Declining acceptability of a food plant for the polyphagous grasshopper Schistocerca americana: the role of food aversion learning

ABSTRACT Successive contacts with a food that was initially acceptable to the acridid Schistocerca americana (Drury) (Orthoptera: Acrididae) were monitored. It was found that acceptability, as measured by meal length, declined with experience until the food, spinach, was completely rejected. Reduced acceptability of spinach was maintained even after feeding for three meals on a fully acceptable and suitable food, a period of 2–3 h. Experiments with spinach-filled capsules placed in the gut suggested that the decline in acceptability was not due to direct feedback from the gut or haemolymph on the sensory or feeding control systems. Food aversion learning is indicated as the mechanism underlying the decline in acceptability.     

Bryophytes as food: comparative consumption and utilization of mosses by a generalist insect herbivore

Compared with angiosperms, bryophytes are seldom fed upon by insects, despite being commonly used for shelter. Bryophytes are assumed to be unpalatable, and three classes of mechanisms have been suggested as possible barriers to bryophagy: chemical defenses, low digestibility, and low nutrient content. However, very few studies have tested these hypotheses. The present study examines pre‐ and post‐ingestive defenses of mosses. The acceptability and quality of four species of moss –Bryum argenteum Hedw. (Bryales: Bryaceae), Climacium americanum Brid. (Leucodontales: Climaciaceae), Leucobryum glaucum (Hedw.) (Dicranales: Leucobryaceae), and Sphagnum warnstorfii Russ. (Sphagnales: Sphagnaceae) – were compared with two control diets using the generalist caterpillar, Trichoplusia ni Hübner (Lepidoptera: Noctuidae: Plusiinae). In no‐choice trials, caterpillars consumed much less of any of the mosses than lettuce or wheat germ. The only moss consumed in sufficient quantities to evaluate post‐ingestive responses was C. americanum. Digestibility, assimilation, and overall utilization efficiency of C. americanum did not differ from that of lettuce, although C. americanum and lettuce were both less digestible than artificial diet. Choice assays using leaf discs showed that ethanol extract of L. glaucum, the least consumed moss, was deterrent, implying that chemical defenses play a major role in deterring feeding on L. glaucum. This study suggested that pre‐ingestive mechanisms are more important than post‐ingestive mechanisms in discouraging herbivory on mosses, and offered evidence that mosses are not simply nutrient poor.     

Seasonal variations in the quality of mopane leaves as a source of browse for mammalian herbivores

This paper deals with seasonal variations in the chemical composition of mopane leaves in terms of their palatability to herbivores. Green mopane leaves were found to be most palatable over summer, a period when, due to very poor rainfall, the veld was at its most desolate. Senescing leaves were at their most palatable in late winter/early spring, this resource maintaining many herbivores for the short period before the spring leaf flush. Four leaf classes were identified during the spring leaf flush: young green leaves were the most palatable followed by very young leaves, mature green leaves and senescent leaves. The flush is independent of rain and coincides with the end of the dry season, a period when most tree species occurring in association with the mopane are leafless and the grass sward is very poor. It begins earlier on heavily browsed trees. Most ungulates present in the area were found to utilize the nutritious young leaves. The importance of this leaf flush is discussed with respect to the management of the area, and the importance of the heavy pressure large mammals are imposing on this veld type is elicited.     

The use of electrophoresis in the separation of two closely related species of terrestrial slugs

The pattern of variation of non-specific esterases, revealed by the use of disc electrophoresis, was used in an investigation of the specific difference between Limax flavus and L. pseudoflavus. The isozyme patterns were used to differentiate between the two taxa both qualitatively and quantitatively. The results were in agreement with the results from studies of the variation in the morphology of the distal portions of the genitalia and of external body colouration and patterning. The biochemical methods allowed a rapid estimation of the difference between the two taxa and were more precise than the use of colour variation.     

