Immature Stages of Tomapoderus(T.) ruficollis Fabricius (Coleoptera: Attelabidae) from Korea

Eggs, larvae and pupae of Tomapoderus (T.) ruficollis Fabricius are described and illustrated. The species is found on host plants, Zelkova serrata Makino and Ulmus davidiana var. japonica Nakai. and is a well‐known forest pest. Taxonomic notes and cradle structure of this species are provided.     

Effects of leaf quality and microhabitat on the survival of a leaf-rolling weevil (Attelabidae)

Female Cycnotrachelus roelofsi (Coleoptera: Attelabidae) construct two types of leaf roll, i.e., cut-off cradles (CCs) and suspended cradles (SCs), in which to lay eggs; these cradles are generally constructed using older and younger leaves, respectively. We conducted two experiments to determine whether the quality of cradle leaves affects egg and larval survival. In the first experiment, we severed all SCs from a tree and placed them on the ground with unmanipulated CCs in the early breeding season. In the second experiment, we resuspended all CCs in a tree with unmanipulated SCs in the late breeding season. We also compared leaf mass per area (LMA), polyphenol content, and nitrogen content between the two cradle types to determine whether there were any differences in leaf quality. Larval mortality, probably caused by cradle herbivory, was significantly greater in severed SCs than in intact CCs in the early season, suggesting that leaf quality had a profound effect on larval mortality in the terrestrial microhabitat. In contrast, larval mortality did not differ between resuspended CCs and intact SCs in the late season, suggesting that leaf quality had little effect on larval mortality in the arboreal microhabitat. LMA was higher in CCs than in SCs, but there were no differences in the nitrogen and polyphenol contents. These results suggest that cradles constructed using mature, tough leaves were more effective against terrestrial cradle herbivores than those constructed using new, soft leaves.     

Multiple oviposition by Apoderus balteatus (Coleoptera, Attelabidae) is associated with larger leaves

ABSTRACT.

• 1 The female weevil of Apoderus balteatus Roelofs cuts the leaf near its base in order to make a leaf-roll‘cradle', in which one, two or three eggs are laid. Cuts are always located about 5 mm from the leaf base, independent of leaf size.
• 2 Larger leaves are made into larger cradles.
• 3 Weevils tend to lay more eggs in large cradles.
• 4 Eggs are laid through different holes and separated by leaf layers in the cradle. Cannibalism by larvae will be prevented by this isolated oviposition.
• 5 Multiple oviposition of this weevil allows larger leaves to be used by multiple larvae. It facilitates more efficient use of larger leaves and economizes on female effort in cradle formation.

     

The release of primordia of marginal buds onbryophyllum crenatum leaves from growth inhibition in relationship to the level of endogenous IAA

The growth of primordia of marginal buds (marginals) which differentiate on leaf margins is correlatively inhibited on intactBryophyllum crenatum plants. Following leaf isolation, the marginals are released from this correlative inhibition, which process is accompanied within 2 to 10 h after leaf isolation with a decrease in the content of endogenous IAA in the leaf blade. This decrease can be enhanced by transversal cutting of the leaf blade into three parts which also results in enhanced subsequent growth of the marginals. The growth which follows after the release of the marginals from correlative inhibition is accompanied in cut leaf blades with an increased content of endogenous IAA in the period from 12 h to 7 d after leaf isolation when compared with uncut leaf blades. The highest content of endogenous IAA was recorded in the middle section, and the lowest IAA content in the basal section of the leaf blade.     

Why are leaf‐cutting ants more common in early secondary forests than in old‐growth tropical forests? An evaluation of the palatable forage hypothesis

