Temperature-size Rule Research Articles

Substrate availability may limit the response of tropical bacterioplankton biomass to warming

AbstractThe response of heterotrophic bacterioplankton to the addition of macrophytic dissolved organic matter (DOM) and temperature was investigated in the Red Sea. We added 40 μmol C L−1 of leachates obtained from seagrass and mangrove leaves to natural bacterial communities, incubated them at three temperatures (25.5°C found in situ plus 3°C below and above that value) and monitored the microbial and biogeochemical responses over 4 d. Seagrass and mangrove DOM, important allochthonous sources in tropical oligotrophic regions, had distinct chemical characteristics compared to unamended seawater, with mangrove substrates containing comparatively more nitrogen and protein‐like fluorescent DOM than seagrass. Specific growth rates (μ) increased twofold in the seagrass and mangrove treatments (1.0 and 0.8 d−1, respectively) relative to the seawater control (0.4 d−1). The biomass of heterotrophic bacteria generally reflected μ changes, reaching maximum values of 16.8 and 17.3 μg C L−1 in the seagrass and mangrove treatments, respectively, compared to just 2.6 μg C L−1 in seawater. The increase in μ values due to experimental warming followed the metabolic theory of ecology, mostly because of enhanced exoenzymatic activity, while cell size decreased as predicted by the temperature–size rule (mean −3% per °C increase). Although the labile nature of the specific seagrass and mangrove DOM leachates was clearly demonstrated, we conclude that tropical heterotrophic bacteria may have limited capability to increase their biomass as a consequence of future warming, even in the presence of high loadings of macrophytic DOM.

Effect of temperature on sexual size dimorphism during the developmental period in the broad-horned flour beetle

Adult size in numerous insects is strongly dependent on temperature. In several cases, a temperature–size rule is observed in which developmental temperature and adult size tradeoff. Although several previous studies have demonstrated the temperature–size rule, only a few have explored the relationship between developmental temperature and weapon traits or sexual size dimorphism. This study was conducted to investigate the size of the broad-horned flour beetle Gnatocerus cornutus when it was developed under different temperatures. G. cornutus males possess weapon traits for male–male combat and exhibit sexual size dimorphism in other morphological traits. Results showed that male weapon size and body size complied with the temperature–size rule. Furthermore, the extent of sex dimorphism in genae width, a weapon-supportive trait, were larger at lower temperatures. Our findings suggest that the temperature–size rule also influences the size of sexual traits.

The temperature-size rule in the context of Dynamic Energy Budget theory

The temperature-size rule (TSR) refers to the observation that a wide variety of organisms reach smaller sizes under warmer temperatures. Although various hypotheses have been stated about the mechanisms underlying TSR, a general understanding is still lacking. In this context, it seems that Dynamic Energy Budget (DEB) theory could be useful, as it constitutes a mechanistic framework that predicts growth and development from individual energetics. However, a critical assumption of this framework is that temperature affects all physiological rates equally, which prevents it from capturing the TSR. It remains an open question if and how the DEB framework can be used to capture the TSR patterns.The model does, however, include a parameter, the food searching rate, that does not need to obey this constraint, as it is a physical rather than a physiological parameter. Here, we aimed to test whether introducing a differing temperature-dependence of food searching rate in DEB models makes it possible to capture the TSR. We parameterized the model using an extensive dataset of Daphnia life history response to temperature. The resulting model was able to capture the decrease of both size at maturity and asymptotic size observed in this dataset.Therefore, DEB models can capture the TSR without any major modification, but only requires to introduce a different function for the temperature dependence of the food searching rate. The mechanism underlying this effect essentially results from increased food limitation at high temperature, which, in combination with the particular allocation scheme of the DEB model, affects differentially growth and development.

Temperature-size responses during ontogeny are independent of progenitors' thermal environments.