Biology of slugs (Agriolimacidae and Arionidae: Mollusca) in New Zealand hill country pastures

Summary Life cycles of the slugs Deroceras reticulatum and Arion intermedius were studied over a 2.5-year period in a sheep-grazed pasture in the Kaimai Range, New Zealand. D. reticulatum approximated a bivoltine phenology, with intervals between consecutive generations ranging from 4 to 7 months and maximum life span from 8 to 12 months. Egg laying occurred in autumn and spring-early summer. Periods of egg hatching were followed by a phase of juvenile growth characterised by linear increase in the logarithm of population mean body weights. This was followed by a phase of reproductive activity. A. intermedius had an annual life cycle. Eggs laid during late summer and autumn hatched during autumn-winter. Immature slugs were characterised by a period of low growth rate during winter followed by a period of rapid growth culminating in reproductive maturity. In both species, the phase of rapid growth rate was associated with enlargement of the hermaphrodite gland. The size of the hermaphrodite gland relative to body weight reached a maximum during the spermatozoon stage, generally coincident with maximum body weight. During the reproductive period of D. reticulatum and A. intermedius the body weight remained relatively stable but the hermaphrodite gland became progressively smaller as the slugs approached the post-reproductive stage. The albumen gland attained maximum weight at the oocyte stage of the hermaphrodite gland, at the onset of reproductive activity. The pasture, strongly dominated by grasses, was heterogenous in slope, vegetation and treading/grazing influences from sheep due to stratification of the habitat into contour tracks, associated with sheep movement around the slope contours, and intertrack areas. High population densities of D. reticulatum were associated with the track edges (kerbs) and the upper intertrack slopes, while densities of A. intermedius were highest for the intertrack slopes. Low densities of both species occurred on the tracks and at sheep campsites. Analysis of cohort life tables indicated that these dispersion patterns were important in the dynamics of the populations. The stage mortality contributing most to the variance in reproductive populations of both species was that between hatching and onset of reproduction; most of the mortality occurred shortly after hatching on south-facing slopes, but tended to occur later in the life cycle on north slopes. For D. reticulatum this mortality was inversely related to initial density and associated, at least in part, with predation by carabids and birds and with treading by sheep. Density relationships, both at whole plot and plot stratum level, indicated that variations in natality had a stabilizing influence on D. reticulatum populations.     

A simple enzyme assay for dry matter digestibility and its value in studying food selection by generalist herbivores

Summary The dry matter digestibility of 94 species of leaf was assayed by a simple method involving sequential treatment with pepsin and fungal cellulase enzymes. It was demonstrated that for foliage from rainforest trees of a wide range of dicotyledonous plant families the assay showed high positive correlation with estimates of dry matter digestibility obtained using rumenliquor from a fistulated steer. Both assays were found to reflect negative correlates of digestibility, notably fibre and condensed tannin, rather than the nutritional value of an item. The higher dry matter digestibility of immature leaves relative to mature leaves appeared to be accounted for by their lower fibre content. It is suggested that the pepsin/cellulase assay offers a cheap, quick, routine method of gaining information on the effects of some types of plant secondary compounds (digestibility reducers) on the food potential of different kinds of foliage to herbivores. Its use in studies of herbivory in rainforest areas in relation to analyses for plant secondary compounds and food selection by herbivores is discussed.Publication 20-018 of the Wisconsin Regional Primate Research Center     

The value of three cereal aphid species as food for a generalist predator

The value of the cereal aphid species Metopolophium dirhodum (Wlk.), Sitobion avenae (F.) and Rhopalosiphum padi (L.) as prey for the linyphiid spider Erigone atra (Bl.) was assessed. Fecundity of females was determined for spiders fed on eight experimental diets: three single‐species aphid diets, a mixed diet of all three aphid species, three mixed diets with each aphid species in combination with fruit flies Drosophila melanogaster (Meig.), and pure D. melanogaster as a high quality comparison diet. The development and survival of first‐instar juveniles fed on three diets of single aphid species, and on a diet of Collembola were compared with those subjected to starvation. Prey value for adult females was assessed by egg production, hatching success and offspring size. In pure diets all three aphid species were of low value to the spiders, causing a rapid decline in egg production and supporting no growth of significance of first‐instar juveniles. No difference in value of aphid species of single‐species aphid diets was found in the fecundity experiment, while a ranking of aphid species of M. dirhodum > R. padi > S. avenae was revealed in the survivorship experiment. A mixed‐aphid diet was not found to be advantageous compared with single‐species aphid diets, and no advantage of including aphids in mixed diets with fruit flies was found. Metopolophium dirhodum and R. padi were neutral in mixed diets, while a diet of S. avenae and fruit flies caused reduced egg production compared with the pure diet of fruit flies, revealing a toxic effect of S. avenae on the spider. The value‐ranking of aphid species in mixed diets was similar to that of single‐species diets. A similar ranking of aphid species was found for different fitness parameters (fecundity of adult females and development of juveniles). A ranking of aphids by offspring size of mothers on aphid‐only diets was S. avenae > M. dirhodum > R. padi. All aphid‐fruit fly diets resulted in larger offspring than a diet of only D. melanogaster, with the overall largest offspring being produced on the diet of M. dirhodum and fruit flies.     

