Can't tell the caterpillars from the trees: countershading enhances survival in a woodland

Perception of the body's outline and three-dimensional shape arises from visual cues such as shading, contour, perspective and texture. When a uniformly coloured prey animal is illuminated from above by sunlight, a shadow may be cast on the body, generating a brightness contrast between the dorsal and ventral surfaces. For animals such as caterpillars, which live among flat leaves, a difference in reflectance over the body surface may degrade the degree of background matching and provide cues to shape from shading. This may make otherwise cryptic prey more conspicuous to visually hunting predators. Cryptically coloured prey are expected to match their substrate in colour, pattern and texture (though disruptive patterning is an exception), but they may also abolish self-shadowing and therefore either reduce shape cues or maintain their degree of background matching through countershading: a gradation of pigment on the body of an animal so that the surface closest to illumination is darker. In this study, we report the results from a series of field experiments where artificial prey resembling lepidopteran larvae were presented on the upper surfaces of beech tree branches so that they could be viewed by free-living birds. We demonstrate that countershading is superior to uniform coloration in terms of reducing attack by free-living predators. This result persisted even when we fixed prey to the underside of branches, simulating the resting position of many tree-living caterpillars. Our experiments provide the first demonstration, in an ecologically valid visual context, that shadowing on bodies (such as lepidopteran larvae) provides cues that visually hunting predators use to detect potential prey species, and that countershading counterbalances shadowing to enhance cryptic protection.     

Geographic variation in oviposition preference for male and female host plants in a geometrid moth: implications for evolution of host choice

Several dioecious plant species exhibit sexual dimorphisms in defensive traits. However, the effects of sexual dimorphism on defense against herbivores remain poorly understood. Eurya japonica (Thunb.) (Theaceae) is a dioecious shrub that shows sexual dimorphism in the chemical defense of flower buds. Female calyces contain higher concentrations of total phenolics and condensed tannins than do male calyces. Male flower buds are edible for a florivore moth, Chloroclystis excisa (Butler) (Lepidoptera: Geometridae), whereas the female flower buds are lethal to the moth larvae. The moths prefer to oviposit on male over female E. japonica flower buds. As the moths also occur in areas lacking E. japonica, we tested whether the oviposition preference for E. japonica flower sex differed between moths sympatric and allopatric with E. japonica. The moths sympatric with E. japonica showed a stronger preference for male E. japonica than the moths allopatric with E. japonica. Our phylogeographic study using mitochondrial cytochrome oxidase subunit I gene sequences revealed little genetic differentiation between moth populations sympatric and allopatric with E. japonica. These results suggest that the adaptive oviposition preference for flower sex of E. japonica has evolved rapidly in C. excisa.     

Biological control: Premises,ecological input and Mimosa pigra in the wetlands of Australia's Top End

The adverse impacts of weeds on natural ecosystems, together with the inadequate outcomes from treating weeds as a symptom, have escalated interest in finding efficacious control methods. With the aim of protecting wetlands from invasive weeds, this contribution uses the woody shrub Mimosa pigra L. (mimosa) as a case history to examine the methodology of classical biological control and the reasons for the widely accepted 75% failure rate. Overall objectives are for all biocontrol agents to have the opportunity to fully express their potential and to insure that limited resources are spent wisely on attainable weed control. The three main conclusions were that (1) the premises on which biocontrol is based has restricted advancement of this method; (2) monitoring is the logical first step to improving the selection of agents and release sites; and (3) it may be more cost-effective to introduce fewer agents that have undergone agent/plant and host/home range pre- and post-release ecological studies. Weed control may remain elusive unless advantage is taken of every beneficial result. Innovative assistance to agents and the integration of different control methods may preserve a role for weed biological control and has the potential to be of great importance for future weed management. It is proposed that the climate in the Top End of the Northern Territory and the lack of competition on the floodplains has contributed to mimosa's invasiveness. Climate may also underly the difficulties faced by agents. Agents appear unable to impart effective control in the dry season because of low numbers which relates to mimosa's poor growth; nor in the wet season, when the impact from high numbers of agents is outstripped by mimosa's growth.     