I evaluated the hypothesis that leaf‐cutting ants are more common in early successional forests than in old‐growth forests because pioneer species, which dominate in early successional habitats, appear more susceptible to leafcutters than shade‐tolerant species, which dominate primary forests (palatable forage hypothesis). The relative importance of pioneer and shade‐tolerant species as plant resources for leaf‐cutting ant colonies was evaluated (1) by literature review of leaf‐cutting ants’ diet, and (2) experimentally, using field assays to determine leafcutter's selectivity. Pioneer species were harvested three times more frequently than shade‐tolerant species and made up the largest component of the diet in all the studies reviewed. The amount harvested was not correlated with the plant species abundance. In addition, leaves from pioneer plants were selected eight times more than leaves from shade‐tolerant species in the field assays. These results support the palatable forage hypothesis. Leafcutters probably select pioneer leaves because of their low level of chemical defenses and high nutrient content. The high availability of pioneer species in early successional forest probably decreases the cost to locate palatable resources. Therefore, early successional habitats support more ant colonies than old‐growth forests. On the other hand, the effective defense mechanisms of mature plant species and the high dispersion of palatable plants could explain the low density of leaf‐cutting ant colonies in old‐growth forests. The palatable forage hypothesis is compared with other hypotheses that explain leaf‐cutting ant density. The preference of foundress queens for forest clearings, the dependence of small colonies on herbs, and the importance of pioneer plant species for mature colonies (palatable forage hypothesis) can be considered complementary, because they focus on different stages of the colony's life history. Consequently, the availability of pioneer plants appears to be one of the most influential factors determining mature leaf‐cutting ant nest densities in Neotropical forests.     

The Multiple Impacts of Leaf‐Cutting Ants and Their Novel Ecological Role in Human‐Modified Neotropical Forests

Herbivory has been identified as a potent evolutionary force, but its ecological impacts have been frequently underestimated. Leaf‐cutting ants represent one of the most important herbivores of the Neotropics and offer an interesting opportunity to address the role played by herbivorous insects through a perspective that embraces population‐ to ecosystem‐level effects. Here we: (1) qualitatively summarize the multiple ways leaf‐cutting ants interact with food plants and their habitats and elucidate the ultimate outcome of such interactions at multiple organization levels; (2) update our understanding of leaf‐cutting ant‐promoted disturbance regimes; and (3) examine potential ecological roles by leaf‐cutting ants within the context of human‐modified landscapes to guide future research agendas. First, we find that leaf‐cutting ants show that some herbivorous insects are able to generate ecologically important disturbance regimes via non‐trophic activities. Second, impacts of leaf‐cutting ants can be observed at multiple spatio‐temporal scales and levels of biological organization. Third, ecosystem‐level effects from leaf‐cutting ants are ecosystem engineering capable not only of altering the abundance of other organisms, but also the successional trajectory of vegetation. Finally, effects of leaf‐cutting ants are context‐dependent, species‐specific, and synergistically modulated by anthropogenic interferences. Future research should examine how leaf‐cutting ants respond to deforestation and influence remaining vegetation in human‐modified landscapes. By promoting either heterogeneity or homogeneity, leaf‐cutting ants operate not only as agricultural pests but also as ecological key players.     

Social Behavior of Larvae of the Neotropical Processionary Weevil Phelypera distigma (Boheman) (Coleoptera: Curculionidae: Hyperinae)

Socially gregarious behavior among free‐living leaf‐eating insect larvae occurs mostly among Lepidoptera, Symphyta, and a few Chyrsomelidae (Coleoptera). However, the Neotropical hyperine curculionid Phelypera distigma has also evolved this lifestyle, exhibiting a suite of social behaviors unique among beetles. The larvae are nomadic processionary foragers that punctuate foraging bouts with rosette‐shaped resting formations (cycloalexy). Larvae also vibrate or bob their heads rapidly when moving, especially when in contact with conspecifics, and this suggests acoustic or vibrational communication. In this study we used observational and experimental approaches to investigate the basis of processionary, cycloalexic,and head‐vibration behavior of this species. Larvae used both trail pheromones and thigmotactic signals to organize themselves into head‐to‐tail processionary columns. The trail pheromone, produced from the center of the abdomen, remains active for up to 4 h. Processions are not consistently led by particular individuals, but dynamically change over time and often temporarily break into two or more subprocessions. Subprocessions reunite through use of the trail pheromone. We found no evidence that head‐bobbing generates attraction through substrate‐borne or acoustic signals, but this behavior functions in direct contact to excite group activity. Time‐lapse videography used to analyze cycloalexic group formation showed that larvae transition from feeding in a line along the leaf margin to cycloalexic formations on the upper leaf surface via a coordinated back‐up movement that brings the posterior tip of their abdomens into contact. We identify three phases of cycloalexic formation: line‐up, back‐up, and an adjustment phase. Complete assembly can be achieved in as little as 5 s, but often the two phases establishing the basic rosette lasts 5–10 min, while the adjustment phase slowly tightens the group over a period of up to an hour. Collectively these studies present the first documented case of chemical trail marking in a beetle, and provide insight into a remarkable social‐behavioral repertoire convergent in key respects with the better‐studied social caterpillars and sawflies.     

