Variation in ovariole number and egg size of species of Dacus (Diptera; Tephritidae) and their relation to host specialization

Abstract. 1. An analysis of the number of ovarioles/ovary is presented for fourteen species of Dacus ranging from broad generalists (D.tryoni (Frogg.)) to strict host specialists (D.aglaia Hardy). For eight species data on egg size and the incubation time of eggs is also presented. Interrelationships among these life history variables and their relationships to host specialization were examined. 2. There was a broad positive relationship between the breadth of the natural host range (i.e. excluding cultivated fruits) and ovariole number (and hence potential fecundity). Highly polyphagous species had thirty-five to forty ovarioles/ovary while the specialists ranged from eight to twenty. 3. D.musae (Tryon), a specialist on banana, was an exception having an ovariole number similar to that of the most polyphagous species. 4. There was no clear relationship between egg size and ovariole number across eight species, though monophages were not fully represented in this analysis. Among these eight species there was no significant relationship between wing length and egg length, nor was there any such relationship among individuals within species. 5. There was considerable variation among species in the incubation time (at 25°C) for eggs, ranging from 24 h for D.cucumis French to 53 h for D.cacuminatus (Hering). This variation was only weakly related to egg size, though D.cucumis produced the largest eggs of any species examined. 6. Possible explanations for a relationship between potential fecundity and host range are discussed in terms of the balance between the abundance of host resources for generalists and their predictability for specialists. However, the observed relationship between ovariole number and host range must remain tentative, given that only fourteen species from one genus have been examined. Considerably more data is required for other dacines, for species in other tephritid subfamilies, and for other phytophagous insect groups to determine whether the relationship is generally applicable.     

Effects of eucalypt nutritional quality on the Bog gum‐Victorian metapopulation of Ctenarytaina bipartita and implications for host and range expansion

1. Novel hosts present phytophagous insects with nutritional challenges which can cause host‐associated divergence. 2. The performance of the Bog gum‐Victorian metapopulation of Ctenarytaina bipartita Burckhardt et al. (Psylloidea:Aphalaridae), a psyllid whose range is being expanded by tree planting, on five species and genotypes of Eucalyptus endemic to south‐eastern Australia was quantified. Settling of females on two non‐hosts was also tested. 3. Female C. bipartita exhibited significant host‐associated plasticity in proctiger length (the body part used for oviposition into apical buds). Psyllids with longer wings and proctigers arose from a primary host. Fecundity varied significantly among hosts and was highest on a novel host. Hosts did not differ significantly in free amino acids (FAAs) but did differ in concentrations of essential amino acids (EAAs). However, nymphs did not differ significantly in EAAs. Surprisingly, fecundity was not related to total FAAs, availability of EAAs or concentrations of EAAs but was related to concentrations of four non‐EAAs. Mean fecundity per host was also positively related to the relative abundance of galloyl groups (associated with hydrolysable tannins). 4. Leaf age was more important to settling than eucalypt species; females settled on young leaves but this response was not related to leaf water content. 5. Planting the rare Eucalyptus kitsoniana in new habitats will expand the range of Bog gum‐Victorian C. bipartita and provide a bridge for the colonisation of novel eucalypts with buds suitable for oviposition. Host expansion by this metapopulation is not constrained by nutritional quality and may result in morphological divergence.     

Selection on body size and sexual size dimorphism differs between host species in a seed-feeding beetle

Sexual size dimorphism varies substantially among populations and species but we have little understanding of the sources of selection generating this variation. We used path analysis to study how oviposition host affects selection on body size in a seed-feeding beetle (Stator limbatus) in which males contribute large ejaculates (nuptial gifts) to females. Females use nutrients in these ejaculates for egg production. Male body size, which affects ejaculate size, affects female fecundity and is thus under fecundity selection similar in magnitude to the fecundity selection on female body size. We show that when eggs are laid on a host on which larval mortality is low (seeds of Acacia greggii) fecundity predicts fitness very well and fecundity selection is the major source of selection on both male and female adult size. In contrast, when eggs are laid on a host on which larval mortality is high (seeds of Parkinsonia florida) fecundity poorly predicts fitness such that fecundity selection is relaxed on both male and female size. However, because egg size affects larval mortality on this poor host (P. florida) there is selection on female size via the female size --> egg size --> fitness path; this selection via egg size offsets the reduction in fecundity selection on female, but not male, body size. Thus, differences in host suitability (due to differences in larval mortality) affect the relative importance of two sources of selection on adult body size; fecundity selection on both male and female body size is lower on the poor quality host (P. florida) relative to the high quality host (A. greggii) whereas selection on female body size via effects of egg size on offspring survival (body size --> egg size --> fitness) is greater on the poor quality host relative to the high quality host. Because selection via the egg size path affects only females the difference in larval survival between hosts shifts the relative magnitude of selection on female vs. male size. Researchers working on other study systems should be alerted to the possible importance of subtle, but consequential, indirect selection on their study organisms.     