Does immune function influence population fluctuations and level of parasitism in the cyclic geometrid moth?

Populations of the autumnal moth, Epirrita autumnata, exhibit cycles with high amplitudes in northernmost Europe, culminating in devastating outbreak densities at favourable sites. Parasitism by hymenopteran parasitoids has been hypothesised to operate with a delayed density dependence capable of producing the observed dynamics. It has also been hypothesised that insects in crowded conditions invest greatly in their immunity as a counter-measure to increased risk of parasitism and pathogen infections. Furthermore, inducible plant defences consequent to grazing by herbivorous insects may be linked to the performance of parasitoids and pathogens through increased immunocompetence of the herbivore feeding on the foliage, in which the defence induction has taken place. At ten sampling sites, we quantified larval abundance, outbreak status and percentage larval parasitism during an extended peak phase of a population cycle. These population level covariates, together with an individual pupal mass, were used to explain differences in the immune defence, measured as an encapsulation reaction to artificial antigen. We also conducted a field study for an investigation of the susceptibility of autumnal moth pupae to naturally occurring pupal parasitoids. We did not find obvious differences between the encapsulation rate of autumnal moths originating from the sites with different past and current larval densities and risks for parasitism. The best ranked statistical models included pupal mass and outbreak status as explanatory variables, although both showed only slight effects on the encapsulation rate. The host resistance test revealed positive relationships between the encapsulation rate, body size and percentage parasitism of the exposed pupae, indicating that pupal parasitoids chose, and/or survived better, in large host individuals irrespective of their encapsulation ability. Thus, our results do not provide support for the hypothesis that variation in the immune function drives or modulates population cycles of autumnal moths. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorised users.     

Faunal composition of geometrid moths changes with altitude in an Andean montane rain forest

Aim The objective of this study was to describe and interpret the changes in faunal composition in the moth family Geometridae (Lepidoptera) along a small‐scale elevational gradient in a tropical montane rain forest. This gradient was compared with a large‐scale latitudinal gradient in Europe. Location Investigations were carried out in the province Zamora‐Chinchipe in southern Ecuador along a gradient ranging from 1040 to 2677 m above sea level at twenty‐two sites. Methods Moths were sampled with light‐traps in three field periods in 1999 and 2000 and subsequently sorted and determined to species or morphospecies. Results We analysed 13,938 specimens representing 1010 species of geometrid moths. The proportional contribution of subtaxa to the local geometrid fauna changes along the elevational gradient at all systematic levels considered. While proportions of species of the subfamilies Ennominae, Sterrhinae and Geometrinae significantly decrease, the proportion of Larentiinae increases with increasing altitude. Changes also occur within the subfamilies Ennominae and Larentiinae. The host–plant specialist ennomine tribes Cassymini, Macariini and Palyadini completely vanish, and the proportion of the tribe Boarmiini decreases at high altitudes. In contrast, the remaining tribes (mostly comprising polyphagous species) either do not show proportional changes (Azelinini, Nacophorini, Nephodiini, Ourapterygini) or even increase (Caberini, ‘Cratoptera group’). Within Larentiinae, the species proportion of the genus Eois decreases, whereas concomitantly the proportion of Eupithecia increases. There is a remarkable similarity between the altitudinal patterns in Ecuador and those found along the latitudinal gradient in Europe. Main conclusions Species of the subfamily Larentiinae seem to be particularly well‐adapted to harsh environmental conditions, towards both high altitudes and latitudes. They might disproportionately profit from lower predation at higher altitudes. Many changes in the faunal composition can be explained by expected host–plant requirements of the species involved. Our results show that diversity estimates based on taxon ratios which are assumed to be constant must be regarded with caution because such ratios can change rapidly along environmental gradients.     

