

The ants L. semenovi has been found to belong to species with endogenous-heterodynamic seasonal life cycles with the obligate diapause induced predominantly by factors internal for a colony, whereas external ecological factors (photoperiods and temperature) produce merely modifying effects by accelerating or delaying the diapause onset. The photoperiodic and temperature regulation of diapause induction in larvae and queens is shown. Under effect of short days and low temperature the periods of larval pupation and queen oviposition in a colony are shortened markedly, i.e., the diapause of larvae and queens occurs earlier. The daily rhythms of temperature 15/25°C and particularly 20/30°C as compared with constant temperatures 20 and 25°C that correspond to the mean circadian temperatures of the thermorhythm, inhibit manifestations of the short day effects by stimulating the non-diapause development and increasing duration of the seasonal development cycle of ant colonies. The L. semenovi photoperiodic reaction is quantitative, as development and pupation of larvae and egg-laying of queens cease sooner or later under both the short and the long days, but in the latter case significantly later. Thus L. semenovi is one more example among very rare ant species that are revealed to have the photoperiodic regulation of the colony development seasonal cycle.

For 3–4 generations, selection of the nymphs most rapidly and slowly developing at 28°C was performed in four families of the red soldier bug Pyrrhocoris apterus L. In each generation, duration of development of nymphs was determined at 5 constant temperatures from 20°C to 28°C. From these data the linear regression coefficient of the development rate against temperature (the development thermolability coefficient) for each generation was calculated. As a result of the selection the average duration of the nymph development was shortened or increased statistically significantly depending on its direction. The artificial selection for the development duration has been established to change not only this parameter, but also the temperature norms of the insect development. At selection for fast development the regression straight line slope (i.e., the regression coefficient value) increased statistically significantly, i.e., development became more dependent on temperature. As a rule, the temperature threshold of development increased. At selection for slow development the values of the regression coefficient and of the threshold decreased, but these differences were not statistically significant. The effect of artificial selection for the development duration on temperature norms of insect development has been revealed for the first time.

Expression of HSP70 is induced by stress factors, including sublethal chilling. However, the role of HSP70 for overcoming the consequences of cold stress is not clear. If it is positive, the level of HSP70 expression might be higher in populations from cold climates. Using the immunoblotting technique we investigated dynamics of HSP70 expression in response to cold stress in two Myrmica species (M. rubra and M. ruginodis) from three localities of different latitudes (50, 60 and 67°N). The results showed that in the more thermophilic species Myrmica rubra, expression of HSP70 after cold stress was higher. Within both species, HSP70 expression did not show a direct relationship with latitude. The southernmost population of M. rubra and northernmost population of M. ruginodis displayed the fastest and the most intense response. However, two other populations of M. rubra were similar in timing and intensity of the response, while in M. ruginodis the intermediate population showed the slowest and weakest response. The data suggest that expression of HSP70 may play essential role for adaptation to cold only in the northernmost population of M. ruginodis

The influence of photoperiod on the thermal requirements for development was discovered for the first time in insects during experiments on the linden-bug, Pyrrhocoris apterus. The effect of photoperiod on the duration of linden-bug development at five constant temperatures (20, 22, 24, 26 and 28°C) was measured and the thermal requirements for development at three photope- riods (14, 17 and 20 h light per day) were calculated. Bugs from four geographic populations were used in these experiments: Pyatigorsk (44°02´N, 43°04´E), Borisovka (50°36´N, 36°01´E), Mikhailov (54°15´N, 39°0´E) and Ryazan (54°36´N, 39°42´E). From the values of individual development times at different temperatures the coefficient of linear regression of development rate (the inverse of the duration) on temperature and the thermal threshold for development were calculated. Both these parameters were found to decrease significantly with decrease in day-length for all four populations studied. It means that at shorter day-lengths nym- phal development is less dependent on temperature compared to the development at longer day-lengths. These effects seem to be adaptive. The development times of nymphs at relatively high temperatures (above 24-25°C) are shorter under long-days than under short days which should be advantageous at the height of summer when the days are long and the weather is warm. In the contrast, at relatively low temperatures (below 24-25°C) the nymphs develop significantly faster under short-days than under long days, which is advantageous at the end of summer as it allows the nymphs to reach the adult stage, the only stage capable of overwintering. The influence of photoperiod on the thermal reaction norm appeared to be more or less gradual, i.e. the shorter the day-length the shal- lower the slope of the regression line of development rate on temperature and the lower the thermal threshold for development. An analysis of the literature shows that this effect of photoperiod on the thermal requirements for development is widespread among insects but has been overlooked by previous authors. The authors conclude that the variation in the development time observed in insects at different seasons, photoperiods or food regimes, or from different populations, etc., are generally due to some modification of the thermal reaction norms and more specifically to differences in the thermal requirements for development.