Geometric morphometrics and leaf phenotypic plasticity: assessing fluctuating asymmetry and allometry in European white oaks (Quercus)

Because plants are unable to move away from unfavourable habitats and environmental perturbations, leaf phenotypic plasticity facilitates light absorption and gas exchange. Oaks (Quercus spp.) are particularly known for their adaptability and plastic phenotypes, and leaf allometry and developmental instability may represent important mechanisms for their adaptation to environments and evolution. Because of its important role in the adaptation of plant populations to different environments, allometry can be involved in diversifying selection. Developmental instability is related to environmental perturbations and stresses by producing random deviations in structures characterized by bilateral symmetry, such as oak leaves. In addition, developmental instability can also arise from genetic bottlenecks or as a result of hybridization. The splitting of symmetric and asymmetric components of variation and their separate analysis allows the variability in leaf shape traits to be summarized, reducing the variation produced by developmental instability. The geometric morphometric approach is a useful method for the study of leaf asymmetry and allometric patterns. This method provides an important tool for the visualization of shape attributes that characterize species with highly variable leaf phenotypic patterns. In this study, leaf shape and size variability of three white oak species was investigated by means of a two‐dimensional landmark‐based method providing improved knowledge of variance partitioning, species discrimination, fluctuating asymmetry and allometric patterns of variation resulting from the different analyses. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 335–348.     

Seedling recruitment in a semi‐arid Patagonian steppe: Facilitative effects of refuse dumps of leaf‐cutting ants

Question: What is the influence of refuse dumps of leaf‐cutting ants on seedling recruitment under contrasting moisture conditions in a semi‐arid steppe? Location: Northwestern Patagonia, Argentina. Methods: In a greenhouse experiment, we monitored seedling recruitment in soil samples from refuse dumps of nests of the leaf‐cutting ant Acromyrmex lobicornis and non‐nest sites, under contrasting moisture conditions simulating wet and dry growing seasons. Results: The mean number of seedling species and individuals were higher in wet than in dry plots, and higher in refuse dump plots than in non‐nest soil plots. The positive effect of refuse dumps on seedling recruitment was greater under low moisture conditions. Both the accumulation of discarded seeds by leaf‐cutting ants and the passive trapping of blowing‐seeds seems not explain the increased number of seeds in refuse dumps. Conversely, refuse dumps have higher water retention capacity and nutrient content than adjacent non‐nest soils, allowing the recruitment of a greater number of species and individual seedlings. Conclusions: Nests of A. lobicornis may play an important role in plant recruitment in the study area, allowing a greater number of seedlings and species to be present, hence resulting in a more diverse community. Moreover, leaf‐cutting ant nests may function as nurse elements, generating safe sites that enhance the performance of neighbouring seedlings mainly during the driest, stressful periods.     

The<Emphasis Type="Italic"> Arabidopsis thaliana rlp</Emphasis> mutations revert the ectopic leaf blade formation conferred by activation tagging of the<Emphasis Type="Italic"> LEP</Emphasis> gene