Patterns of introduction of non-indigenous non-marine snails and slugs in the Hawaiian Islands

The native snails of the Hawaiian Islands are disappearing. One cause is predation by introduced carnivorous snails. Habitat destruction/modification is also important, facilitating the spread of other non-indigenous snails and slugs. Eighty-one species of snails and slugs are recorded as having been introduced. Thirty-three are established: 12 freshwater, 21 terrestrial. Two or three species arrived before western discovery of the islands (1778). During the nineteenth century about one species per decade, on average, was introduced. The rate rose to about four per decade during the twentieth century, with the exception of an especially large number introduced in the 1950s as putative biocontrol agents against the giant African snail, Achatina fulica. The geographical origins of these introductions reflect changing patterns of commerce and travel. Early arrivals were generally Pacific or Pacific rim species. Increasing trade and tourism with the USA, following its annexation of Hawaii, led to an increasing proportion of American species. More general facilitation of travel and commerce later in the twentieth century led to a significant number of European species being introduced. African species dominated the 1950s biological control introductions. The process continues and is just part of the homogenization of the unique faunas of tropical Pacific islands.     

The influence of large mammalian herbivores on growth form and utilization of mopane trees, Colophospermum mopane, in Botswana's Northern Tuli Game Reserve

Mopane veld is of great value to the general ungulate spectrum in times of drought, and it is capable of retaining its dominance in a community even in the presence of extremely heavy browsing pressure imposed by large browsing mammals. Scrub mopane (hedges) has been regarded as resulting from excessive browsing pressure by large mammals, especially elephants. Both the nutrient and chemical composition of mopane twig bark were investigated, the seasonal results being related back to the seasonal utilization of branches by large mammals. Mopane twigs were most palatable in winter. Eland feed on mopane throughout the year irrespective of palatability. Elephants were rarely present in the scrub mopane area before the onset of spring rains, when the major mopane leaf flush occurs independently of rainfall. The impact of both species was not excessive and recruitment of mopane seedlings does occur. Herbivore browsing is responsible for a mopane morph which buds early and continues to produce accessible, nutritious leaves even when heavily browsed. Many browsing ungulates are reliant on this resource during the stressful transition from spring to summer in south-eastern Botswana.     

Interactions between a fungal pathogen, foliar properties, and generalist herbivores

American [Castanea dentata (Marsh) Borkh.] × Chinese [Castanea mollissima Blume] chestnut (Fagac, ae) hybrids are a novel system in which to study influences of phytopathogenic fungi and woody plant hybridization on herbivore susceptibility, as the hybrids are well characterized with regard to resistance to the chestnut blight fungus [Cryphonectria parasita (Murr) Barr (Endothia) Diaporthales: Valsaceae] and variability is present. We chose two groups of resistance‐rated backcross chestnut that shared an F1 parent and had different American parents. Foliage from both backcross groups and the parent trees was sampled on three dates for use in feeding assays with gypsy moth larvae [Lymantria dispar (L.) [Lepidoptera: Lymantriidae], adult Japanese beetles [Popillia japonica Newman (Coleoptera: Scarabaeidae)], and fall webworm larvae [Hyphantria cunea Drury (Lepidoptera: Arctiidae)], respectively. Foliar analyses were performed concurrently and included carbohydrate, tannin, and nitrogen content, toughness, and density. Blight resistance had almost no effect on herbivore performance or foliar chemistry. When the parent trees and backcross groups were compared, however, significant differences in gypsy moth performance and Japanese beetle consumption were evident. There were no differences in fall webworm consumption. Most foliar characteristics measured differed among chestnut genotypes at some point in the season, and all varied seasonally. No clear pattern emerges with respect to the relationship among blight resistance, herbivore susceptibility, foliar properties, and plant genotype, and more research is needed to separate these effects.     

