Two major intraspecific patterns of adult size variation are plastic temperature-size (T-S) responses and latitude-size (L-S) clines. Yet, the degree to which these co-vary and share explanatory mechanisms has not been systematically evaluated. We present the largest quantitative comparison of these gradients to date, and find that their direction and magnitude co-vary among 12 arthropod orders (r(2) = 0.72). Body size in aquatic species generally reduces with both warming and decreasing latitude, whereas terrestrial species have much reduced and even opposite gradients. These patterns support the prediction that oxygen limitation is a major controlling factor in water, but not in air. Furthermore, voltinism explains much of the variation in T-S and L-S patterns in terrestrial but not aquatic species. While body size decreases with warming and with decreasing latitude in multivoltine terrestrial arthropods, size increases on average in univoltine species, consistent with predictions from size vs. season-length trade-offs.

Body size is central to ecology at levels ranging from organismal fecundity to the functioning of communities and ecosystems. Understanding temperature‐induced variations in body size is therefore of fundamental and applied interest, yet thermal responses of body size remain poorly understood. Temperature–size (T–S) responses tend to be negative (e.g. smaller body size at maturity when reared under warmer conditions), which has been termed the temperature–size rule (TSR). Explanations emphasize either physiological mechanisms (e.g. limitation of oxygen or other resources and temperature‐dependent resource allocation) or the adaptive value of either a large body size (e.g. to increase fecundity) or a short development time (e.g. in response to increased mortality in warm conditions). Oxygen limitation could act as a proximate factor, but we suggest it more likely constitutes a selective pressure to reduce body size in the warm: risks of oxygen limitation will be reduced as a consequence of evolution eliminating genotypes more prone to oxygen limitation. Thus, T–S responses can be explained by the ‘Ghost of Oxygen‐limitation Past’, whereby the resulting (evolved) T–S responses safeguard sufficient oxygen provisioning under warmer conditions, reflecting the balance between oxygen supply and demands experienced by ancestors. T–S responses vary considerably across species, but some of this variation is predictable. Body‐size reductions with warming are stronger in aquatic taxa than in terrestrial taxa. We discuss whether larger aquatic taxa may especially face greater risks of oxygen limitation as they grow, which may be manifested at the cellular level, the level of the gills and the whole‐organism level. In contrast to aquatic species, terrestrial ectotherms may be less prone to oxygen limitation and prioritize early maturity over large size, likely because overwintering is more challenging, with concomitant stronger end‐of season time constraints. Mechanisms related to time constraints and oxygen limitation are not mutually exclusive explanations for the TSR. Rather, these and other mechanisms may operate in tandem. But their relative importance may vary depending on the ecology and physiology of the species in question, explaining not only the general tendency of negative T–S responses but also variation in T–S responses among animals differing in mode of respiration (e.g. water breathers versus air breathers), genome size, voltinism and thermally associated behaviour (e.g. heliotherms).

Major biological and biogeographical rules link body size variation with latitude or environmental temperature, and these rules are often studied in isolation. Within multivoltine species, seasonal temperature variation can cause substantial changes in adult body size, as subsequent generations experience different developmental conditions. Yet, unlike other size patterns, these common seasonal temperature-size gradients have never been collectively analysed. We undertake the largest analysis to date of seasonal temperature-size gradients in multivoltine arthropods, including 102 aquatic and terrestrial species from 71 global locations. Adult size declines in warmer seasons in 86% of the species examined. Aquatic species show approximately 2.5-fold greater reduction in size per °C of warming than terrestrial species, supporting the hypothesis that greater oxygen limitation in water than in air forces aquatic species to exhibit greater plasticity in body size with temperature. Total percentage change in size over the annual cycle appears relatively constant with annual temperature range but varies between environments, such that the overall size reduction in aquatic-developing species (approx. 31%) is almost threefold greater than in terrestrial species (approx. 11%). For the first time, we show that strong correlations exist between seasonal temperature-size gradients, laboratory responses and latitudinal-size clines, suggesting that these patterns share common drivers.

Aim Body size is a master trait with significant ecological importance. Seasonal changes in body size within diverse ectothermic species can result from different environmental conditions experienced during ontogeny in subsequent generations. Whilst intraspecific changes in adult size have been well studied under controlled experimental conditions and across geographical ranges, comprehensive analyses of temporal changes are lacking, and there remains considerable unexplained variation in body size responses within aquatic taxa. Using planktonic copepods as an exemplar taxon, we quantify variation in adult body mass within seasonally varying marine and freshwater environments. We describe how size variation relates to temperature, food concentration (chlorophyll-a) and life-history characteristics, including feeding strategy.Location Global.Methods Using a meta-analytic approach we extract quantitative data from published literature on seasonal size responses of copepods. We analyse competing models to determine the best predictors of these responses, and compare the relative importance of temperature and chlorophyll-a concentration in explaining variation in body size. ResultsWe quantify 140 seasonal size responses from 33 different global locations, representing 48 planktonic copepod species from four taxonomic orders. We find that temperature (r 2 5 0.50), rather than food (r 2 5 0.22), is the dominant explanatory variable of changes in adult body size across seasons. A striking outcome is that calanoid copepods, which utilize feeding currents to capture prey, exhibit a four-fold greater reduction in adult body mass per 8C (23.66%) compared with cyclopoid copepods (20.91%), which are ambush feeders. By contrast, species body size or reproductive strategy did not explain variation in the seasonal temperature-size response.Main conclusions Our findings lead us to suggest that feeding strategies may play a significant role in dictating the magnitude of seasonal temperature-size responses in copepods, with potential implications for other ectotherms with diverse feeding methods. Seasonal temperature-size responses were typically much more variable than responses in laboratory studies that provided excess food, suggesting that field conditions modify the temperature-size response.

Sexual size dimorphism (SSD) is often affected by environmental conditions, but the effect of temperature on SSD in ectotherms still requires rigorous investigation. We compared the plastic responses of size-at-maturity to temperature between males and females within 85 diverse arthropod species, in which individuals of both sexes were reared through ontogeny under identical conditions with excess food. We find that the sexes show similar relative (proportional) temperature-body size (T-S) responses on average. The high degree of similarity occurs despite an analysis that includes a wide range of animal body sizes, variation in degree of SSD and differences in the sign of the T-S response. We find no support for Rensch's rule, which predicts greater variation in male size, or indeed the reverse, greater female size variation. SSD shows no systematic temperature dependence in any of the 17 arthropod orders examined, five of which (Diptera, Orthoptera, Lepidoptera, Coleoptera and Calanoida) include more than six thermal responses. We suggest that the same proportional T-S response may generally have equivalent fitness costs and benefits in both sexes. This contrasts with effects of juvenile density, and food quantity/quality, which commonly result in greater size plasticity in females, suggesting these variables have different adaptive effects on SSD.