Egg production across a 40-week period in the phasmid Sipyloidea sp. (Diapheromeridae) from a tropical rain forest, north Queensland, Australia

Abstract  In this study we report the results from the first long-term (40 weeks) study of stick-insect fecundity and distribution under natural conditions of which we are aware. We used the number of eggs falling into 72 × 0.5 m² traps to ask: 'Was egg production in Sipyloidea sp. uniform across the sample period'? and 'Was there evidence of host plant species preference or avoidance'? We collected a total of 213 Sipyloidea sp. eggs. The number of eggs caught per week was not uniform and an exponential decay model was the best-fit relationship between egg production and time, indicative of a steep decline from high to low (but continuous) egg production across the study period. Continuous egg production differs from other insect species in tropical areas that often show distinct seasonal differences between wet and dry seasons, timed to leaf production in host plants. The distribution of eggs within traps was aggregated, and more traps than expected from Poisson probabilities received no eggs, or six or more eggs. The concentration of eggs within particular traps was not related to the identity of canopy plant species, however. We suggest that continuous egg production in Sipyloidea sp. may be related to the wider range of plant species available as food resources for the polyphagous Sipyloidea , compared with other tropical insect species.     

HOST EFFECTS ON FERTILITY AND REPRODUCTIVE SUCCESS OF DENDROCTONUS PONDEROSAE HOPKINS (COLEOPTERA: SCOLYTIDAE)

Mountain pine beetles from lodgepole and limber pine in western Canada were crossbred. We compared data about reproductive success and fecundity of parents as well as development, mortality, and fertility of their progeny to determine whether there was reproductive isolation among beetle populations in these hosts. Three factors, directly or indirectly related to the host, influenced reproductive performance of parents (reproductive success, egg gallery length, fecundity, and number of eggs laid per centimeter of gallery) as well as the mortality, dry weight, and fat content of the progeny: (1) the host species in which progeny were reared, (2) the host species in which the female parent was reared, and (3) whether both parents originated from the same or different host species. Limber pine appears to be a better host for Dendroctonus ponderosae reproduction and survival than lodgepole pine. Nonetheless, beetles reared from lodgepole and limber pine can reproduce in either host and will mate with each other. Progeny of all crosses were fertile. Thus, there is no apparent barrier to prevent beetles from the two host species from interbreeding in the field.     

Intraspecific variability in egg maturation patterns and associated life‐history trade‐offs in a polyembryonic parasitoid wasp

1. Life‐history theory predicts a trade‐off between the resources allocated to reproduction and those allocated to survival. Early maturation of eggs (pro‐ovigeny) is correlated with small body size and low adult longevity in interspecific comparisons among parasitoids, demonstrating this trade‐off. The handful of studies that have tested for similar correlations within species produced conflicting results. 2. Egg maturation patterns and related life‐history traits were studied in the polyembryonic parasitoid wasp, Copidosoma koehleri (Hymenoptera: Encyrtidae). Although the genus Copidosoma was previously reported to be fully pro‐ovigenic, mean egg loads of host‐deprived females almost doubled within their first 6 days of adulthood. 3. The initial egg‐loads of newly emerged females were determined and age‐specific realised fecundity curves were constructed for their clone‐mate twins. The females' initial egg loads increased with body size, but neither body size nor initial egg load was correlated with longevity and fecundity. 4. The variation in initial egg loads was lowest among clone‐mates, intermediate among non‐clone sisters and highest among non‐sister females. The within‐clone variability indicates environmental influences on egg maturation, while the between‐clone variation may be genetically based. 5. Ovaries of host‐deprived females contained fewer eggs at death (at ～29 days) than on day 6. Their egg loads at death were negatively correlated with life span, consistent with reduced egg production and/or egg resorption. Host deprivation prolonged the wasps' life span, suggesting a survival cost to egg maturation and oviposition. 6. It is concluded that adult fecundity and longevity were not traded off with pre‐adult egg maturation.     