Specialist and generalist natural enemies as an explanation for geographical gradients in population cycles of northern herbivores

Within Fennoscandia, two well-studied groups of herbivores exhibit clear geographical gradients in their population dynamics. Populations of a forest lepidopteran ( Epirrita autumnata , the autumnal moth) and voles of the genera Microtus and Clethrionomys show pronounced multi-annual cycles in the north but become more stable towards the south. Here we review empirical and theoretical studies on these species, mainly regarding the biological mechanisms that are assumed to generate the pattern of population dynamics in both systems. We conclude that the specialist/generalist predation hypothesis offers a common explanation for the population cycles and their geographical gradients irrespective of whether a herbivorous insect or small mammals are concerned. According to this hypothesis, originally developed for the Fennoscandian voles, but now applied also to E. autumnata , population cycles are generated by specialist natural enemies (predators for the voles and parasitoids for E. autumnata ). Furthermore, the dynamic shift from cycles to stability is assumed to be caused by an increase in the density and diversity of generalist natural enemies from north to south in Fennoscandia.     

Evolutionary relationships among the Pulmonate land snails and slugs (Pulmonata, Stylommatophora)

We have incorporated an additional 56 species of land snails and slugs in our ribosomal (r) RNA molecular phylogeny. The new taxa include representatives of several important groups. The molecular tree now includes 160 species of stylommatophoran land snails and slugs in 144 genera in 61 families. In the rDNA tree, the Stylommatophora are principally divided into an 'achatinoid' and a 'non-achatinoid' clade. Within these clades, several major land snail groups, including the Orthurethra, Elasmognatha, Limacoidea, and Helicoidea, are supported. Overall, the rDNA molecular phylogeny has remained stable following the incorporation of the additional taxa, with these additions having little impact on the major evolutionary patterns in the tree. Taxonomic coverage of the Orthurethra, Orthalicidae, Camaenidae, and Bradybaenidae is increased significantly. The camaenids and bradybaenids form a complex, and both appear to be paraphyletic. Several families of uncertain affinity, such as the Sagdidae and Thyrophorellidae, are included for the first time. The Sagdidae are shown to belong to the Helicoidea, and the Thyrophorellidae to the Achatinoidea.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 593–610.     

Palatability of monoterpenoid compositional types of Satureja douglasii to a generalist molluscan herbivore,Ariolimax dolichophallus

The differential palatability of monoterpenoid compositional types of Satureja douglasii to a native slug herbivore, Ariolimax dolichophallus, was determined. Types containing a high proportion of bicyclic monoterpenoids (camphene, camphor) were more palatable than those containing high proportions of p-menthane monoterpenoids (isomenthone, pulegone or carvone). In the case of individual monoterpenoids characteristic of these types, camphor proved to be most palatable and pulegone the least; carvone and isomenthone were intermediate. Differential herbivory on the compositional types was observed under field conditions. It is suggested that differential herbivory could have been important in determining the patterns of monoterpenoid variation and their distribution in S. douglasii.     

Flexibility of digestive responses in two generalist herbivores,the tortoises Geochelone carbonaria and Geochelone denticulata

Summary To test the prediction that digestive responses digestibility, intake and passage time-of generalist herbivores vary with different diets, feeding trials were conducted in Venezuela with two sympatric tortoise species, Geochelone carbonaria and G. denticulata. Three single-species diets (two fruit, one foliage) were fed to both species. For a given diet, digestibility, mass-specific intake and passage time did not differ between the two tortoise species, nor did they vary by sex or body mass within each species. However, the digestive parameters varied for tortoises feeding on the different diets. The responses ranged from nearly abandoning cell wall fermentation and depending entirely on extraction of cell contents to relying heavily on cell wall fermentation. Therefore, these generalist herbivores have flexible digestive responses that are influenced by diet, not fixed digestive responses that limit the diet, as previously observed in other generalist herbivores. A three-part classification of herbivores (specialist, specialized mixed feeder and opportunistic mixed feeder) is suggested as an approach to understanding flexible and inflexible digestive strategies in herbivores.     