Warming generally induces faster developmental and growth rates, resulting in smaller asymptotic sizes of adults in warmer environments (a pattern known as the temperature-size rule). However, whether temperature-size responses are affected across generations, especially when thermal environments differ from one generation to the next, is unclear. Here, we tested temperature-size responses at different ontogenetic stages and in two consecutive generations using two soil-living Collembola species from the family Isotomidae: Folsomia candida (asexual) and Proisotoma minuta (sexually reproducing). We used individuals (progenitors; F0) from cultures maintained during several generations at 15 °C or 20 °C, and exposed their offspring in cohorts (F1) to various thermal environments (15 °C, 20 °C, 25 °C and 30 °C) during their ontogenetic development (from egg laying to first reproduction; i.e., maturity). We measured development and size traits in the cohorts (egg diameter and body length at maturity), as well as the egg diameters of their progeny (F2). We predicted that temperature-size responses would be predominantly determined by within-generation plasticity, given the quick responsiveness of growth and developmental rates to changing thermal environments. However, we also expected that mismatches in thermal environments across generations would constrain temperature-size responses in offspring, possibly due to transgenerational plasticity. We found that temperature-size responses were generally weak in the two Collembola species, both for within- and transgenerational plasticity. However, egg and juvenile development were especially responsive at higher temperatures and were slightly affected by transgenerational plasticity. Interestingly, plastic responses among traits varied non-consistently in both Collembola species, with some traits showing plastic responses in one species but not in the other and vice versa. Therefore, our results do not support the view that the mode of reproduction can be used to explain the degree of phenotypic plasticity at the species level, at least between the two Collembola species used in our study. Our findings provide evidence for a general reset of temperature-size responses at the start of each generation and highlight the importance of measuring multiple traits across ontogenetic stages to fully understand species' thermal responses.

Diet Affects the Temperature-Size Relationship in the Blowfly Aldrichina grahami.

In warmer environments, most ectotherms exhibit a plastic reduction in body size (the temperature-size rule, TSR). However, in such environments, growth is usually accelerated and would be expected to result in maturation at a larger body size, leading to increases in fecundity, survival, and mating success, compared to maturation at a smaller size (the 'life-history puzzle'). To explore these mechanisms, we reared Aldrichina grahami at 20 °C, 25 °C, and 30 °C, and added a nutritional challenge by using dilutions of pork liver paste to provide diets that ranged in quality from high (undiluted) to moderate (1/8), low (1/16), and poor (1/24). Larvae were randomly sampled for weighing from hatching. Growth curves were fitted to the relationships between growth rate and weight for the third instar larvae. Our results showed that body size was affected by an interaction between temperature and diet, and that following or not following the TSR can vary depending on underfeeding. Moreover, when the TSR was followed as temperature increased, there was a cross-over point that divided the two growth curves into early and later stages, which could be used to help understand the life-history puzzle in warmer temperatures, with the instantaneous growth rate being faster in the early stages of development and then slower in later stages. This study reminds us that animals have evolved to cope with multiple simultaneous environmental changes, and it has thus offered a better understanding of life-history puzzles.

Voltinism and life‐history traits of the invasive fall armyworm Spodoptera frugiperda (Lepidoptera: Noctuidae) feeding on corn

AbstractVoltinism and life‐history traits of the invasive fall armyworm (FAW) Spodoptera frugiperda were investigated under semi‐natural conditions for a period of 2 years. The FAW invaded the corn field in the suburbs of Nanchang (28°46′ N, 115°50′ E) in early summer and produced six complete generations. FAW had the characteristics of short developmental time, high survival rates and strong fecundity. The development time of female pupae was significantly faster than that of male pupae, resulting in the emergence of female pupae earlier than male pupae. Except for the sixth generation in 2021, there was no significant difference between female and male sex ratio, which was close to 1:1. FAW showed male‐bias sexual size dimorphism with male pupae being significantly larger than female pupae. Unlike pupal weight, in most generations, male adult weighed significantly less than females, because the weight loss of male pupae during metamorphosis was significantly greater than that of female pupae. The temporal variation of pupal weight did not conform to the temperature–size rule. Compared with 22.8°C, the 29.2°C high temperature not only significantly reduced the development time of larvae but also significantly increased pupal weight. The adult fecundity feeding on fresh corn leaves was higher than that feeding on live corn plants in most generations. In most generations, pupal weight was positively correlated with larval development time and adult weight was positively correlated with fecundity. In conclusion, climate differences between generations and years have significant effects on developmental time, body weight, sexual size dimorphism and fecundity of the FAW. These results add to the understanding of the evolution of life‐history traits in the FAW and may have important implications for predicting population dynamics of the FAW and optimising control strategies.

