Temperature,size, reproductive allocation,and life‐history evolution in a gregarious caterpillar

The present study aimed to investigate the relationship between growth rate, final mass, and larval development, as well as how this relationship influences reproductive trade‐offs, in the context of a gregarious life‐style and the need to keep an optimal group size. We use as a model two sympatric populations of the pine processionary moth Thaumetopoea pityocampa, which occur in different seasons and thus experience different climatic conditions. Thaumetopoea pityocampa is a strictly gregarious caterpillar throughout the larval period, which occurs during winter in countries all over the Mediterranean Basin. However, in 1997, a population in which larval development occurs during the summer was discovered in Portugal, namely the summer population (SP), as opposed to the normal winter population (WP), which coexists in the same forest feeding on the same host during the winter. Both populations were monitored over 3 years, with an assessment of the length of the larval period and its relationship with different climatic variables, final mass and adult size, egg size and number, colony size, and mortality at different life stages. The SP larval period was reduced as a result of development in the warmer part of the year, although it reached the same final mass and adult size as the WP. Despite an equal size at maturity, a trade‐off between egg size and number was found between the two populations: SP produced less but bigger eggs than WP. This contrasts with the findings obtained in other Lepidoptera species, where development in colder environments leads to larger eggs at the expense of fecundity, but corroborates the trend found at a macro‐geographical scale for T. pityocampa, with females from northern latitudes and a colder environment producing more (and smaller) eggs. The results demonstrate the importance of the number of eggs in cold environments as a result of an advantage of large colonies when gregarious caterpillars develop in such environments, and these findings are discussed in accordance with the major theories regarding size in animals. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 340–349.     

Heritability of life‐history traits in the spruce budworm

The heritability of life‐history traits is of particular importance for insects that are very dependent on host conditions. Severe defoliation caused by the spruce budworm negatively impacts its food source, which in turn imposes environmental constraints on the insect. The heritability of those traits can help elucidate this species' evolutionary process. Heritability also helps identify which traits exhibit significant additive variance and can be key to understanding natural selection effects. Individuals were reared under laboratory conditions over three generations on an artificial diet. Heritability was estimated by parent–offspring regression. Fertility and fecundity demonstrated significant heritability followed by larval development, while pupal mass showed minimal heritable variation. These results suggest an important percent of additive variance in life‐history traits. This study contributes to our understanding of the relationship of this forest pest to its environmental conditions. This study also reveals an important genetic architectural structure of life‐history traits in the spruce budworm.     

Characterization of five microsatellite loci in the Pine Processionary Moth Thaumetopoea pityocampa (Lepidoptera Notodontidae Thaumetopoeinae)

Five microsatellite markers were developed for the lepidopteran species Thaumetopoea pityocampa using an enrichment protocol. All loci could be amplified with no evidence of null alleles and will be useful for population genetic studies. The number of alleles ranged from three to 12 for a population of 30 individuals. Observed heterozygosities ranged from 0.53 to 0.80. No significant heterozygote deficiency was detected. Four markers might be of interest for Th. wilkinsoni.     

A large‐scale latitudinal pattern of life‐history traits in a strictly univoltine damselfly

1. Variation in thermal conditions and season length along latitudinal gradients affect body size‐related traits over different life stages. Selection is expected to optimise these size traits in response to the costs and benefits. 2. Egg, hatchling, larval and adult size in males and females were estimated along a latitudinal gradient of 2730 km across Europe in the univoltine damselfly Lestes sponsa, using a combination of field‐collection and laboratory‐rearing experiments. In the laboratory, individuals were grown in temperatures and photoperiod simulating those at the latitude of origin, and in common‐garden conditions. 3. The size of adults sampled in nature was negatively correlated with latitude. In all populations the females were larger than the males. Results from simulated and common‐garden rearing experiments supported this pattern of size difference across latitudes and between sexes, suggesting a genetic component for the latitudinal size trend and female‐biased size dimorphism. In contrast, hatchling size showed a positive relationship with latitude, but egg size, although differing between latitudes, showed no such relationship. 4. The results support a converse Bergmann cline, i.e. a negative body size cline towards the north. This negative cline in body size is probably driven by progressively stronger seasonal time and temperature constraints towards the higher latitudes and by the obligate univoltine life cycle of L. sponsa. As egg size showed no relationship with latitude, other environmental factors besides temperature, such as desiccation risk, probably affect this trait.     