Cumulative consumption of the lake macrophyte Elodea by abundant generalist invertebrate herbivores

Previous investigations have shown that macrophyte biomass can be substantially reduced by invertebrate herbivores but have not provided evidence for the links between the magnitude of the observed damage and the densities of herbivores. The results of this study support the hypothesis that the abundant occurrence of the epiphytic generalist herbivores may result in their cumulative consumption which, in turn, can be regarded as the mechanism responsible for often observed relatively high level of herbivory on freshwater macrophytes. The percentage of Elodea sp. biomass consumed by invertebrates was estimated for six European lakes, based on analysis of gut contents, daily rations and the density of epiphytic herbivores. Although the daily ration of these invertebrates when feeding upon Elodea averaged only 14.6% of their dry mass, their biomass was relatively high (from 0.163 to 1.161 g DW per 100 g DW plant). The estimated percentage of Elodea biomass consumed during one summer month by epiphytic invertebrates ranged from 0.5 to 5.9%. These values, after extrapolating to the whole growing season would mean that the biomass of Elodea lost to herbivory was between about 2 and 23%, an estimate which are within the range of consumption reported by other authors.     

The acceptability of meadow plants to the slug Deroceras reticulatum and implications for grassland restoration

Background and Aims

Despite the selective pressure slugs may exert on seedling recruitment there is a lack of information in this context within grassland restoration studies. Selective grazing is influenced by interspecific differences in acceptability. As part of a larger study of how slug–seedling interactions may influence upland hay meadow restoration, an assessment of relative acceptability is made for seedlings of meadow plants to the slug, Deroceras reticulatum.

Methods

Slug feeding damage to seedling monocultures of 23 meadow species and Brassica napus was assessed in microcosms over 14 d. The severity and rate of damage incurred by each plant species was analysed with a generalized additive mixed model. Plant species were then ranked for their relative acceptability.

Key Results

Interspecific variation in relative acceptability suggested seedlings of meadow species form a hierarchy of acceptability to D. reticulatum. The four most acceptable species were Achillea millefolium and the grasses Holcus lanatus, Poa trivialis and Festuca rubra. Trifolium pratense was acceptable to D. reticulatum and was the second highest ranking forb species. The most unacceptable species were mainly forbs associated with the target grassland, and included Geranium sylvaticum, Rumex acetosa, Leontodon hispidus and the grass Anthoxanthum odoratum. A strong positive correlation was found for mean cumulative feeding damage and cumulative seedling mortality at day 14.

Conclusions

Highly unacceptable species to D. reticulatum are unlikely to be selectively grazed by slugs during the seedling recruitment phase, and were predominantly target restoration species. Seedlings of highly acceptable species may be less likely to survive slug herbivory and contribute to seedling recruitment at restoration sites. Selective slug herbivory, influenced by acceptability, may influence community-level processes if seedling recruitment and establishment of key functional species, such as T. pratense is reduced.     

Optimal foraging and community structure: implications for a guild of generalist grassland herbivores

Summary A particular linear programming model is constructed to predict the diets of each of 14 species of generalist herbivores at the National Bison Range, Montana. The herbivores have body masses ranging over seven orders of magnitude and belonging to two major taxa: insects and mammals. The linear programming model has three feeding constraints: digestive capacity, feeding time and energy requirements. A foraging strategy that maximizes daily energy intake agrees very well with the observed diets. Body size appears to be an underlying determinant of the foraging parameters leading to diet selection. Species that possess digestive capacity and feeding time constraints which approach each other in magnitude have the most generalized diets. The degree that the linear programming models change their diet predictions with a given percent change in parameter values (sensitivity) may reflect the observed ability of the species to vary their diets. In particular, the species which show the most diet variability are those whose diets tend to be balanced between monocots and dicots. The community-ecological parameters of herbivore body-size ranges and species number can possibly be related to foraging behavior.     

The enemy release and EICA hypothesis revisited: incorporating the fundamental difference between specialist and generalist herbivores

The success of invasive plants has been attributed to their escape from natural enemies and subsequent evolutionary change in allocation from defence to growth and reproduction. In common garden experiments with Senecio jacobaea, a noxious invasive weed almost worldwide, the invasive populations from North America, Australia, and New Zealand did indeed allocate more resources to vegetative and reproductive biomass. However, invasive plants did not show a complete change in allocation from defence to growth and reproduction. Protection against generalist herbivores increased in invasive populations and pyrrolizidine alkaloids, their main anti‐herbivore compounds, did not decline in invasive populations but were higher overall compared with native populations. In contrast, invasive plants lost additional protection against specialist herbivores adapted to pyrrolizidine alkaloids. Hence, the absence of specialist herbivores in invasive populations resulted in the evolution of lower protection against specialists and increased growth and reproduction, but also allowed a shift towards higher protection against generalist herbivores.