Does the temperature–size rule apply to idiobiont parasitoids?

AbstractIn most ectotherms, several life‐history traits, including body size, respond to environmental conditions through the temperature–size rule (TSR). The mechanisms underlying the TSR are still being debated, but studying idiobiont insect parasitoids, which develop with a fixed amount of resources, may shed light on this relationship. In this study, we conducted experiments to determine how the developmental temperature affects various characteristics of male and female Trichogramma euproctidis (Girault) (Hymenoptera: Trichogrammatidae), an idiobiont egg parasitoid of Lepidoptera. The study tested various hypotheses, including the cellular or oxygen diffusion hypotheses and the resource acquisition hypothesis, to understand whether T. euproctidis follows the TSR. The developmental time of both male and female T. euproctidis decreased with increasing temperature. Both males and females displayed a unimodal distribution for size, dry mass, and lipid content, with individuals at lower and higher temperatures being smaller, weighing less, and containing fewer lipids. Female lifetime fecundity increased from 13 to 24 °C and then decreased at 31 °C. Additionally, the number and size of gametes in male and female T. euproctidis displayed a unimodal distribution with increasing temperature. Trichogramma euproctidis deviates from the TSR as it follows a non‐linear reaction norm with an optimal developmental temperature. This result supports the hypothesis that for species following TSR and having unlimited access to food resources, the resource acquisition hypothesis is a significant mechanism explaining the TSR. With climate change affecting temperature, understanding the TSR is crucial, and research on insect parasitoids may help reveal how the interplay between environmental temperature and resource allocation affects the TSR in natural populations.

Fish weight reduction in response to intra‐ and interspecies competition under climate change

AbstractAs described by the temperature–size rule paradigm, fish living in warmer temperatures grow faster but have a smaller mature body size. However, the changes in the body size of fish communities in the western North Pacific, which is one of the most active fishing grounds, remain unclear. This study aimed to investigate changes in the body size of fish assemblages in the western North Pacific and whether fish sizes were potentially driven by the temperature–size rule, bottom‐up effects and intra‐ and interspecies competition at a community scale. We evaluated the fish weight data of 6 stocks of 4 species from 1978 to 2018 and 17 stocks of 13 species from 1995/1997 to 2018. Weight reduction in the fish assemblage was observed in the 1980s and was associated with the biomass peak of the Japanese sardine (Sardinops melanostictus), indicating the effect of intra‐ and interspecies competition. Another weight reduction was observed in the 2010s, which was associated with a moderate increase in the biomass of the Japanese sardine and chub mackerel (Scomber japonicus). Our analyses indicate that stronger stratifications in the surface layers during the 2010s potentially reduced the nutrient supply from the subsurface to the surface layers. This limitation in food availability forced intra‐ and interspecies competition under a moderate increase in fish biomass. Our findings underscore the critical significance of integrating the impacts of species competition and climate change on fish sizes to improve fishery management at a community level.

Multispecies population-scale emergence of climate change signals in an ocean warming hotspot

Abstract Ocean waters of the Northeast US continental shelf have warmed rapidly in recent years, with sea surface temperatures rising 2.5 times faster than those of the global oceans. With this strong warming trend, the frequency and duration of marine heatwaves have increased. These temperature changes stood out as a distinct warm temperature regime during the 2010s. During this decade, fish population characteristics also differed from the past. Species distribution shifts were detected for many species, demonstrating one way species could adapt to warming conditions. However, for most species, distribution shifts were insufficient to avoid warmer surface or bottom temperatures. As species occupied warmer habitats, growth patterns aligned with expectations for warming temperatures. Consistent with the temperature-size rule, some species exhibited faster growth at early life stages but plateaued at smaller body sizes; other species, however, experienced reduced growth across all ages, indicating thermal stress. Finally, population productivity indexed by the recruit-to-spawner ratio declined significantly during the 2010s for some populations. Changes in these three processes—distribution, growth, and productivity—indicate the emergence of climate change signals across multiple Northeast US fish populations. These effects create new challenges for fishery managers and industry participants operating in the context of non-stationarity and uncertainty.