Stable and variable life‐history responses to temperature and photoperiod in the beet webworm,Loxostege sticticalis

The beet webworm, Loxostege sticticalis L. (Lepidoptera: Crambidae), unlike many temperate insects and despite its wide distribution range, has a geographically stable value of the critical photoperiod for diapause induction. It has thus been hypothesized that the species could adjust its life cycle to different climates in an alternative way, which should be reflected in geographical variation and/or environmental plasticity of some other ecophysiological trait. Three remote populations of the beet webworm were studied. The insects were reared from egg to adult at several combinations of temperature and photoperiod in order to measure development times of all the immature stages and pre‐pupal body weight, and to characterize the sensitivity of these life‐history traits to the two ecological factors. The thermal reaction norms for immature development appeared to be significantly different in the three populations. There was also a significant effect of photoperiod on development time as well as on the thermal sensitivity and lower temperature threshold for larval development. Pre‐pupae from the northernmost population were heavier and their body mass was more strongly affected by photoperiod than in the other two, but attainment of a greater weight under short‐day conditions, especially combined with higher temperatures, was common for all the three populations. Nevertheless, all the discovered geographical and environmentally induced differences in life‐history traits were very small and their adaptive significance remains problematic.     

Latitudinal variation in sexual dimorphism in life‐history traits of a freshwater fish

Sexual dimorphism is common across the animal kingdom, but the contribution of environmental factors shaping differences between the sexes remains controversial. In ectotherms, life‐history traits are known to correlate with latitude, but sex‐specific responses are not well understood. We analyzed life‐history trait variation between the sexes of European perch (Perca fluviatilis L.), a common freshwater fish displaying larger female size, by employing a wide latitudinal gradient. We expected to find sex‐dependent latitudinal variation in life‐history variables: length at age, length increment, and size at maturity, with females showing consistently higher values than males at all latitudes. We further anticipated that this gender difference would progressively decrease with the increasingly harsh environmental conditions toward higher latitude. We hypothesized that growth and length increment would decrease and size/age at maturity would increase at higher latitudes. Our results confirmed female‐biased sexual size dimorphism at all latitudes and the magnitude of sexual dimorphism diminished with increase in latitude. Growth of both sexes decreased with increase in latitude, and the female latitudinal clines were steeper than those of males. Hence, we challenge two predominant ecological rules (Rensch's and Bergmann's rules) that describe common large‐scale patterns of body size variation. Our data demonstrate that these two rules are not universally applicable in ectotherms or female‐biased species. Our study highlights the importance of sex‐specific differences in life‐history traits along a latitudinal gradient, with evident implications for a wide range of studies from individual to ecosystems level.     

Size‐related life‐history traits in geometrid moths: a comparison of a temperate and a tropical community

1. Comparative studies on insect life histories are facilitated by the increasing availability of reliable phylogenies but are hampered by the scarcity of comparable data. Fortunately, morphological proxies of some life‐history traits can be measured on preserved specimens. 2. This study compared values of size‐related life‐history traits among a tropical (Ugandan) and a temperate (Estonian) assemblage of geometrid moths. 3. A comparative analysis based on an originally derived phylogeny revealed that tropical moths were, on average, larger than temperate ones. Tropical moths also had somewhat lower relative abdomen masses than temperate ones. This indicates that the tropical rather than the temperate moths tend to use an income (rather than capital) breeding strategy. Nevertheless, no difference was found in a related index of pro‐ovigeny. When body size was accounted for, tropical moths were found to lay smaller eggs than temperate ones. 4. The differences between the two compared areas are consistent with selection on higher mobility of the moths imposed by the more diverse tropical vegetation. Relatively larger eggs of temperate moths may constitute an adaptation to overcome the presumably stronger quantitative defences of their host plants. 5. Overall, however, we conclude that the differences in ecologically relevant size‐related traits are relatively low among moth assemblages of a tropical and a temperate forest region, indicating that these environments may not impose radically different selective pressures on insect life histories.     