Activation tagging of the gene LEAFY PETIOLE ( LEP) with a T-DNA construct induces ectopic leaf blade formation in Arabidopsis, which results in a leafy petiole phenotype. In addition, the number of rosette leaves produced prior to the onset of bolting is reduced, and the rate of leaf initiation is retarded by the activation tagged LEP gene. The ectopic leaf blade results from an invasion of the petiole region by the wild-type leaf blade. In order to isolate mutants that are specifically disturbed in the outgrowth of the leaf blade, second site mutagenesis was performed using ethane methanesulphonate (EMS) on a transgenic line that harbours the activation-tagged LEP gene and exhibits the leafy petiole phenotype. A collection of revertant for leafy petiole ( rlp) lines was isolated that form petiolated rosette leaves in the presence of the activated LEP gene, and could be classified into three groups. The class III rlp lines also display altered leaf development in a wild-type (non-transgenic) background, and are probably mutated in genes that affect shoot or leaf development. The rlp lines of classes I and II, which represent the majority of revertants, do not affect leaf blade outgrowth in a wild-type (non-transgenic) background. This indicates that LEP regulates a subset of the genes involved in the process of leaf blade outgrowth, and that genetic and/or functional redundancy in this process compensates for the loss of RLP function during the formation of the wild-type leaf blade. More detailed genetic and morphological analyses were performed on a selection of the rlp lines. Of these, the dominant rlp lines display complete reversion of (1) the leafy petiole phenotype, (2) the reduction in the number of rosette leaves and (3) the slower leaf initiation rate caused by the activation-tagged LEP gene. Therefore, these lines are potentially mutated in genes for interacting partners of LEP or in downstream regulatory genes. In contrast, the recessive rlp lines exhibit a specific reversion of the leafy petiole phenotype. Thus, these lines are most probably mutated in genes specific for the outgrowth of the leaf blade. Further functional analysis of the rlp mutations will contribute to the dissection of the complex pathways underlying leaf blade outgrowth.Communicated by G. Jürgens     

Larval descriptions of five Byctiscus species (Coleoptera: Rhynchitidae) in Korea

The genus Byctiscus reported in Korea in five of nine species (B. lacunipennis, B. rugosus, B. puberulus puberulus, B. princeps and B. coerulans) is described for the last instar larvae. In addition, the cradle structure and host‐plants are provided.     

Quantifying leaf venation patterns: two-dimensional maps

The leaf vasculature plays crucial roles in transport and mechanical support. Understanding how vein patterns develop and what underlies pattern variation between species has many implications from both physiological and evolutionary perspectives. We developed a method for extracting spatial vein pattern data from leaf images, such as vein densities and also the sizes and shapes of the vein reticulations. We used this method to quantify leaf venation patterns of the first rosette leaf of Arabidopsis thaliana throughout a series of developmental stages. In particular, we characterized the size and shape of vein network areoles (loops), which enlarge and are split by new veins as a leaf develops. Pattern parameters varied in time and space. In particular, we observed a distal to proximal gradient in loop shape (length/width ratio) which varied over time, and a margin-to-center gradient in loop sizes. Quantitative analyses of vein patterns at the tissue level provide a two-way link between theoretical models of patterning and molecular experimental work to further explore patterning mechanisms during development. Such analyses could also be used to investigate the effect of environmental factors on vein patterns, or to compare venation patterns from different species for evolutionary studies. The method also provides a framework for gathering and overlaying two-dimensional maps of point, line and surface morphological data.     

Anatomy of vegetative organs,inflorescence axis and pedicel in the Neoregelia bahiana complex (Bromeliaceae): taxonomic and ecological importance

Delimitation of genera and species in Bromeliaceae is often problematic, for example in the Neoregelia bahiana complex which is distributed throughout the rocky fields of the Espinhaço Range, Brazil. Considering that the anatomical characterisation of different organs is potentially important for taxonomic and ecological interpretation of this complex, we analysed roots, stems (stolon), leaves, inflorescence axes (peduncle) and pedicels in individuals from different populations. In all the studied individuals, the roots are composed of velamen, a heterogeneous cortex, and a polyarch vascular cylinder with sclerenchymatous pith. The stolon features a parenchymatous cortex and collateral vascular bundles randomly distributed in the vascular cylinder. This organ may increase in diameter by the formation of new vascular bundles and a multi‐layered cork. The leaf blade has epidermal cells with U‐shaped thickened walls and peltate scales occur on the adaxial surface. The mesophyll consists of mechanical and water‐storage hypodermis and a heterogeneous chlorenchyma. The inflorescence axis and the pedicel have a parenchymatous cortex and vascular bundles randomly distributed in an aerenchyma. Some variable leaf characters, such as presence of air lacunae in the mesophyll, are related to the size of the individuals and were interpreted as phenotypic variations related mainly to sunlight incidence. In contrast, leaf characters such as lamina shape, distribution of the peltate scales, and number of cell layers forming the water‐storage hypodermis distinguish the populations of the Serra do Cabral and one population of the Diamantina (Minas Gerais) from the remaining studied populations, suggesting the existence of more than one taxon.     