Parasitoid assemblages reared from geometrid defoliators (lepidoptera: geometridae) of larch and fir in the alps

1 Geometrid larvae were collected from larch and fir in the Swiss Alps in 1991–94. Eight geometrid species were found in sufficient numbers to rear out parasitoids: Agriopis aurantiaria, Bupalus piniarius, Epirrita autumnata, Eupithecia lariciata, Odontopera bidentata, Lycia isabellae and Semiothisa liturata were collected from larch, and Puengeleria capreolaria was obtained from fir. 2 Parasitoid species belonging to five different guilds and four families were obtained; however, the taxonomic status of some of these is not completely resolved. 3 The parasitoid complex of the larch‐feeding species was totally different from that of Tortricidae and Tenthredinidae, which feed on the same host tree. In general, there was little overlap in the parasitoid complexes of the larch geometrids, with the most dominant parasitoid of each species reared from only one host. By contrast, many parasitoid species found during our study are also known to attack the same hosts or closely related hosts on different host tree species in different environments, suggesting that host specificity in geometrid parasitoids is more related to host taxonomy than to host plant or habitat.     

Sex-specific reproductive trade-offs in the gregarious fucoid macroalga Ascophyllum nodosum

The existence of sex-specific reproductive trade-offs is well established in plants. They usually occur because females invest more resources into reproduction than males, and have to compensate by sacrificing growth or defence. Investigations into by-sex differences in resource allocation by seaweeds are comparatively scarce. A small number of authors report differences between the sexes in some red algae, but known by-sex differences in brown algae are largely confined to the reproductive structures. In this study, sex-specific reproductive trade-offs are investigated in the common temperate brown alga Ascophyllum nodosum from three distinct populations. Quantified investments into growth, defence, and reproduction of a large number of individuals (n = 720) selected across a full annual cycle are presented in parallel with feeding trials using a common gastropod herbivore and a comprehensive assessment of the biotic and abiotic stressors impacting on A. nodosum at three sites. These reveal that sex-specific reproductive trade-offs occur in two of the three populations, as females invest more into reproduction than males and are subsequently less chemically defended for the months post gamete release. Feeding trials confirm that this leaves females more vulnerable to grazing pressure during these months, although mortality and competitive ability appear unaffected in the field. Possible causes of the trade-offs made by females are discussed, and new avenues of investigation are identified which could reveal interesting parallels between seaweeds and higher plants.     

A field study with geometrid moths to test the coevolution hypothesis of red autumn colours in deciduous trees

Red autumn colouration of trees is the result of newly synthesized anthocyanin pigments in senescing autumn leaves. As anthocyanin accumulation is costly and the trait is not present in all species, anthocyanins must have an adaptive significance in autumn leaves. According to the coevolution hypothesis of autumn colours, red autumn leaves warn herbivorous insects – especially aphids that migrate to reproduce in trees in the autumn – that the tree will not be a suitable host for their offspring in spring due to a high level of chemical defence or lack of nutrients. The signalling allows trees to avoid herbivores and herbivores to choose better host trees. In this study the coevolution hypothesis was tested with four deciduous tree species that have red autumn leaf colouration – European aspen (Populus tremula L.) (Salicaceae), rowan (Sorbus aucuparia L.) (Rosaceae), mountain birch [Betula pubescens ssp. czerepanovii (NI Orlova) Hämet‐Ahti], and dwarf birch (Betula nana L.) (Betulaceae), and with two generalist herbivores, the autumnal moth [Epirrita autumnata (Borkhausen)] and the winter moth [Operophtera brumata (L.)] (both Lepidoptera: Geometridae). Anthocyanin concentrations of autumn leaves were determined from leaf samples and the growth performance parameters of the moth larvae on the study trees were measured in the spring. Trees with higher anthocyanin concentration in the autumn were predicted to be low‐quality food for the herbivores. Our results clearly showed that anthocyanin concentration was not correlated with the growth performance of the moths in any of the studied tree species. Thus, our study does not support the coevolution hypothesis of autumn colours.     