The cost of being able to fly: a study of wing polymorphism in two species of crickets

Summary The widespread occurrence of wing polymorphisms in insects suggests that the possession of wings and ability to fly adversely affect components of the insect's life characteristics that contribute to its Darwinian fitness. This hypothesis was tested by an analysis of the differences in life history parameters of the macropterous and micropterous morphs of the two cricket species G. firmus and A. fasciatus. In both species there were no differences in development time or adult survival between the two morphs. Significant differences in head width were not consistent between the two species but in both sexes of G. firmus and females of A. fasciatus (insufficient males for analysis) long-winged individuals weighed more than short-winged individuals with the same head width. In both species egg production is delayed in macropterous females. The cumulative fecundity of the micropterous morph is greater than the macropterous morph in both species but only in G. firmus is the difference statistically significant. A. fasciatus frequently loose their wings but no such loss has been observed in G. firmus. There is a significant increase in egg production after the loss of the wings. These results are in accord with those of Tanaka (1976) for the cricket, Pteronemobius taprobanensis.Breeding experiments indicate that in G. firmus the wing polymorphism is under genetic control. The decrease in fecundity is sufficiently large that genotypes producing only macropterous offspring could only persist in highly unstable environments where continuous dispersal was imperative for survival. However, the reproductive cost of a genotype producing a small percentage of macropterous individuals is slight. The fitness that accrues to a genotype producing a few dispersing offspring is likely to offset the small reproductive cost and hence wing polymorphisms should be favoured.     

The relationship of body size to breadth of diet in some Lepidoptera

Abstract. 1. We have explored the question of whether breadth of diet in moths is associated with adaptations other than host selection behaviour, with particular reference to body size.
2. Generalist species in Britain and New York are larger than specialist species.
3. This relationship is probably not due to latitudinal clines, nor to higher fecundity in generalist species.
4. We suggest that the large bodies of generalists buffer them against environmentally-induced physiological stress.     

Crown defoliation decreases reproduction and wood growth in a marginal European beech population

Background and AimsAbiotic and biotic stresses related to climate change have been associated with increased crown defoliation, decreased growth and a higher risk of mortality in many forest tree species, but the impact of stresses on tree reproduction and forest regeneration remains understudied. At the dry, warm margin of species distributions, flowering, pollination and seed maturation are expected to be affected by drought, late frost and other stresses, eventually resulting in reproduction failure. Moreover, inter-individual variation in reproductive performance versus other performance traits (growth, survival) could have important consequences for population dynamics. This study investigated the relationships among individual crown defoliation, growth and reproduction in a drought-prone population of European beech, Fagus sylvatica.MethodsWe used a spatially explicit mating model and marker-based parentage analyses to estimate effective female and male fecundities of 432 reproductive trees, which were also monitored for basal area increment and crown defoliation over 9 years.Key ResultsFemale and male fecundities varied markedly between individuals, more than did growth. Both female fecundity and growth decreased with increasing crown defoliation and competition, and increased with size. Moreover, the negative effect of defoliation on female fecundity was size-dependent, with a slower decline in female fecundity with increasing defoliation for the large individuals. Finally, a trade-off between growth and female fecundity was observed in response to defoliation: some large trees maintained significant female fecundity at the expense of reduced growth in response to defoliation, while some other defoliated trees maintained high growth at the expense of reduced female fecundity.ConclusionsOur results suggest that, while decreasing their growth, some large defoliated trees still contribute to reproduction through seed production and pollination. This non-coordinated decline of growth and fecundity at individual level in response to stress may compromise the evolution of stress-resistance traits at population level, and increase forest tree vulnerability.     

Does host determine short‐range flight capacity of trichogrammatids?