Phenotypic divergence in reproductive traits of a moth population experiencing a phenological shift

Allochrony that is reproductive isolation by time may further lead to divergence of reproductive adaptive traits in response to different environmental pressures over time. A unique “summer” population of the pine processionary moth Thaumetopoea pityocampa, reproductively isolated from the typical winter populations by allochronic differentiation, is here analyzed. This allochronically shifted population reproduces in the spring and develops in the summer, whereas “winter” populations reproduce in the late summer and have winter larval development. Both summer and winter populations coexist in the same pine stands, yet they face different climatic pressures as their active stages are present in different seasons. The occurrence of significant differences between the reproductive traits of the summer population and the typical winter populations (either sympatric or allopatric) is thus hypothesized. Female fecundity, egg size, egg covering, and egg parasitism were analyzed showing that the egg load was lower and that egg size was higher in the summer population than in all the studied winter populations. The scales that cover the egg batches of T. pityocampa differed significantly between populations in shape and color, resulting in a looser and darker covering in the summer population. The single specialist egg parasitoid species of this moth was almost missing in the summer population, and the overall parasitism rates were lower than in the winter population. Results suggest the occurrence of phenotypic differentiation between the summer population and the typical T. pityocampa winter populations for the life‐history traits studied. This work provides an insight into how ecological divergence may follow the process of allochronic reproductive isolation.     

Direct and indirect genetic effects in life‐history traits of flour beetles (Tribolium castaneum)

Indirect genetic effects (IGEs) are the basis of social interactions among conspecifics, and can affect genetic variation of nonsocial and social traits. We used flour beetles (Tribolium castaneum) of two phenotypically distinguishable populations to estimate genetic (co)variances and the effect of IGEs on three life‐history traits: development time (DT), growth rate (GR), and pupal body mass (BM). We found that GR was strongly affected by social environment with IGEs accounting for 18% of the heritable variation. We also discovered a sex‐specific social effect: male ratio in a group significantly affected both GR and BM; that is, beetles grew larger and faster in male‐biased social environments. Such sex‐specific IGEs have not previously been demonstrated in a nonsocial insect. Our results show that beetles that achieve a higher BM do so via a slower GR in response to social environment. Existing models of evolution in age‐structured or stage‐structured populations do not account for IGEs of social cohorts. It is likely that such IGEs have played a key role in the evolution of developmental plasticity shown by Tenebrionid larvae in response to density. Our results document an important source of genetic variation for GR, often overlooked in life‐history theory.     

Population differences in early life‐history traits in grayling

In this paper we test population differences in early life‐history traits in three grayling Thymallus thymallus populations. The grayling shared ancestors some 80–90 years ago. We performed common‐garden experiments at three temperatures (mimicking population‐specific summer temperatures), and measured survival and growth rates during early development. We found significant additive genetic variance in size (length and yolk‐sac volume) measured at hatching, swim‐up and termination of the experiment, and significantly different reaction norms for growth rate and survival during the period of first feeding. In general, each population did best at the temperature experienced in nature. These differences in early life‐history traits suggest that natural selection has resulted in local adaptation in a time period of 13–18 generations.     

Variation of life‐history traits of the Asian corn borer,Ostrinia furnacalis in relation to temperature and geographical latitude

