Different currencies for calculating resource phenology result in opposite inferences about trophic mismatches.

Shifts in phenology are among the key responses of organisms to climate change. When rates of phenological change differ between interacting species they may result in phenological asynchrony. Studies have found conflicting patterns concerning the direction and magnitude of changes in synchrony, which have been attributed to biological factors. A hitherto overlooked additional explanation are differences in the currency used to quantify resource phenology, such as abundance and biomass. Studying an insectivorous bird (the sanderling) and its prey, we show that the median date of cumulative arthropod biomass occurred, on average, 6.9 days after the median date of cumulative arthropod abundance. In some years this difference could be as large as 21 days. For 23 years, hatch dates of sanderlings became less synchronized with the median date of arthropod abundance, but more synchronized with the median date of arthropod biomass. The currency-specific trends can be explained by our finding that mean biomass per arthropod specimen increased with date. Using a conceptual simulation, we show that estimated rates of phenological change for abundance and biomass can differ depending on temporal shifts in the size distribution of resources. We conclude that studies of trophic mismatch based on different currencies for resource phenology can be incompatible with each other.

Site-specific length-biomass relationships of arctic arthropod families are critical for accurate ecological inferences.

Arthropods play a crucial role in terrestrial ecosystems, for instance in mediating energy fluxes and in forming the food base for many organisms. To better understand their functional role in such ecosystem processes, monitoring of trends in arthropod biomass is essential. Obtaining direct measurements of the body mass of individual specimens is laborious. Therefore, these data are often indirectly acquired by utilizing allometric length-biomass relationships based on a correlative parameter, such as body length. Previous studies have often used such relationships with a low taxonomic resolution and/or small sample size and/or adopted regressions calibrated in different biomes. Despite the scientific interest in the ecology of arctic arthropods, no site-specific family-level length-biomass relationships have hitherto been published. Here we present 27 family-specific length-biomass relationships from two sites in the High Arctic: Zackenberg in northeast Greenland and Knipovich in north Taimyr, Russia. We show that length-biomass regressions from different sites within the same biome did not affect estimates of phenology but did result in substantially different estimates of arthropod biomass. Estimates of daily biomass at Zackenberg were on average 24% higher when calculated using regressions for Knipovich compared to using regressions for Zackenberg. In addition, calculations of daily arthropod biomass at Zackenberg based on order-level regressions from frequently cited studies in literature revealed overestimations of arthropod biomass ranging from 69.7% to 130% compared to estimates based on regressions for Zackenberg. Our results illustrate that the use of allometric relationships from different sites can significantly alter the biological interpretation of, for instance, the interaction between insectivorous birds and their arthropod prey. We conclude that length-biomass relationships should be locally established rather than being based on global relationships.

Population-specific effects of temperature and photoperiod on development and body mass in Cassida vibex (Coleoptera: Chrysomelidae).

An interplay of genetic divergence and phenotypic plasticity in shaping geographic variation is increasingly receiving attention in the entomological literature. Two major environmental variables that govern life histories are temperature and photoperiod. Studies of thermal and photoperiodic reaction norms help us understand how insect diversity evolved and how insects respond to environmental change. We studied survival, development, and body mass in three geographic populations of the beetle Cassida vibex reared in the laboratory under several combinations of constant temperature (16, 19, 22, 25, and 28 °C) and photoperiod (short-day and long-day). The three collection sites are situated along a climatic gradient and separated by hundreds of kilometers. Each population subtly but significantly differs in the absolute values of survival rate, developmental rate, and body mass as well as in the thermal and photoperiodic plasticity of these traits, but the geographic differences do not form a latitudinal cline. The southernmost population from a relatively warm climate survives worse at low temperatures than the other two, but the overall survival is lowest in the latitudinally intermediate population. Short-day conditions tend to accelerate postembryonic development and increase the slope of the developmental rate-temperature relationship, especially so in the intermediate population, followed by the southernmost population and then by the northernmost population. The latter, which inhabits a harsh climate, has the fastest and most temperature-sensitive development, regardless of photoperiod, and attains the largest body mass among the three populations. The intermediately located and photoperiodically plastic population, which lives in a cool but mild climate, in contrast, has the smallest body size. Hence, although the importance of short-day conditions as a seasonal cue increases poleward, the photoperiodic responses do not always become more pronounced in colder, high-latitude environments. Our results emphasize that insect life-history traits can exhibit quite sophisticated patterns of variation along climatic gradients.