Biological traits, such as body size, fecundity and fertility, of egg parasitoids are strongly influenced by host species, and the use of a single‐host species for consecutive generations has been showed to be detrimental for egg parasitoid biology. Besides biological traits, behaviour of egg parasitoids is also an important parameter for implying their performance in the field, but information about the effect of host on the behaviour of egg parasitoids is still scarce. Thus, this work aimed at determining the influence of host species on the short‐range flight capacity of three egg parasitoids Trichogrammatoidea annulata De Santis, Trichogramma atopovirilia Oatman and Platner and Trichogramma bruni Nagajara reared on Anagasta kuehniella (Zeller), Corcyra cephalonica (Stainton) and Sitotroga cerealella (Olivier), for first, 10 and 28 consecutive generations. Trichogrammatids emerged inside a tubular dark flight chamber, under controlled conditions, and short‐range flying response was based on the proportions of adults trapped to the sticky top of the chamber and the ones found on the bottom, which were checked for defective wings. Our data clearly demonstrate that trichogrammatid short‐range flight depends on the fasctidious host species. All three egg parasitoids had their flight capacity significantly reduced when emerged from S. cerealella. The highest flying capacity of T. annulata was registered when reared on C. cephalonica, while for T. atopovirilia was on A. kuehniella and T. bruni on both C. cephalonica and A. kuehniella. Parasitoid flight capacity is affected when reared on the same host over generations; however, it was not possible to define a pattern for any of the trichogrammatids. Our findings are of great relevance for quality control of mass‐reared egg parasitoids because measuring only behavioural traits with use of flight chambers can be more practical and less time‐consuming than assessing biological parameters.     

RELATIONSHIPS BETWEEN GENETIC VARIABILITY AND LIFE-HISTORY FEATURES OF BONY FISHES

Correlations between genetic variation and life-history variables were obtained for 80 species of bony fishes as a means of testing the hypothesis that genetic variation is directly related to 1) opportunity for balancing selection, as indicated by fecundity, and 2) environmental variation, as indicated by capacity for population increase. Genetic data were taken from the literature, and data on longevity, age at maturity, egg size, body size, and lifetime fecundity were taken from the literature where available and were otherwise estimated from other variables. Average heterozygosity does not increase significantly with increasing fecundity. However, heterozygosity is significantly associated with short generation times, quick maturation, small maximum size, and small eggs. Thus, heterozygosity appears to increase on a demographic continuum toward maximum values in species that are most strongly selected for maximizing the intrinsic rate of increase. Such species are associated with less stable environments. Thus, the results indicate a predominate role for environmental variation in controlling genetic variation of bony fishes.     

Reproductive cost of flight capability: a comparison of life history traits in wing dimorphic planthoppers

Abstract. 1. Reproductive costs associated with flight capability were evaluated in the wing dimorphic planthopper, Prokelisia dolus Wilson, by comparing the life history of traits of winged (macropterous) and flightless (brachypterous) females under controlled laboratory conditions.
2. Macropters with large thoraces and fully developed wings maintain a greater investment in flight apparatus than brachypters with small thoraces and reduced wings.
3. Associated with greater flight capability in the macropter of P.dolus are shorter adult life, decreased total fecundity, and delayed age at first reproduction compared to brachypterous females.
4. Under field conditions where mortality is high, the difference in realized fecundity between the two wing forms living on similar resources is further exaggerated with the brachypter having the greater advantage.
5. When the life history traits of the wing forms of P. dolus are compared with traits for nine other species of planthoppers, two similarities emerge. First, the preoviposition period of the macropterous wing form is always longer than that for the brachypter resulting in a reproductive delay. Second, most studies show that macropters are less fecund than brachypters.
6. There is no general tendency among planthopper species for macropterous adults to live fewer days or develop more slowly as nymphs compared to their flightless counterparts.
7. The reproductive delay and reduced fecundity of the volent wing form of planthoppers supports the notion that flight capability is costly and that phenotypic trade-offs between flight and reproduction exist.     