Breaking the Law: Is It Correct to Use the Converse Bergmann Rule in Ceroglossus chilensis? An Overview Using Geometric Morphometrics.

The converse Bergmann's rule is a pattern of body size variation observed in many ectothermic organisms that contradicts the classic Bergmann's rule and suggests that individuals inhabiting warmer climates tend to exhibit larger body sizes compared to those inhabiting colder environments. Due to the thermoregulatory nature of Bergmann's rule, its application among ectotherms might prove to be more complicated, given that these organisms obtain heat by absorbing it from their habitat. The existence of this inverse pattern therefore challenges the prevailing notion that larger body size is universally advantageous in colder climates. Ceroglossus chilensis is a native Chilean beetle that has the largest latitudinal range of any species in the genus, from 34.3° S to 47.8° S. Within Chile, it continuously inhabits regions extending from Maule to Aysen, thriving on both native and non-native forest species. Beyond their remarkable color variation, populations of C. chilensis show minimal morphological disparity, noticeable only through advanced morphological techniques (geometric morphometrics). Based on both (1) the "temperature-size rule", which suggests that body size decreases with increasing temperature, and (2) the reduced resource availability in high-latitude environments that may lead to smaller body sizes, we predict that C. chilensis populations will follow the converse Bergmann's rule. Our results show a clear converse pattern to the normal Bergmann rule, where smaller centroid sizes were found to be measured in the specimens inhabiting the southern areas of Chile. Understanding the prevalence of the converse Bergmann's rule for ectotherm animals and how often this rule is broken is of utmost importance to understand the underlying mechanisms allowing organisms to adapt to different environments and the selective pressures they face.

Temperature seasonality drives taxonomic and functional homogenization of tropical butterflies

AbstractAimTo better understand the potential impact of climate change on butterfly assemblages across a tropical island, we model the potential for taxonomic and functional homogenization and determine climate‐ and trait‐mediated shifts in projected species distributions.LocationPuerto Rico.MethodsWe used thousands of museum records of diurnal Lepidoptera to model current (1970–2000) and forecast future (2061–2080) species distributions and combined these to test for taxonomic and functional homogenization. We then quantified climatic‐mediated effects on current and forecasted taxonomic and functional composition and, specifically, whether temperature was a primary driver, as predicted by the temperature–size rule and the thermal melanism hypotheses. Finally, we measured wing traits important in thermoregulation (size and colour) and determined trait‐mediated changes in forecasted species distributions over time.ResultsBased on ensemble model outputs, taxonomic and functional richness and turnover were predicted to vary across the island's complex topography. Our models projected an increase in taxonomic and functional richness over time, and a decrease in taxonomic and functional turnover – a signature of biotic homogenization. Under future climate scenarios, models projected a decrease in wing length and an increase in wing brightness at higher elevations. One variable, temperature seasonality, was the strongest predicted driver of both the current spatial distribution and the projected per cent change over time for not only wing traits but also taxonomic and functional richness and turnover.Main conclusionsThe species distribution models generated here identify several priority regions and species for future research and conservation efforts. Our work also highlights the role of seasonality and climatic variability on diverse tropical Lepidoptera assemblages, suggesting that climatic variability may be an important, albeit overlooked, driver of climate change responses.

Distinct impacts of feeding frequency and warming on life history traits affect population fitness in vertebrate ectotherms.