Life‐history traits from four geographical populations (tropical Ledong population [LD], subtropical Guangzhou [GZ] and Yongxiu populations, and temperate Langfang population [LF]) of the Asian corn borer, Ostrinia furnacalis were investigated at a wide range of temperatures (20–32°C). The larval and pupal times were significantly decreased with increasing rearing temperature, and growth rate was positively correlated with temperature. The relationship between body weight and rearing temperature in O. furnacalis did not follow the temperature–size rule (TSR); all populations exhibited the highest pupal and adult weights at high temperatures or intermediate temperatures. However, development time, growth rate, and body weight did not show a constant latitudinal gradient. Across all populations at each temperature, female were significantly bigger than males, showing a female‐biased sexual size dimorphism (SSD). Contrary to Rensch's rule, the SSD tended to increase with rising temperature. The subtropical GZ population exhibited the largest degree of dimorphism while the temperate LF exhibited the smallest. Male pupae lose significantly more weight at metamorphosis compared to females. The proportionate weight losses of different populations were significantly different. Adult longevity was significantly decreased with increasing temperature. Between sexes, all populations exhibit a rather female‐biased adult longevity. Finally, we discuss the adaptive significance of higher temperature‐inducing high body weight in the moth's life history and why the moth exhibits the reverse TSR.     

rangeMapper: a platform for the study of macroecology of life‐history traits

Aim To introduce rangeMapper, an R package for the study of the macroecological patterns of life‐history traits, and demonstrate its capabilities using three case studies. The first case study addresses an important topic in conservation biology: biodiversity hotspots. Specifically, we investigate the congruence between global hotspots of three parameters that describe avian diversity: species richness, endemic species richness and relative body mass diversity. The second case study investigates a topic of relevance for macroecology: the inter‐specific relationship between range size and body size for avian assemblages, and how it varies geographically. The third case study tackles a methodological problem in macroecology: the influence of map resolution on statistical inference, i.e. the question of whether and how the relationship between species richness and body mass varies with map resolution. Innovation rangeMapper offers a tight integration of spatial and statistical tools for macroecological projects and it relies on a high‐performance database engine which makes it suitable for managing projects using a large number of species. rangeMapper's architecture follows closely the concepts described by Gaston et al. (2008 Journal of Biogeography, 35 , 483–500) and its flexibility allows for both complex data manipulation procedures and easy implementation of new functions. By choosing case studies to cover various technical and conceptual issues we demonstrate rangeMapper's capabilities to address a wide array of questions. Main conclusion rangeMapper ( http://cran.r‐project.org/package=rangeMapper ) is an open source front end software which can be used to address questions in both fundamental ecological research and conservation biology.     

Maternal and environmental influences on egg size and juvenile life‐history traits in Pacific salmon

Life‐history traits such as fecundity and offspring size are shaped by investment trade‐offs faced by mothers and mediated by environmental conditions. We use a 21‐year time series for three populations of wild sockeye salmon (Oncorhynchus nerka) to test predictions for such trade‐offs and responses to conditions faced by females during migration, and offspring during incubation. In years when their 1100 km upstream migration was challenged by high water discharges, females that reached spawning streams had invested less in gonads by producing smaller but not fewer eggs. These smaller eggs produced lighter juveniles, and this effect was further amplified in years when the incubation water was warm. This latter result suggests that there should be selection for larger eggs to compensate in populations that consistently experience warm incubation temperatures. A comparison among 16 populations, with matching migration and rearing environments but different incubation environments (i.e., separate spawning streams), confirmed this prediction; smaller females produced larger eggs for their size in warmer creeks. Taken together, these results reveal how maternal phenotype and environmental conditions can shape patterns of reproductive investment and consequently juvenile fitness‐related traits within and among populations.     

Altitudinal variation in life‐history traits in the lawn ground cricket,Polionemobius mikado

It has been reported that the lawn ground cricket, Polionemobius mikado (Orthoptera: Trigonidiidae), shows variations in voltinism and life‐history traits along latitude in Japan, but whether it shows variations along altitude has not been examined. The present study aims to determine whether there is variation in voltinism, body size and the incidence of embryonic diapause among populations of P. mikado at different altitudes in Kyoto, Japan. Six populations collected from 70 to 800 m a.s.l. showed a positive relationship between the adult head width and the altitude of sampling locality. This body size cline is attributed to the increasing proportion of larger univoltine individuals from low to high altitude. The incidence of diapause of the strain from altitude 800 m was approximately 100% when reared under natural photoperiods and temperatures from spring to summer at an altitude of 70 m, and was significantly higher than that of the strain from 70 m. These differences in life‐history traits suggest that the population of P. mikado at a higher altitude has adapted to a shorter length of the season available for growth.     