Stagewise resolution of temperature-dependent embryonic and postembryonic development in the cowpea seed beetle Callosobruchus maculatus (F.).

BACKGROUND: The thermal plasticity of life-history traits receives wide attention in the recent biological literature. Of all the temperature-dependent traits studied, developmental rates of ectotherms are especially often addressed, and yet surprisingly little is known about embryonic responses to temperature, including changes in the thermal thresholds and thermal sensitivity during early development. Even postembryonic development of many cryptically living species is understood superficially at best. RESULTS: This study is the first to estimate the exact durations of developmental stages in the cowpea seed beetle C. maculatus from oviposition to adult emergence at five permissive constant temperatures from 20 to 32 °C. Early embryonic development was tracked and documented by means of destructive sampling and subsequent confocal imaging of fluorescently stained specimens. Late embryonic and early larval development was studied with the use of destructive sampling and light microscopy. Well-resolved temporal series based on thousands of embryos allowed precise timing of the following developmental events: formation of the blastoderm; formation, elongation, and retraction of the germ band; dorsal closure; the onset and completion of sclerotization of the cuticle; hatching, and penetration of the first-instar larva into the cowpea seed. Pupation and adult eclosion were observed directly through an incision in the seed coat. The thermal phenotype of C. maculatus was found to vary in the course of ontogeny and different stages scaled disproportionately with temperature, but pitfalls and caveats associated with analyses of relative durations of individual stages are also briefly discussed. CONCLUSION: Disproportionate changes in developmental durations with temperature may have important implications when study design requires a high degree of synchronization among experimental embryos or when the occurrence of particular stages in the field is of interest, as well as in any other cases when development times need to be estimated with precision. This work provides one of the first examples of integration of embryological techniques with ecophysiological concepts and will hopefully motivate similar projects in the future. While experiments with Drosophila continue to be the main source of information on animal development, knowledge on other model species is instrumental to building a broader picture of developmental phenomena.

Effects of Temperature and Photoperiod on the Immature Development in Cassida rubiginosa Müll. and C. stigmatica Sffr. (Coleoptera: Chrysomelidae).

Tortoise beetles (Cassida and related genera) are a large cosmopolitan group that includes several pests of agricultural crops and natural enemies of weeds but their biology and ecology remain poorly known. Using a set of environmental chambers, we address simultaneous effects of temperature and photoperiod on immature development and adult body mass in two European species, C. rubiginosa and C. stigmatica. Consistent with its broader distribution range, the former species is less susceptible to low rearing temperatures, develops faster and has a larger body mass than the latter. However, C. rubiginosa seems to be less adapted to late-season conditions as a short-day photoperiod accelerates its immature development to a lesser extent than it does in C. stigmatica, which nevertheless results in greater larval mortality and slightly but significantly smaller adults. By contrast, in C. stigmatica, which is more likely to encounter late-season conditions due to its slower life cycle, short-day acceleration of development is achieved at no cost to survivorship and final body mass. The experiment with C. stigmatica was repeated during two consecutive years with different methods and the main results proved to be well reproducible. In addition, laboratory results for C. rubiginosa agree with field data from literature.

Thermal reaction norms can surmount evolutionary constraints: comparative evidence across leaf beetle species.

One of the leitmotifs of the ecophysiological research on ectotherms is the variation and evolution of thermal reaction norms for biological rates. This long-standing issue is crucial both for our understanding of life-history diversification and for predicting the phenology of economically important species. A number of properties of the organism's thermal phenotype have been identified as potential constraints on the evolution of the rate-temperature relationship. This comparative study addresses several such constraints by testing whether the actual interspecific variation of thermal reaction norms across nearly hundred leaf beetle species agrees with the expected patterns. The results show that developmental rate and its temperature-dependent parameters are similar in closely related species and that the variation pattern depends on the taxonomic scale, the thermal reaction norms being mostly parallel for the representatives of distant subclades but intersecting more often farther down the phylogenetic tree. The parallel shift disagrees with the putative ubiquity of a positive slope-threshold relationship, whereby thermal reaction norms should normally intersect, and even more contradicts with the common-intersection hypothesis. The ability to develop in cooler conditions is not traded off at higher temperatures, which is an exception to the "warmer is better" principle. A comparison of high- and low-quality data indicates that some of these discrepancies with earlier findings may stem from a likely presence of noise in previous analyses, which may have affected the variation patterns observed. Overall, the failure to support the universality of the predicted patterns suggests that the evolution of thermal reaction norms in leaf beetles has largely overcome the hypothesized constraints.