Nest covering in plovers: How modifying the visual environment influences egg camouflage

Camouflage is one of the most widespread antipredator defences, and its mechanistic basis has attracted considerable interest in recent years. The effectiveness of camouflage depends on the interaction between an animal's appearance and its background. Concealment can therefore be improved by changes to an animal's own appearance, by behaviorally selecting an optimal background, or by modifying the background to better match the animal's own appearance. Research to date has largely focussed on the first of these mechanisms, whereas there has been little work on the second and almost none on the third. Even though a number of animal species may potentially modify their environment to improve individual‐specific camouflage, this has rarely if ever been quantitatively investigated, or its adaptive value tested. Kittlitz's plovers (Charadrius pecuarius) use material (stones and vegetation) to cover their nests when predators approach, providing concealment that is independent of the inflexible appearance of the adult or eggs, and that can be adjusted to suit the local surrounding background. We used digital imaging and predator vision modeling to investigate the camouflage properties of covered nests, and whether their camouflage affected their survival. The plovers' nest‐covering materials were consistent with a trade‐off between selecting materials that matched the color of the eggs, while resulting in poorer nest pattern and contrast matching to the nest surroundings. Alternatively, the systematic use of materials with high‐contrast and small‐pattern grain sizes could reflect a deliberate disruptive coloration strategy, whereby high‐contrast material breaks up the telltale outline of the clutch. No camouflage variables predicted nest survival. Our study highlights the potential for camouflage to be enhanced by background modification. This provides a flexible system for modifying an animal's conspicuousness, to which the main limitation may be the available materials rather than the animal's appearance.     

Empirical tests of the role of disruptive coloration in reducing detectability

Disruptive patterning is a potentially universal camouflage technique that is thought to enhance concealment by rendering the detection of body shapes more difficult. In a recent series of field experiments, artificial moths with markings that extended to the edges of their 'wings' survived at higher rates than moths with the same edge patterns inwardly displaced. While this result seemingly indicates a benefit to obscuring edges, it is possible that the higher density markings of the inwardly displaced patterns concomitantly reduced their extent of background matching. Likewise, it has been suggested that the mealworm baits placed on the artificial moths could have created differential contrasts with different moth patterns. To address these concerns, we conducted controlled trials in which human subjects searched for computer-generated moth images presented against images of oak trees. Moths with edge-extended disruptive markings survived at higher rates, and took longer to find, than all other moth types, whether presented sequentially or simultaneously. However, moths with no edge markings and reduced interior pattern density survived better than their high-density counterparts, indicating that background matching may have played a so-far unrecognized role in the earlier experiments. Our disruptively patterned non-background-matching moths also had the lowest overall survivorship, indicating that disruptive coloration alone may not provide significant protection from predators. Collectively, our results provide independent support for the survival value of disruptive markings and demonstrate that there are common features in human and avian perception of camouflage.     

Experiments on the responses of the larval stages of the geometrid moth Ectropis excursaria to light and gravity

The intensity-response curve to light in a vertical plane (where gravity was a conflicting stimulus to light) and a horizontal plane (where gravity was not a conflicting stimulus) suggests that the climbing behaviour of the first instar Ectropis excursaria (Guénée) is consistent with a summation of response to gravity and a response to light. Response to gravity was affected by the curvature of the crawling surface and the level of food deprivation of the caterpillars. On a 2 cm diameter crawling surface orientation to gravity occurred at an inclination between 20° and 30° from the horizontal. In a centrifugal field, a behaviour analogous to negative geotaxis occurred at accelerations of 3.5 ms^–2. Analysis of the geotactic turning tendencies of the caterpillar on flat inclined planes suggests that they exhibit meta-geotactic behaviour from inclinations of 40 to 90°. Negative geotaxis was exhibited by the caterpillars of later stadia although a response to gravity was lacking in the prepupal wandering caterpillars. The pre-pupal stage was also characterized by a reversal of the response to light.