An invasive species imposes selection on life‐history traits of a native frog

As well as their direct ecological impacts on native taxa, invasive species can impose selection on phenotypic attributes (morphology, physiology, behaviour, etc.) of the native fauna. In anurans, body size at metamorphosis is a critical life‐history trait: for most challenges faced by post‐metamorphic anurans, larger size at metamorphosis probably enhances survival. However, our studies on Australian frogs (Limnodynastes convexiusculus) show that this pattern can be reversed by the arrival of an invasive species. When metamorph frogs first encounter invasive cane toads (Bufo marinus), they try to eat the toxic invader and, if they are able to do so, are likely to die from poisoning. Because frogs are gape‐limited predators, small metamorphs cannot ingest a toad and thus survive long enough to disperse away from the natal pond (and thus from potentially deadly toads). These data show that larger size at metamorphosis can reduce rather than increase anuran survival rates, because larger metamorphs are more easily able to ingest (and thus be poisoned by) metamorph cane toads. Our results suggest that patterns of selection on life‐history traits of native taxa (such as size and age at metamorphosis, seasonal timing of breeding and duration of pondside aggregation prior to dispersal) can be modified by the arrival of an invasive species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 329–336.     

Susceptibility of the pine processionary caterpillar Thaumetopoea pityocampa (Lepidoptera: Thaumetopoeidae) toward δ-endotoxins of Bacillus thuringiensis under laboratory conditions

A series of natural crystal proteins from B. thuringiensis subsp. Alesti 12–25, caucasicus, galleriae 11–67, galleriae 6–96, kenyae, and shondungensis and spore‐crystal preparations from finitimus 11–66 and from a recombinant strain of B. thuringiensis subsp. kurstaki expressing Cry 1 Ga1 only, were assessed as a toxic agent for the pine processionary caterpillar, Thaumetopoea pityocampa. Some preparations had a thoroughly investigated composition and contained Cry1Aa, Cry1Ab2, Cry1Ab7, Cry1D, Cry1F, Cry 1 Ga1, Cry9Aa, Cry26 crystal proteins, whereas crystals of B. thuringiensis subsp. caucasicus, kenyae, and shondungensis harboured predominantly unidentified toxins distant from commonly used prototypes. Bioassays were based on the simultaneous assignment of each treatment to groups of 20 full sibling first‐instar larvae, obtained from broods of a population from North‐western Italy. The toxin was applied to pine needles by the leaf dipping method and the effect was registered in both feeding inhibition and mortality. B. thuringiensis subsp. caucasicus, kenyae, galleriae 6–96, alesti, and galleriae 11–67 gave the best results in terms of both feeding inhibition and larval mortality. Broods tested in B. thuringiensis bioassays showed a substantial variation in susceptibility to the toxins, suggesting the potential development of resistance in the population.     

Geographical variability in life‐history traits of a midslope dogfish: the brier shark Deania calcea

Deania calcea (n = 420) were collected from the catch of deep‐water trawlers in the southern and eastern scalefish and shark fishery in southern Australia during the years 2008–2011. The total length (L_T) range varied between sexes, females being larger (n = 264; 280–1530 mm) than males (n = 156; 310–921 mm). The reproductive cycle in this population is non‐continuous and asynchronous. The estimated L_T at which 50% of males are mature is 807 mm and is 914 mm for females. Populations of D. calcea in higher latitudes appear to mature at a larger size than conspecifics in lower latitudes, in both the northern and southern hemispheres. Litters ranged from three to 10 embryos with a 1:1 sex ratio, but litter size does not increase with maternal L_T. Deania calcea shows geographical variability in its biological parameters and gathering information on life‐history traits of populations is vital to understand the trade‐offs made by this species in response to environmental conditions and to predict intraspecific spatial differences. Such information is a basis for specific spatial management to protect populations from excessive fishing.     