Body size shifts in ectotherms are mostly attributed to the Temperature Size Rule (TSR) stating that warming speeds up initial growth rate but leads to smaller size when food does not limit growth. Investigating the links between temperature, growth, and life history traits is key to understand the adaptive value of TSR, which might be context dependent. In particular, global warming can affect food quantity or quality which is another major driver of growth, fecundity, and survival. However, we have limited information on how temperature and food jointly influence life history traits in vertebrate predators and how changes in different life history traits combine to influence fitness and population demography. We investigate (1) whether TSR is maintained under different food conditions, (2) if food exacerbates or dampens the effects of temperature on growth and life history traits and (3) if food influences the adaptive value of TSR. We combine experiments on the medaka with Integral Projection Models to scale from life history traits to fitness consequences. Our results confirm that warming triggers a higher initial growth rate and a lower adult size, reduces generation time and increases mean fitness. A lower level of food exacerbates the effects of warming on growth trajectories. Although lower feeding frequency increased survival and decreased fecundity, it did not influence the effects of warming on fish development rates, fecundity, and survival. In contrast, feeding frequency influenced the adaptive value of TSR, as, under intermittent feeding, generation time decreased faster with warming and the increase in growth rate with warming was weaker compared to continuously fed fish. These results are of importance in the context of global warming as resources are expected to change with increasing temperatures but, surprisingly, our results suggest that feeding frequency have a lower impact on fitness at high temperature.

Supplemental oxygen does not improve growth but can enhance reproductive capacity of fish.

Fish tend to grow faster as the climate warms but attain a smaller adult body size following an earlier age at sexual maturation. Despite the apparent ubiquity of this phenomenon, termed the temperature-size rule (TSR), heated scientific debates have revealed a poor understanding of the underlying mechanisms. At the centre of these debates are prominent but marginally tested hypotheses which implicate some form of 'oxygen limitation' as the proximate cause. Here, we test the role of oxygen limitation in the TSR by rearing juvenile Galaxias maculatus for a full year in current-day (15°C) and forecasted (20°C) summer temperatures while providing half of each temperature group with supplemental oxygen (hyperoxia). True to the TSR, fish in the warm treatments grew faster and reached sexual maturation earlier than their cooler conspecifics. Yet, despite supplemental oxygen significantly increasing maximum oxygen uptake rate, our findings contradict leading hypotheses by showing that the average size at sexual maturation and the adult body size did not differ between normoxia and hyperoxia groups. We did, however, discover that hyperoxia extended the reproductive window, independent of fish size and temperature. We conclude that the intense resource investment in reproduction could expose a bottleneck where oxygen becomes a limiting factor.

Body size and trophic level increase with latitude, and decrease in the deep-sea and Antarctica, for marine fish species.

The functional traits of species depend both on species' evolutionary characteristics and their local environmental conditions and opportunities. The temperature-size rule (TSR), gill-oxygen limitation theory (GOLT), and temperature constraint hypothesis (TCH) have been proposed to explain the gradients of body size and trophic level of marine species. However, how functional traits vary both with latitude and depth have not been quantified at a global scale for any marine taxon. We compared the latitudinal gradients of trophic level and maximum body size of 5,619 marine fish from modelled species ranges, based on (1) three body size ranges, <30, 30-100, and >100 cm, and (2) four trophic levels, <2.20, 2.20-2.80, 2.81-3.70, >3.70. These were parsed into 5° latitudinal intervals in four depth zones: whole water column, 0-200, 201-1,000, and 1,001-6,000 m. We described the relationship between latitudinal gradients of functional traits and salinity, sea surface and near seabed temperatures, and dissolved oxygen. We found mean body sizes and mean trophic levels of marine fish were smaller and lower in the warmer latitudes, and larger and higher respectively in the high latitudes except for the Southern Ocean (Antarctica). Fish species with trophic levels ≤2.80 were dominant in warmer and absent in colder environments. We attribute these differences in body size and trophic level between polar regions to the greater environmental heterogeneity of the Arctic compared to Antarctica. We suggest that fish species' mean maximum body size declined with depth because of decreased dissolved oxygen. These results support the TSR, GOLT and TCH hypotheses respectively. Thus, at the global scale, temperature and oxygen are primary factors affecting marine fishes' biogeography and biological traits.