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Résumé Un comportement d'escalade du premier stade d'E. excursaria correspondant à l'addition des réponses à la gravité et à la lumière, est suggéré par la courbe d'intensité de la réponse à la lumière, dans un plan vertical (où la gravité constitue un stimulus en opposition avec la lumière) et un plan horizontal (où la gravité n'est pas en opposition). La réponse à la gravité a été modifiée par la courbure de la surface à escalader et du degré de privation d'aliment des chenilles. Sur une surface d'escalade de 2 cm de diamètre, l'orientation par rapport à la gravité se produit entre 20 et 30° par rapport à l'horizontale. Das run champ de centrifugation, un comportement analogue à une géotaxis négative se produit pour une accélération de 3,5 m s^–2. L'analyse des tendances géotactiques des chenilles à tourner sur des plans inclinés suggère qu'elles présentent un comportement métagéotactique pour les inclinaisons de 40 à 90°. Des expériences d'immobilisation laissent penser que les antennes jouent un rôle dans la perception de la gravité. Les chenilles de stades ultérieurs ont présenté une géotaxie négative bien que la réponse à la gravité disparaisse chez les chenilles vagabondes en prénymphe. La prénymphose est aussi caractérisée par une inversion des réponses à la lumière.

     

Herbivore-induced plant vaccination. Part I. The orchestration of plant defenses in nature and their fitness consequences in the wild tobacco Nicotiana attenuata

A plant's responses to attack from particular pathogens and herbivores may result in resistance to subsequent attack from the same species, but may also affect different species. Such a cross-resistance, called immunization or vaccination, can benefit the plant, if the fitness consequences of attack from the initial attacker are less than those from subsequent attackers. Here, we report an example of naturally occurring vaccination of the native tobacco plant, Nicotiana attenuata, against Manduca hornworms by prior attack from the mirid bug, Tupiocoris notatus (Dicyphus minimus), which results from the elicitation of two categories of induced plant responses. First, attack from both herbivore species causes the plants in nature to release predator-attracting volatile organic compounds (VOCs), and the attracted generalist predator, Geocoris pallens, preferentially attacks the less mobile hornworm larvae. Second, attack from both mirids and hornworms increases the accumulation of secondary metabolites and proteinase inhibitors (PIs) in the leaf tissue, which is correlated with the slow growth of Manduca larvae. Mirid damage does not significantly reduce the fitness of the plant in nature, whereas attack from the hornworm reduces lifetime seed production. Consequently, plants that are attacked by mirids realize a significant fitness advantage in environments with both herbivores. The combination of growth-slowing direct defenses and predator-attracting indirect defenses results in greater hornworm mortality on mirid-attacked plants and provides the mechanism of the vaccination phenomenon.     

Intraspecific variation in herbivore‐induced plant volatiles influences the spatial range of plant–parasitoid interactions

Chemical information influences the behaviour of many animals, thus affecting species interactions. Many animals forage for resources that are heterogeneously distributed in space and time, and have evolved foraging behaviour that utilizes information related to these resources. Herbivore‐induced plant volatiles (HIPVs), emitted by plants upon herbivore attack, provide information on herbivory to various animal species, including parasitoids. Little is known about the spatial scale at which plants attract parasitoids via HIPVs under field conditions and how intraspecific variation in HIPV emission affects this spatial scale. Here, we investigated the spatial scale of parasitoid attraction to two cabbage accessions that differ in relative preference of the parasitoid Cotesia glomerata when plants were damaged by Pieris brassicae caterpillars. Parasitoids were released in a field experiment with plants at distances of up to 60 m from the release site using intervals between plants of 10 or 20 m to assess parasitism rates over time and distance. Additionally, we observed host‐location behaviour of parasitoids in detail in a semi‐field tent experiment with plant spacing up to 8 m. Plant accession strongly affected successful host location in field set‐ups with 10 or 20 m intervals between plants. In the semi‐field set‐up, plant finding success by parasitoids decreased with increasing plant spacing, differed between plant accessions, and was higher for host‐infested plants than for uninfested plants. We demonstrate that parasitoids can be attracted to herbivore‐infested plants over large distances (10 m or 20 m) in the field, and that stronger plant attractiveness via HIPVs increases this distance (up to at least 20 m). Our study indicates that variation in plant traits can affect attraction distance, movement patterns of parasitoids, and ultimately spatial patterns of plant–insect interactions. It is therefore important to consider plant‐trait variation in HIPVs when studying animal foraging behaviour and multi‐trophic interactions in a spatial context.     