Temperature-dependent development in Chrysomela vigintipunctata (Coleoptera: Chrysomelidae), a stenothermal early-season breeder.

Chrysomela vigintipunctata (Scopoli) is a univoltine leaf beetle commonly encountered on willows across the Palearctic forest zone. The preimaginal development in this species takes place during a short time period, from May to June, because larvae are unable to consume mature leaves of the host plant. Therefore, the diet quality imposes a time constraint, and it was expected that the temperature dependence of development in C. vigintipunctata should be adaptively adjusted to the shortness and cool conditions of the favorable season. It was experimentally determined that this leaf beetle was stenothermal at the larval stage, required 275.5 degree-days above the threshold of 9.0°C for total development from oviposition to adult emergence, and attained greater body mass at lower temperatures. However, in all of these aspects, the thermal ecology of C. vigintipunctata was similar to that of two related multivoltine species, C. populi and C. scripta. The interspecific similarity of thermal reaction norms for development rate and body size suggests that these reaction norms in C. vigintipunctata were unlikely to have been shaped by selection favoring faster development or growth early in the season. The results are discussed in terms of the "ecological fitting" concept, which states that a species may be successful in exploiting novel environments while retaining ecophysiological traits evolved elsewhere.

Egg-hatching synchrony and larval cannibalism in the dock leaf beetle Gastrophysa viridula (Coleoptera: Chrysomelidae).

Females of leaf beetles and many other herbivorous insects lay eggs in coherent batches. Hatchlings emerge more or less simultaneously and often prey on their late-hatching clutchmates. It is not certain, however, whether this synchrony of hatching is a mere by-product of cannibalism or whether an additional synchronizing factor exists. The following simple experiment was aimed at determining the causal relationship between cannibalism and simultaneous larval emergence. Egg clutches of the dock leaf beetle Gastrophysa viridula were split into two halves. These halves were either kept as coherent groups in two separate dishes or, alternatively, only one half remained whole, whereas the other one was divided into single eggs, each of which was incubated in a separate dish. Halving of a clutch into coherent groups only slightly disrupted the synchrony of emergence. The consequence of individual isolation was more dramatic. Half-clutches consisting of disconnected solitary eggs required almost twice as much time for complete emergence of all larvae, which was significantly more than cannibalism as a sole synchronizing factor might explain. Moreover, survival rates were the same in coherent half-clutches (in the presence of cannibalism) and among isolated individuals. This group effect and the small contribution of cannibalism suggest the existence of an additional synchronizing factor. Possible mechanisms underpinning this phenomenon are discussed.

Analyses of Developmental Rate Isomorphy in Ectotherms: Introducing the Dirichlet Regression.

Temperature drives development in insects and other ectotherms because their metabolic rate and growth depends directly on thermal conditions. However, relative durations of successive ontogenetic stages often remain nearly constant across a substantial range of temperatures. This pattern, termed 'developmental rate isomorphy' (DRI) in insects, appears to be widespread and reported departures from DRI are generally very small. We show that these conclusions may be due to the caveats hidden in the statistical methods currently used to study DRI. Because the DRI concept is inherently based on proportional data, we propose that Dirichlet regression applied to individual-level data is an appropriate statistical method to critically assess DRI. As a case study we analyze data on five aquatic and four terrestrial insect species. We find that results obtained by Dirichlet regression are consistent with DRI violation in at least eight of the studied species, although standard analysis detects significant departure from DRI in only four of them. Moreover, the departures from DRI detected by Dirichlet regression are consistently much larger than previously reported. The proposed framework can also be used to infer whether observed departures from DRI reflect life history adaptations to size- or stage-dependent effects of varying temperature. Our results indicate that the concept of DRI in insects and other ectotherms should be critically re-evaluated and put in a wider context, including the concept of 'equiproportional development' developed for copepods.

Photoperiod modifies thermal reaction norms for growth and development in the red poplar leaf beetle Chrysomela populi (Coleoptera: Chrysomelidae).

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°Ð¡. 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.