Morphological Variation in Leaf Dissection of Rheum palmatum Complex (Polygonaceae)

Aims

Rheum palmatum complex comprises all taxa within section Palmata in the genus Rheum, including R. officinale, R. palmatum, R. tanguticum, R. tanguticum var. liupanshanense and R. laciniatum. The identification of the taxa in section Palmata is based primarily on the degree of leaf blade dissection and the shape of the lobes; however, difficulties in species identification may arise from their significant variation. The aim of this study is to analyze the patterns of variation in leaf blade characteristics within and among populations through population-based sampling covering the entire distribution range of R. palmatum complex.

Methods

Samples were taken from 2340 leaves from 780 individuals and 44 populations representing the four species, and the degree of leaf blade dissection and the shape of the lobe were measured to yield a set of quantitative data. Furthermore, those data were statistically analyzed.

Important Findings

The statistical analysis showed that the degree of leaf blade dissection is continuous from lobed to parted, and the shape of the lobe is also continuous from broadly triangular to lanceolate both within and between populations. We suggested that taxa in section Palmata should be considered as one species. Based on the research on the R. palmatum complex, we considered that the quantitative characteristics were greatly influenced by the environment. Therefore, it is not reliable to delimitate the species according to the continuously quantitative vegetative characteristics.     

Taxonomy of the Genus Eusphalerum Kraatz (Coleoptera: Staphylinidae: Omaliinae) from South Korea

The eusphalerine single genus, Eusphalerum Kraatz contains about 230 species distributed in the Holearctic and Oriental regions. In East Asia, 74 species of the genus are studied and Zanetti reported six species from North Korea in 1993. They are known to feed pollen on flowers of shrubs and trees. As a result, they have peculiar molar structure. Members of the Eusphalerum are characterized by the combination of following features: body spindle‐shape and parallel‐side; eyes large and protruded, temple short; mandibles subtriangular and edentate, molar and scissorial area fused, medial surface of molar with grinding cones; maxillary palpomere 4 as wide as palpomere 3; antennae gradually wide apically, antennomere 10 almost as wide as long; tarsomeres 1–4 short and broad, ventral surface setose densely, tarsomere 5 as long as or longer than basal four combined. In this paper, we report four South Korean Eusphalerum species and provide illustrations of the diagnostic characters including first line drawings of spermatheca and accessary sclerite. The specimens studied are deposited in the Chungnam National University Insect Collection (CNUIC), Daejeon, Korea.     

Disease defence through generations: leaf‐cutter ants and their symbiotic bacteria

Microbial ecology of animals is taking on significance in the modern dialogue for the biology of species. Similar to a nuclear genome, the entire bacterial assemblage maintains an ancestral signal of the host's evolution leading to cophylogeny between the host and the microbes they harbour (Brucker & Bordenstein 2012b). The stability of such associations is of great interest as they provide a means for species to acquire new traits and genetic diversity that their own genomes lack (McFall‐Ngai et al. 2013). The role of gut microbiota, for example, in host health and nutrition is widely recognized and a shared characteristic among animals. The role of bacteria colonizing the outside surfaces of animals is less well understood, but rather than random colonization, these microbes on skin, cuticles, scales and feathers in many cases provide benefits to the host. The symbiosis of leaf‐cutter ants, their fungus gardens and their microbiota is a fascinating and complex system. Whether culture‐independent bacterial diversity on the cuticle of leaf‐cutter ants is high or highly constrained by subcuticular gland secretions is one prominent question. In this issue of Molecular Ecology, Andersen et al. (2013) show that leaf‐cutting ants, Acromyrmex echinatior, maintain a dominant and colony‐specific bacterium called Pseudonocardia on their cuticles (the laterocervical plates in particular). This bacterium is involved in protecting the ants and their fungal gardens from disease. Other fungus‐gardening attine species as well as soil and vegetation can harbour Pseudonocardia. However, it was previously unknown how stable the bacterial strain–ant colony association was through the lifetime of the colony.     