The dietary overlap between red kangaroos (Macropus rufus) and sheep (Ovis aries) in the arid rangelands of Australia

Abstract The interaction between red kangaroos and sheep in the more arid areas of Australia was examined by means of a large-scale manipulative experiment. The diets of red kangaroos and sheep grazing together and separately in large paddocks were studied to assess whether competition for food might influence diet selection in these species. Diets and the amount of intra- and interspecific dietary overlap varied in accordance with pasture conditions. At pasture biomasses ranging from 50 to 200 g m^?2 (dry weight [wt]), the diets of red kangaroos and sheep overlapped by 58–73%, with forbs and grasses being the major items in the diets of both species. The diets of both sheep and red kangaroos under different experimental regimes were similar. In dry times (pasture biomass 40–50 g m^?2 (dry wt) more shrubs were eaten by both species and the amount of dietary overlap tended to be lower (52–66%). The diets of kangaroos were similar between paddocks. However, in paddocks containing both species sheep consumed proportionately more chenopodaceous shrubs and fewer grasses than those in ‘kangaroo-free’ paddocks.     

Performance of Danaini larvae is affected by both exotic host plants and abiotic conditions

The consequences of the introduction of invasive plants for the diet of herbivorous insects have been little explored in nature where, potentially, abiotic and biotic factors operate. In this study, we examined the larval performance of two Neotropical Danaini butterflies when using either a native or an exotic Apocynaceae species as host plant in both field and laboratory experiments. Hosts greatly differ in their amount of latex exudation and other physicochemical traits, as well as in the amount of evolutionary time they have interacted with herbivores. First, herbivore performance on the hosts was investigated under laboratory conditions. Larvae of both Danaini species took more time to develop on the exotic host; larval survivorship did not vary between hosts. Second, first instar survivorship on both hosts was evaluated in two field sites, one site per host. To do so, in both sites half of the larvae were bagged (protected against both abiotic and biotic factors) while the remainder were nonbagged (exposed). The interaction between larval exposure with the use of the exotic host reduced larval survival. We concluded that the combined effects of host plant traits and abiotic factors reduced survival of herbivores in field conditions. Therefore, the performance of herbivores when using hosts of different origins should be considered together with the multiple ecological factors found in natural environments, as these factors can modify the result of plant–herbivore interactions.     

The role of herbivory in the macroevolution of vertebrate hormone dynamics

Vertebrates have high species‐level variation in circulating hormone concentrations, and the functional significance of this variation is largely unknown. We tested the hypothesis that interspecific differences in hormone concentrations are partially driven by plant consumption, based on the prediction that herbivores should have higher basal hormone levels to ‘outcompete’ plant endocrine disruptors. We compared levels of glucocorticoids (GCs), the hormones with the most available data, across 166 species. Using phylogenetically informed comparisons, we found that herbivores had higher GC levels than carnivores. Furthermore, we found that the previously described negative relationship between GC levels and body mass only held in herbivores, not carnivores, and that the effect of diet was greatest at extreme body sizes. These findings demonstrate the far‐reaching effects of diet on animal physiology, and provide evidence that herbivory influences circulating hormone concentrations. We urge future direct testing of the relationship between phytochemical load and GC levels.     