It has been established that duration of brood development and success of rearing of the first worker individuals by founder females of the ant Lasius niger from the northern population (St. Petersburg) depend on temperature conditions to a larger extent than of individuals from the southern areal (Borisovka Village, Belgorod Region) The development rate of individuals from the northern population is lower as compared with individuals from the southern population in the low temperature range (17–20°C), but is higher at elevated temperatures (22–27°C). This leads to a greater slope of the temperature regression line of the development rate on the temperature, to lower values of the sum of effective temperatures, and to a higher temperature threshold of development in the northern population. Lower temperatures have a stronger effect on development of larvae from the northern population by inducing diapause in the greater number of individuals as compared with larvae from the southern population. The L. niger founder females require, on average, higher temperatures than the females from the southern populations of this species for rearing the first worker ants during the same summer season (i.e., without winter diapause). Thus, ants of the southern populations, when rearing the first brood, use a strategy of faster development at higher temperatures, while low temperatures restrict to the greater extent the development of their brood as compared with ants from the southern populations. It is concluded that one of the ways of physiological adaptation of ants for inhabitance on the North is such change of the norm of response to temperature, at which development becomes more temperature-dependent, the physiological response to higher temperatures increasing at the cost of its decrease at lower temperatures.

The mean body size of workers of the holarctic ant Leptothorax acervorum increases with latitude. Workers from populations near the Polar Circle were 10% larger than workers from central Europe. This gradient does not appear to be associated with variation in colony size. According to controlled rearing experiments with brood from populations in Cape Kartesh, Karelia (67°N) and Erlangen, Germany (49.7°N), larger adult body size in boreal populations is not an epiphenomenon of slow cell growth and larger cell size at lower temperatures. Larger workers survived longer without food both at room temperature and < 0°C, suggesting that selection for increased fasting endurance in boreal habitats might lead to this Bergmann's rule-like pattern in an ectothermic ant.

Colonies of M. rubra, M. ruginodis and M. scabrinodis were collected in four geographic regions: Kiev, Ukraine (50.5°N, 30.5°E - first two species), Vladimir, Russia (56.2°N, 40.4°E - only last species), St. Petersburg, Russia (59.3°N, 30.3°E - all three species) and Chupa, Murmansk prov., Russia (66.3°N, 33.7°E - last two species). After artificial overwintering experimental cultures consisting of 150 workers and one queen were established and kept at 16, 18, 20, 22, 24 and 26°C under long (22 h) day lengths. The workers reared eggs laid by queens into rapid (non-diapause) brood pupae and diapause larvae, which were removed and counted. The results showed the distinct latitudinal variation in the temperature effects on rapid brood rearing and in the thermal requirements for development. First, the period during which new rapid brood pupae appeared was found to be longer and the total number of pupae produced to be greater in ants from more southern populations. The number of diapause larvae reared by ant cultures was also usually greater, in ants from southern sites. Second, low temperatures reduced the period of rapid brood production and the number of pupae reared to a greater degree in ants from northern populations. It means that northern Myrmica colonies rear rapid brood under lower temperatures evidently worse in comparison with ants from southern regions. Third, eggs and larvae from more northern sites appeared to develop faster than southern brood at temperatures above 16-18°C. This was because brood development in northern populations was more temperature dependent, i.e. characterised by higher slopes of regression lines of development rate on temperature. The sum of effective temperatures decreased with the advance to North. The higher slopes were always associated with higher thermal thresholds for development. We conclude that the reaction norm of Myrmica colonies, in response to temperature, changes according to the local climate in such a way that brood rearing, growth and development of individuals become more temperature dependent in more severe environments with colder and shorter summers. This lead to the increase of the physiological and developmental responses at higher temperatures at the expense of a decrease within lower temperature range. In fact Myrmica colonies from northern populations need on average higher temperatures in their nests for successful production of new adults as compared to southern ants.