Conodont size in time and space: Beyond the temperature-size rule

The temperature-size rule (TSR) states that ectotherms mature at smaller adult body size in warmer conditions. Such a rule may have the potential to explain size response of fossil organisms to past temperature variations, but its validity in deep time has been seldom tested. The generality of this rule was investigated here by compiling data documenting the size record of three conodont genera (Palmatolepis, Ancyrodella and Polygnathus) at different spatial and temporal scales during the Late Frasnian and the Famennian, characterized by short- and long-term temperature variations. Statistical models were used to investigate the relationship between conodont size and oxygen isotope values, considered as paleotemperature proxies. Comparison of evolutionary models further allowed to test alternative modes of size variation such as stasis or punctuation.The TSR was not validated as a general rule explaining size variation in these fossil records, being only observed as a large-scale geographic trend during a time-slice. The only strong support for temperature being the driver of temporal variations was found regarding the size of Palmatolepis during the Kellwasser period, but the relationship was reverse to the expectation of the TSR. The absence of general TSR pattern is probably due to the interference of many other factors (demography and mortality patterns, temperature tolerance, size reduction due to stress) whose relative importance may depend on the time interval and the genus considered. Rather than a correlation with environmental proxies, evolutionary models suggested the occurrence of a synchronous shift in Palmatolepis size around 369 Ma (Palmatolepis termini conodont Zone) in several outcrops, raising questions about the environmental forcing beyond this shift. Departures from the expected TSR may thus provide relevant insights into the complex interplay of physical, tectonic and eco-evolutionary processes impacting size evolution in deep time.

How temperature affects the body size of terrestrial tardigrades

Abstract Many vertebrates, both homeo- and poikilothermic, show a significant relationship between body size and environmental temperature. Whether such an association may exist in microscopic invertebrates has been less explored. Therefore, we decided to analyse terrestrial Tardigrada from various habitats worldwide to examine whether these animals reveal any relationship pattern between body size and environmental temperature. Data on minimum, maximum, and mean body sizes were extracted from original descriptions or sometimes from later re-descriptions of the species. Minimum, maximum, and mean temperature data from the type localities of the species were retrieved from WorldClim 2. In general, accounting for geographic and phylogenetic confounding factors, the body size of terrestrial tardigrades decreased as the environmental temperature increased. The same tendency was observed for most of the genera when additional analyses were carried out separately for each genus. This is the first biogeographical analysis demonstrating that terrestrial tardigrades generally conform to the temperature–size rule.

Mechanisms of increasing predation by planktivorous fish with rising temperature may explain the temperature–body size relationships in zooplankton

The temperature–size rule (TSR) has been consistently observed in numerous studies, showing that ectotherms reared at higher temperatures experience accelerated growth during the juvenile stage and ultimately reach smaller sizes and younger ages at maturity. One explanation for this response is that it occurs when the effect of temperature on mortality, including predation, outweighs its effect on food intake and metabolism. While several studies have found that the latter effect is close to the expected result based on the Q10 = 2 assumption, confirmation of this hypothesis requires evidence that the effect of temperature on mortality exceeds the Q10 = 2 threshold. To test this hypothesis, we conducted experiments with two fish species: rudd and Malabar danio. We examined the capture rate, which serves as a proxy for mortality, as well as the standard metabolic rate (SMR) and several parameters characterising the mobility of the fish and their planktonic prey (Daphnia) at different temperatures. The results strongly supported our hypothesis, as the capture rate increased significantly more than expected based on the Q10 = 2 assumption, especially for the danio. This substantial effect cannot be attributed solely to the thermal sensitivity of the SMR, as the Q10 for the SMR was only around 2. The most likely explanation seems to be a much more pronounced increase in the fish’s mobility and resulting reaction field volume compared to its planktonic prey at elevated temperatures. This increased mobility leads to an improved attack rate by the fish, which exceeds the prediction made by the Q10 = 2 assumption. This mechanism may explain not only the TSR pattern in zooplankton, but also their reduced mean body size and density at population and community levels at elevated temperatures, and may hypothetically be observed at other predator–prey interfaces.

Body mass decline in a Mediterranean community of solitary bees supports the size shrinking effect of climatic warming.