Females of Cotesia vestalis,a parasitoid of diamondback moth larvae,learn to recognise cues from aphid‐infested plants to exploit honeydew

1. To maximise their reproductive success, the females of most parasitoids must not only forage for hosts but must also find suitable food sources. These may be nectar and pollen from plants, heamolymph from hosts and/or honeydew from homopterous insects such as aphids. 2. Under laboratory conditions, females of Cotesia vestalis, a larval parasitoid of the diamondback moth (Plutella xylostella) which does not feed on host blood, survived significantly longer when held with cruciferous plants infested with non‐host green peach aphids (Myzus persicae) than when held with only uninfested plants. 3. Naïve parasitoids exhibited no preference between aphid‐infested and uninfested plants in a dual‐choice test, but those that had been previously fed aphid honeydew significantly preferred aphid‐infested plants to uninfested ones. 4. These results suggest that parasitoids that do not use aphids as hosts have the potential ability to learn cues from aphid‐infested plants when foraging for food. This flexible foraging behaviour could allow them to increase their lifetime reproductive success.     

Facing herbivory on the climb up: Lost opportunities as the main cost of herbivory in the wild yam Dioscorea praehensilis

Plants with simple architecture and strong constraints on their growth may offer critical insights into how growth strategies affect the tolerance of plants to herbivory. Although Dioscorea praehensilis, a wild yam of African forests, is perennial, both aerial apparatus and tuber are annually renewed. Each year, the tuber produces a single stem that climbs from the ground to the forest canopy. This stem bears no leaves and no branches until it reaches optimal light conditions. Once in the canopy, the plant's production fuels the filling of a new tuber before the plant dies back to the ground. We hypothesized that if deprived of ant defense, the leafless growth phase is a vulnerable part of the cycle, during which a small amount of herbivory entails a high cost in terms of loss of opportunity. We compared the growth of stems bearing ants or not as well as of intact stems and stems subjected to simulated or natural herbivory. Ants reduce herbivory; herbivory delays arrival to the canopy and shortens the season of production. Artificially prolonging the stem growth to the canopy increased plant mortality in the following year and, in surviving plants, reduced the stem diameter and likely the underground reserves produced. Tuber size is a key variable in plant performance as it affects both the size of the aerial apparatus and the duration of its single season of production. Aerial apparatus and tuber are thus locked into a cycle of reciprocal annual renewal. Costs due to loss of opportunity may play a major role in plant tolerance to herbivory, especially when architectural constraints interact with ecological conditions to shape the plant's growth strategy.     

Toxicity of acephate to larvae of gypsy moth as a function of host plant and bioassay method

We examined toxicity of acephate to third-instar gypsy moth, Lymantria dispar (L.) (Lepidoptera: Lymantriidae), under different conditions of administration method, availability of food to larvae during bioassay, host plant, and activity of detoxifying enzymes. Larvae that had been fed field-collected foliage of white alder (Alnus rhombifolia Nutt.) were less susceptible 48 h after treatment with topically applied acephate if they were allowed to continue feeding on foliage during the bioassay period (LD₅₀= 60.6 μg/g larva ) than if they were not (LD₅₀= 13.5 μg/g larva ). All surviving larvae were replaced on their original food plant after the 48-h bioassay; of these, 14.4% of the larvae not fed during treatment died before pupation, compared with 1.3% of the larvae fed alder during treatment. The LD₅₀ obtained for topically treated larvae reared and treated on Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, (51.1 μg/g larva) was comparable to that obtained for larvae fed alder (60.0 μg/g larva) throughout treatment. Larvae treated orally with acephate, however, were slightly more susceptible when reared on Douglas-fir (LC₅₀, 20.3 ppm ) than when reared on alder (LC₅₀, 27.0 ppm ). Post-treatment mortality in orally treated larvae was 10.3% in those fed alder and 9.5% in those fed Douglas-fir. Higher cytochrome P-450 activities in larvae reared on Douglas-fir apparently did not enhance tolerance to acephate. Both sexes of orally treated larvae took significantly longer to pupate than did controls on both foliage types, as did topically treated males fed Douglas-fir. Pupal weight generally was slightly, but not always significantly, higher in treated than untreated larvae under all dietary and treatment regimes.