Hypoxia and life‐history traits in a eurytopic African cichlid

This study quantified variation in key life‐history traits of the widespread African mouth‐brooding cichlid Pseudocrenilabrus multicolor victoriae. Egg size, number, batch reproductive effort, size at maturity and brooding efficiency were compared among field populations across a wide range of dissolved oxygen (DO) concentrations from extreme hypoxia to normoxia. In the laboratory, a similar suite of characters was quantified in F1 of low‐ and high‐DO origin reared under low or high DO. In general, females from low‐DO habitats and females reared under low DO were characterized by a smaller size at maturity and no difference in batch reproductive effort when compared with females from high‐DO habitats or females reared under normoxia. A trade‐off between egg size and number was evident in the field and in the laboratory‐rearing experiment, but the direction of the trade‐off differed. Egg size was negatively correlated with egg number across field populations; females collected from low‐DO sites generally had more, smaller eggs relative to females from high‐DO sites. In the laboratory‐rearing experiment, F1 females of high‐DO origin produced larger, fewer eggs than F1 females of low‐DO origin, lending support to the field results and suggesting a heritable component to these traits. There was also an element of developmental plasticity, F1 females raised under low DO produced larger, fewer eggs compared with F1 females raised under high DO (regardless of population) suggesting that DO may interact with other variables to determine egg size in the field.     

Evolution of larval competitiveness and associated life‐history traits in response to host shifts in a seed beetle

Resource competition is frequently strong among parasites that feed within small discrete resource patches, such as seeds or fruits. The properties of a host can influence the behavioural, morphological and life‐history traits of associated parasites, including traits that mediate competition within the host. For seed parasites, host size may be an especially important determinant of competitive ability. Using the seed beetle, Callosobruchus maculatus, we performed replicated, reciprocal host shifts to examine the role of seed size in determining larval competitiveness and associated traits. Populations ancestrally associated with either a small host (mung bean) or a large one (cowpea) were switched to each other's host for 36 generations. Compared to control lines (those remaining on the ancestral host), lines switched from the small host to the large host evolved greater tolerance of co‐occurring larvae within seeds (indicated by an increase in the frequency of small seeds yielding two adults), smaller egg size and higher fecundity. Each change occurred in the direction predicted by the traits of populations already adapted to cowpea. However, we did not observe the expected decline in adult mass following the shift to the larger host. Moreover, lines switched from the large host (cowpea) to the small host (mung bean) did not evolve the predicted increase in larval competitiveness or egg size, but did exhibit the predicted increase in body mass. Our results thus provide mixed support for the hypothesis that host size determines the evolution of competition‐related traits of seed beetles. Evolutionary responses to the two host shifts were consistent among replicate lines, but the evolution of larval competition was asymmetric, with larval competitiveness evolving as predicted in one direction of host shift, but not the reverse. Nevertheless, our results indicate that switching hosts is sufficient to produce repeatable and rapid changes in the competition strategy and fitness‐related traits of insect populations.     

Size‐induced phenotypic reaction norms in a parasitoid wasp: an examination of life‐history and behavioural traits

The amount of resources available during development often affects body size, causing phenotypic variation in life‐history traits and reproductive behaviours. However, past studies have seldom examined the reaction norms of both life‐history and behavioural traits versus body size. We measured the phenotypic plasticity of several life‐history (age‐specific egg load, egg size, longevity) and behavioural (oviposition rate, host marking rate, walking speed) traits of the egg parasitoid Telenomus podisi Ashmead (Hymenoptera: Scelionidae) in response to body size variation. We predicted that life‐history traits would show more evidence of size compensation than behavioural traits, resulting in fewer positively‐sloped size versus trait reaction norms among the former. As predicted by life‐history models, smaller wasps appear to shift resource allocation towards early‐life reproduction, having a similar egg load to large individuals 9 days after emergence. Surprisingly, longevity was unaffected by body size. However, egg size, the number of offspring produced during oviposition bouts, and the rate of subsequent egg synthesis were greater for larger individuals. In addition, as predicted, the reaction norms of behavioural traits versus body size were all positively sloped. Thus, despite possible adaptive compensatory plasticity of life‐history traits by small individuals, behavioural constraints directly related to body size would contribute to maintaining a positive size–fitness relationship.