Leaf‐shape remodeling: programmed cell death in fistular leaves of Allium fistulosum

Some species of Allium in Liliaceae have fistular leaves. The fistular lamina of Allium fistulosum undergoes a process from solid to hollow during development. The aims were to reveal the process of fistular leaf formation involved in programmed cell death (PCD) and to compare the cytological events in the execution of cell death to those in the unusual leaf perforations or plant aerenchyma formation. In this study, light and transmission electron microscopy were used to characterize the development of fistular leaves and cytological events. Terminal deoxynucleotidyl transferase‐mediated dUTP nick end labeling (TUNEL) assays and gel electrophoresis were used to determine nuclear DNA cleavage during the PCD. The cavity arises in the leaf blade by degradation of specialized cells, the designated pre‐cavity cells, in the center of the leaves. Nuclei of cells within the pre‐cavity site become TUNEL‐positive, indicating that DNA cleavage is an early event. Gel electrophoresis revealed that DNA internucleosomal cleavage occurred resulting in a characteristic DNA ladder. Ultrastructural analysis of cells at the different stages showed disrupted vacuoles, misshapen nuclei with condensed chromatin, degraded cytoplasm and organelles and emergence of secondary vacuoles. The cell walls degraded last, and residue of degraded cell walls aggregated together. These results revealed that PCD plays a critical role in the development of A. fistulosum fistular leaves. The continuous cavity in A. fistulosum leaves resemble the aerenchyma in the pith of some gramineous plants to improve gas exchange.     

LeafJ: An ImageJ Plugin for Semi-automated Leaf Shape Measurement

High throughput phenotyping (phenomics) is a powerful tool for linking genes to their functions (see review¹ and recent examples^2-4). Leaves are the primary photosynthetic organ, and their size and shape vary developmentally and environmentally within a plant. For these reasons studies on leaf morphology require measurement of multiple parameters from numerous leaves, which is best done by semi-automated phenomics tools^5,6. Canopy shade is an important environmental cue that affects plant architecture and life history; the suite of responses is collectively called the shade avoidance syndrome (SAS)⁷. Among SAS responses, shade induced leaf petiole elongation and changes in blade area are particularly useful as indices⁸. To date, leaf shape programs (e.g. SHAPE⁹, LAMINA¹⁰, LeafAnalyzer¹¹, LEAFPROCESSOR¹²) can measure leaf outlines and categorize leaf shapes, but can not output petiole length. Lack of large-scale measurement systems of leaf petioles has inhibited phenomics approaches to SAS research. In this paper, we describe a newly developed ImageJ plugin, called LeafJ, which can rapidly measure petiole length and leaf blade parameters of the model plant Arabidopsis thaliana. For the occasional leaf that required manual correction of the petiole/leaf blade boundary we used a touch-screen tablet. Further, leaf cell shape and leaf cell numbers are important determinants of leaf size¹³. Separate from LeafJ we also present a protocol for using a touch-screen tablet for measuring cell shape, area, and size. Our leaf trait measurement system is not limited to shade-avoidance research and will accelerate leaf phenotyping of many mutants and screening plants by leaf phenotyping.     

Phylloclade anatomy in Phyllanthus section Choretropsis (Phyllanthaceae)

The phylloclade anatomy of nine Brazilian species of the genus Phyllanthus section Choretropsis was studied, and its systematic significance was analysed. The arrangement of epidermal cells varied according to the species, with the guard cells of the stomatal apparatus distributed randomly or with the longitudinal axis parallel to the branch. In both branch types, the stomata were distributed on the phylloclade surface, except in P. gladiatus which showed hypostomatic branches. In most of the species, phylloclades developed a vascular system surrounded by clusters of fibres and cortical parenchyma, which could differentiate into palisade‐like tissue and ground tissue like a true leaf blade. The distribution of the vascular system followed the cylindrical or flattened diameter shape of the branch. In subsection Choretropsis, the cylindrical branches of P. choretroides, contrary to other species, developed a second ring formed by clusters of fibres surrounded by large parenchymal cells. In subsection Applanata, the palisade‐like cells of P. flagelliformis phylloclades acquired an isodiametric shape. The highest level of specialization was observed in P. gladiatus, which developed phylloclades with a dorsiventral structure, a main vein, and small sets of lateral vascular bundles like a typical leaf blade. The tissue arrangement of different species showed a gradual pattern, which was correlated with the different branch morphology and plant architecture. Despite the transitional pattern, it was suggested that some of the phylloclades had an independent origin and convergent evolution in the Choretropsis section. The anatomy of different branch types was shown to be useful, either in isolation or in association with other morphological characters, to delimit the species and to further understand their relationships and phylogeny. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 91–102.