The worker ants Myrmica rubra kept at long (18–20 h) days (LD) for 2–3 weeks cause the diapause termination (reactivation) in larvae and queens even under short-day (12 h) conditions (SD). The experiments performed have shown this effect to be due to a distant action of an activating stimulus that has the chemical nature and able to be spread from one laboratory nest with ants to another by means of diffusion through a double perforated net-covered partition or with an artificial air flow. The transfer of this stimulus does not require tactile contacts or food exchange between workers. The chemical nature of the activating stimulus was confirmed by the experiments, in which reactivation was produced by killed LD workers that were ground in a mortar and added twice a day to the nest with diapause ants kept at SD. The detected pheromone is the first distantly perceived primer found in social insects.

Metabolic respiration by groups of resting Myrmica ruginodis and M. scabrinodis worker ants from five sites representing a range of latitudes, have been compared by measuring rates of CO2 production—standardised by fat-free weight—at 5 and 25 °C. M. ruginodis which lives in cooler habitats than M. scabrinodis consistently produced more CO2 .A t 5°C ants of both species from southern latitudes were metabolically more active than those from more northerly latitudes, whereas at 25°C the situation was reversed. Estimates of Q10 were positively correlated with latitude indicating that the respiratory metabolism of northern populations increases relatively more in response to rising temperatures than southern populations. Values of Q10 at different latitudes were the same for both species. The results are discussed in terms of seasonal fluctuations of temperature at different latitudes. © 1999 Published by Elsevier Science Ltd. All rights reserved.

Regression lines of development rate on temperature appeared significantly different between long (22 h) and short (12 h) day conditions and intersected each other at 23.8°С. Thus, the rate of growth and development was higher at temperatures below the intersection point under short-day but above the intersection point it was higher under long day. Ecological relevance of this effect seems as follows: in autumn, as nights become longer and average daily temperature decreases, larvae have to speed up their development because it is only imago that overwinters. Conversely, midsummer offers long days and usually higher temperature, so again it is advantageous to develop as fast as possible in order to have at least one more generation per year. These results are compared with other studies showing interactions between photoperiod and temperature, and some possible general patterns are outlined. The lower thermal threshold for larval development depended on photoperiodic conditions; therefore rate isomorphy must be violated in this species. Development at higher temperatures generally resulted in smaller adults, as is usual with ectotherms according to the "temperature-size rule", but body weight depended significantly on temperature only under short day. Our estimates of the lower temperature thresholds for growth and development in both cases did conform to the generalization made previously by Walters and Hassall (2006) in spite of another formula used by us. We briefly discuss this phenomenon and argue that relative position of these thresholds can be explained mathematically and per se may lack any biological sense.

The thermal norms of egg development were studied in the ground beetles Amara communis, A. nitida, Carabus granulatus, Platynus assimilis, Poecilus versicolor, Pterostichus oblongopunctatus (spring breeding species), P. melanarius, and P. niger (autumn breeding species). The adults were collected in soil traps near Arkhangelsk, St. Petersburg, Moscow, and Bryansk in May–June. Females and males were kept in pairs at 20°C and 22 h light per day. The eggs laid by females were kept at constant temperatures (12, 14, 16, 18, 20, and 22 ± 0.1°C). The development time for each egg was determined accurate to 0.5 day. From the values of the individual development rate (a reciprocal of development time) at all the temperatures, the thermal constants for development were calculated: the coefficient of linear regression (CLR) of the development rate on temperature, the sum of degree-days, and the thermal threshold for development (TTD). The egg development time was found to vary significantly between the species, except for A. communis, A. nitida, and C. granulatus. The values of CLR (i.e., the slopes of the regression lines) and TTD varied rather distinctly between the species and populations that revealed differences in the thermal requirements for egg development. In the spring breeding species the mean egg development time was significantly shorter, and the CLR and TTD values were on average higher than those of the autumn breeding species. Intraspecific latitudinal variation of the development time and the thermal requirements for egg development was revealed. The eggs laid by beetles from the Arkhangelsk population developed faster than those of the same species from the southern populations at all experimental temperatures. These differences were not great but statistically significant. At the same time, the differences between the CLR and TTD values for the northern and southern populations of the same species were non-significant in most cases. Thus, the main ecophysiological adaptation of carabid beetles during their northward expansion was the shortening of development time within the entire range of favorable temperatures, while the CLR and TTD values only insignificantly changed.