Comparative phylogenetic analysis of the evolution of semelparity and life history in salmonid fishes

The selective pressures involved in the evolution of semelparity and its associated life-history traits are largely unknown. We used species-level analyses, independent contrasts, and reconstruction of ancestral states to study the evolution of body length, fecundity, egg weight, gonadosomatic index, and parity (semelparity vs. degree of iteroparity) in females of 12 species of salmonid fishes. According to both species-level analysis and independent contrasts analysis, body length was positively correlated with fecundity, egg weight, and gonadosomatic index, and semelparous species exhibited a significantly steeper slope for the regression of egg weight on body length than did iteroparous species. Percent repeat breeding (degree of iteroparity) was negatively correlated with gonadosomatic index using independent contrasts analysis. Semelparous species had significantly larger eggs by species-level analysis, and the egg weight contrast for the branch on which semelparity was inferred to have originated was significantly larger than the other egg weight contrasts, corresponding to a remarkable increase in egg weight. Reconstruction of ancestral states showed that egg weight and body length apparently increased with the origin of semelparity, but fecundity and gonadosomatic index remained more or less constant or decreased. Thus, the strong evolutionary linkages between body size, fecundity, and gonadosomatic index were broken during the transition from iteroparity to semelparity. These findings suggest that long-distance migrations, which increase adult mortality between breeding episodes, may have been necessary for the origin of semelparity in Pacific salmon, but that increased egg weight, leading to increased juvenile survivorship, was crucial in driving the transition. Our analyses support the life-history hypotheses that a lower degree of repeat breeding is linked to higher reproductive investment per breeding episode, and that semelparity evolves under a combination of relatively high juvenile survivorship and relatively low adult survivorship.     

The role of fecundity and reproductive effort in defining life‐history strategies of North American freshwater mussels

Selection is expected to optimize reproductive investment resulting in characteristic trade‐offs among traits such as brood size, offspring size, somatic maintenance, and lifespan; relative patterns of energy allocation to these functions are important in defining life‐history strategies. Freshwater mussels are a diverse and imperiled component of aquatic ecosystems, but little is known about their life‐history strategies, particularly patterns of fecundity and reproductive effort. Because mussels have an unusual life cycle in which larvae (glochidia) are obligate parasites on fishes, differences in host relationships are expected to influence patterns of reproductive output among species. I investigated fecundity and reproductive effort (RE) and their relationships to other life‐history traits for a taxonomically broad cross section of North American mussel diversity. Annual fecundity of North American mussel species spans nearly four orders of magnitude, ranging from < 2000 to 10 million, but most species have considerably lower fecundity than previous generalizations, which portrayed the group as having uniformly high fecundity (e.g. > 200000). Estimates of RE also were highly variable, ranging among species from 0.06 to 25.4%. Median fecundity and RE differed among phylogenetic groups, but patterns for these two traits differed in several ways. For example, the tribe Anodontini had relatively low median fecundity but had the highest RE of any group. Within and among species, body size was a strong predictor of fecundity and explained a high percentage of variation in fecundity among species. Fecundity showed little relationship to other life‐history traits including glochidial size, lifespan, brooding strategies, or host strategies. The only apparent trade‐off evident among these traits was the extraordinarily high fecundity of Leptodea, Margaritifera, and Truncilla, which may come at a cost of greatly reduced glochidial size; there was no relationship between fecundity and glochidial size for the remaining 61 species in the dataset. In contrast to fecundity, RE showed evidence of a strong trade‐off with lifespan, which was negatively related to RE. The raw number of glochidia produced may be determined primarily by physical and energetic constraints rather than selection for optimal output based on differences in host strategies or other traits. By integrating traits such as body size, glochidial size, and fecundity, RE appears more useful in defining mussel life‐history strategies. Combined with trade‐offs between other traits such as growth, lifespan, and age at maturity, differences in RE among species depict a broad continuum of divergent strategies ranging from strongly r‐selected species (e.g. tribe Anodontini and some Lampsilini) to K‐selected species (e.g. tribes Pleurobemini and Quadrulini; family Margaritiferidae). Future studies of reproductive effort in an environmental and life‐history context will be useful for understanding the explosive radiation of this group of animals in North America and will aid in the development of effective conservation strategies.     

Rarity in Australian desert lizards

Most species of Australian desert lizards are uncommon. Possible causes of rarity are examined, including body size as measured by snout–vent length (SVL), fecundity, number of sites occupied, habitat niche breadth, microhabitat niche breadth, dietary niche breadth, and average total niche overlap with other species. Rare species tend to be larger with lower fecundities than abundant species and they occur at fewer sites. Many, but not all, uncommon species are specialists, either in habitat, microhabitat, or diet. The niche breadth hypothesis, which states that abundant species should be generalists whereas specialized species should be rare, is tested, but rejected as a general explanation for rarity. Some uncommon species exhibit high overlap with other species suggesting that they may experience diffuse competition. However, no single cause of rarity can be identified, but rather each species has its own idiosyncratic reasons for being uncommon. Multivariate analyses show distinct ecological differences between abundant and uncommon species.     