The long-known, widely documented inverse relationship between body size and environmental temperature ("temperature-size rule") has recently led to predictions of body size decline following current climatic warming ("size shrinking effect"). For keystone pollinators such as wild bees, body shrinking in response to warming can have significant effects on pollination processes but there is still little direct evidence of the phenomenon because adequate tests require controlling for confounding factors linked to climate change (e.g., habitat change). This paper assesses the shrinking effect in a community of solitary bees from well-preserved habitats in the core of a large nature reserve experiencing climatic warming without disturbances or habitat changes. Long-term variation in mean body mass was evaluated using data from 1704 individual bees (137 species, 27 genera, 6 families) sampled over 1990-2023. Climate warmed at a fast rate during this period, annual mean of daily maximum temperature increasing 0.069°C/year on average during 2000-2020. Changes in bee body mass verified expectations from the size shrinking effect. The mean individual body mass of the community of solitary bees declined significantly, irrespective of whether the analysis referred to the full species sample or only to the subset of species that were sampled in both the old (1990-1997) and recent (2022-2023) periods. On average, body mass declined ~0.7%·year-1 , or an estimated average cumulative reduction of ~20 mg per individual bee from 1990 to 2023. Proportional size reduction was greatest among large-bodied species, ranging from around -0.6%·year-1 for the smallest species to -0.9%·year-1 for the largest ones. Declining rate was steeper for cavity-nesting than ground-nesting species. The pollination and mating systems of bee-pollinated plants in the study region are probably undergoing significant alterations as a consequence of supra-annual decline in bee body mass.

Exceptions to the temperature–size rule: no Lilliput Effect in end‐Permian ostracods (Crustacea) from Aras Valley (northwest Iran)

AbstractThe body size of marine ectotherms is often negatively correlated with ambient water temperature, as seen in many clades during the hyperthermal crisis of the end‐Permian mass extinction (c. 252 Ma). However, in the case of ostracods, size changes during ancient hyperthermal events are rarely quantified. In this study, we evaluate the body size changes of ostracods in the Aras Valley section (northwest Iran) in response to the drastic warming during the end‐Permian mass extinction at three taxonomic levels: class, order, species. At the assemblage level, the warming triggers a complete species turnover in the Aras Valley section, with larger, newly emerging species dominating the immediate post‐extinction assemblage for a short time. Individual ostracod species and instars do not show dwarfing or a change in body size as an adaptation to the temperature stress during the end‐Permian crisis. This may indicate that the ostracods in the Aras Valley section might have been exceptions to the temperature–size rule (TSR), using an adaptation mechanism that does not involve a decrease in body size. This adaptation might be similar to the accelerated development despite constant instar body sizes that can be observed in some recent experimental studies of ostracod responses to thermal stress.

Does the temperature–size rule apply to marine protozoans?

Abstract The temperature–size rule suggests that there is a negative relationship between the size (volume) of an ectothermic organism and the environmental temperature experienced during its development We question how to validate this hypothesis for the particular case of some algivorous protozoans because in many of these unicellular organisms, body size is directly related to the consumption of prey and, on most occasions, the true size of the cell is uncertain. In our opinion, to approach this question, the actual size of the protozoan should be measured when the prey are fully digested. To prove our arguments, we designed a series of experiments with the heterotrophic dinoflagellate Oxyrrhis marina, including functional and numerical responses, time‐dependent acclimation responses and the estimation of the protozoan volume during absence of food. We found that after the digestion of the prey Rhodomonas salina, the size of our long‐term thermally acclimated strains of Oxyrrhis marina was the same regardless of the temperature (16 and 22°C). We believe that previous reports showing protozoan size reduction because of temperature might actually reflect imbalances between ingestion and digestion of prey, perhaps amplified by insufficient acclimation. In support of the previous argument, we found evidence that short‐term temperature exposure experiments may suggest a negative relationship between well‐fed protozoan size and temperature, although this was not observed after long‐term acclimation. In light of the results presented here, we suggest that the actual evidence supporting the temperature–size rule in algivorous protozoans may not be sufficiently conclusive because most of the present studies largely ignore the effect of prey presence on the grazer's volume and are usually based on short‐term responses to temperature. Read the free Plain Language Summary for this article on the Journal blog.