Phenotypic plasticity and fitness in egg parasitoids

Several proxies can be used to estimate the fitness of egg parasitoids: size, longevity, fecundity, mating ability and vagility. All these proxies are positively correlated with the lifetime fitness gain of an individual. However, the phenotypic plasticity, which is the ability of a genotype to produce distinct phenotypes depending on environmental conditions, may alter the value of these proxies. Host related factors (host species and quality), competition and duration of development will influence the proxies expressed by a given phenotype. When the phenotype is modified by the temperature at which an individual develops, the resulting phenotype will vary based on the life history strategy of the parasitoid. While in koinobiont species an increase in size is positively correlated with longevity and fecundity, the reverse occurs with idiobiont parasitoids. Using size as a proxy could thus be misleading depending on the factors that influence the size of the adult.     

Effects of diet and host access on fecundity and lifespan in two fruit fly species with different life‐history patterns

The reproductive ability of female tephritids can be limited and prevented by denying access to host plants and restricting the dietary precursors of vitellogenesis. The mechanisms underlying the delayed egg production in each case are initiated by different physiological processes that are anticipated to have dissimilar effects on lifespan and reproductive ability later in life. The egg‐laying abilities of laboratory‐reared females of the Mediterranean fruit fly (Ceratitis capitata Wiedmann) and melon fly (Bactrocera cucurbitae Coquillett) from Hawaii are delayed or suppressed by limiting access to host fruits and dietary protein. In each case, this is expected to prevent the loss of lifespan associated with reproduction until protein or hosts are introduced. Two trends are observed in each species: first, access to protein at eclosion leads to a greater probability of survival and a higher reproductive ability than if it is delayed and, second, delayed host access reduces lifetime reproductive ability without improving life expectancy. When host access and protein availability are delayed, the rate of reproductive senescence is reduced in the medfly, whereas the rate of reproductive senescence is generally increased in the melon fly. Overall, delaying reproduction lowers the fitness of females by constraining their fecundity for the remainder of the lifespan without extending the lifespan. © 2013 The Royal Entomological Society     

Reproductive strategies of sub-Antarctic lithodid crabs vary with habitat depth

We present the first fecundity and egg size data for three species of lithodid crab from Antarctic waters south of the Polar Front, caught in the fisheries operating around the island of South Georgia. In all species, fecundity was observed to increase with body size, but reproductive allocation was found to differ significantly between species. The highest relative fecundity (eggs^−g body weight) and smallest egg size was found in Paralomis spinosissima, which is found in shallower waters, whereas the lowest relative fecundity values and largest eggs were recorded in the deeper living species, Neolithodes diomedeae. Evidence is presented that closely related sympatric species may employ quite different reproductive strategies that suit their depth distribution whilst indicating the overriding evolutionary adaptation of reproductive traits to temperature and food availability.     

Latitudinal gradients and the shaping of life‐history traits in a gregarious caterpillar

The present study aimed to investigate how the impact of several factors linked to geography would shape life‐history traits in a gregarious species, using the pine processionary moth (PPM) Thaumetopoea pityocampa as a model system. PPM has a wide geographical distribution over the Mediterranean Basin, and it is a strictly gregarious species throughout larval development, where the total reproductive output of each female forms a colony. We reviewed both published and unpublished data on PPM from all over its distribution in the Mediterranean Basin and extracted data on fecundity, egg size, egg parasitoid mortality, flight period, and development time. These life‐history traits were then related to location, expressed as latitude and altitude, local average temperatures, and host tree species. We found that PPM fecundity increaseed with latitude, concomitant with an increase in the length of development and an earlier onset of adult flight. These results are the opposite of that found in other Lepidoptera species with a wide geographical distribution, as well as in insects in general. We propose that a large colony size in PPM is important at higher latitudes because this confers an advantage for thermoregulation and tent building in areas where larvae have to face harsher conditions during the winter, thus shifting the optimal trade‐off between the number and size of eggs with latitude. However, host tree species also affected the relationship between egg number and size and the optimal outcome of these traits is likely a compromise between different selection pressures. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 224–236.