Variation in quality of mountain birch and tea-leaved willow for mammal and insect herbivores: differences among trees, among sites and between tree species

We evaluated palatability of winter dormant tree twigs to the mountain hare Lepus tunidus and the grey-sided vote Clethrionomys rufocanus with captive animals We tested differences among trees within sites, among sites, and between two tree species (the mountain birch Betula pubescens ssp czerepanovii and the tea-leaved willow Salix phylicifolia ) In one of the sites, we also measured growth rates of autumnal moth Epirrita autumnata larvae on the same trees that were used m preference trials with the hares and voles The differences m palatability to hares and voles were greatest at the level of tree Species, both hares and voles preferred birches over willows, but with the hare there was some overlap in palatability between the tree species There were also large and significant differences among sites and among trees within sites Within sites, variation in tree palatability seemed to be larger among willows than among birches Hares and voles may select willows at least partly on the same basis, but there was no correlation between palatability of the trees to the mammals and the growth rate of the larvae of the autumnal moth on the same trees Variation in twig palatability to the hares and voles was so large at all levels (among trees within sites, among sites and between tree species) that we suggest it also has implications for food selection of these mammalian herbivores under natural conditions     

Size class structure and tree dispersion patterns in old-growth cedar-hemlock forests of the northern Rocky Mountains (USA)

Summary The relationship between the food selection of four leaf beetle species (Phratora vitellinae, Plagiodera versicolora, Lochmaea capreae, Galerucella lineola) and the phenolic glycosides of willow (Salix spp.) leaves was tested in laboratory food choice experiments. Four willow species native to the study area (Eastern Finland) and four introduced, cultivated willows were tested.The willow species exhibited profound differences in their phenolic glycoside composition and total concentration. The food selection patterns of the leaf beetles followed closely the phenolic glycoside spectra of the willow species. Both the total amount and the composition of phenolic glycosides affected the feeding by the beetles. Phenolic glycosides apparently have both stimulatory and inhibitory influences on leaf beetle feeding depending on the degree of adaptation of a particular insect. Very rare glycosides or exceptional combination of several glycoside types seem to provide certain willow species with high level of resistance against most herbivorous insects. Analogously the average absolute amount of leaf beetle feeding was lower on the introduced willows than on the native species to which the local herbivores have a good opportunity to become adapted.     

Effects on heather of heavy grazing by mountain hares

Continual heavy grazing of small patches of heather by mountain hares produces conspicuous, short, lawn-like bushes. Their probable development is described. The shoots of such heather bushes maintained a juvenile appearance, contained more nitrogen and phosphorus than the surrounding heather and, to judge from their chemical composition, were more digestible. The generalization that tree and shrub twigs of juvenile growth form are more unpalatable to hares than twigs of mature growth form did not apply in this case.     

Do plant secondary compounds determine feeding preferences of snowshoe hares?

Summary We investigated the food preferences of captive snowshoc hares (Lepus americanus) in winter to test three hypotheses proposed to explain food choices by hares: (1) that food choice is related to the protein content of twigs; 92) that defensive chemicals present in twigs are negatively correlated with hare food preferences; and (3) that hares eat less-preferred but protein-rich twigs when their diet is buffered by large amounts of palatable food. Hares exhibited striking and consistent preferences for different species and, in general, preferred mature twigs to juvenile growth stages. Preferences across species among mature twigs were not, however, the same as preferences for juvenile growth stages across species. None of the three hypotheses adequately explained food choice by hares. Hares did not (1) select twigs that were high in protein content. They also did not (2) consistently select twigs that were low in resins or phenols. Finally (3), hares generally ate less, not more of non-preferred twigs in the presence of a protein and energy rich alternative food, commercial rabbit chow. Food preferences of hares must presumably have some chemical basis, but no simple theory has yet explained what this is. We suggest that hares may not be under severe dietary constraints imposed by chemical defenses in winter.     

Phenolic glucosides as feeding cues for willow-feeding leaf beetles

The effects of individual phenolic glucosides and total glucoside fractions on the feeding behaviour of three willowfeeding leaf beetles (Coleoptera: Chrysomelidae) were tested in the laboratory. Feeding preferences of the tested leaf beetles were strongly influenced by certain phenolic glucosides which are typical secondary compounds of willows (Salicaceae:Salix). Two of the tested leaf beetles,Galerucella lineola andLochmaea capreae showed rather similar responses to glucoside treatments. Both of them were strongly stimulated by total glucoside fractions fromSalix triandra and by its major glucoside salidroside. The third species,Phatora vitellinae, was attracted most by the fractions fromS. myrsinifolia andS. pentandra, and by two related salicylate glucosides, tremulacin and salicortin. Food selection pattern of the tested beetles in the laboratory concords fairly well with their distributions in the field and with the occurrence of phenolic glucosides in their host willows. Phenolic glucoside extracts stimulated more feeding than individual pure glucosides. This indicates that different compounds have synergistic effects in the feeding behaviour of leaf beetles. Our results clearly show that willow leaf beetles select their food based on phenolic glucosides of their host plants.     

Plant phenology-mediated indirect effects: The gall midge opens the phenological window wider for a leaf beetle

We examined whether larvae of the gall midge Rabdophaga rigidae (Diptera: Cecidomyiidae) can modify the seasonal dynamics of the density of a leaf beetle, Plagiodera versicolora (Coleoptera: Chrysomelidae), by modifying the leaf flushing phenology of its host willow species, Salix serissaefolia and Salix eriocarpa (Salicaceae). To test this, we conducted field observations and a laboratory experiment. The field observations demonstrated that the leaf flushing phenology of the willows and the seasonal dynamics of the beetle density differed between shoots with stem galls and shoots without them. On galled shoots of both willow species, secondary shoot growth and secondary leaf production were promoted; consequently, leaf production showed a bimodal pattern and leaf production periods were 1 to 2 months longer than on non‐galled shoots. The adult beetle density on galled shoots was thus enhanced late in the season, and was found to change seasonally, synchronizing with the production of new leaves on the host willow species. From the results of our laboratory experiment, we attributed this synchrony between adult beetle density and willow leaf flush to beetles’ preference to eat new leaves rather than old. Indeed, beetles consumed five times more of the young leaves when they were fed both young and old leaves. These results indicate that stem galls indirectly enhance the adult beetle density by enhancing food quality and quantity late in the beetle‐feeding season. We therefore conclude that midge galls widen the phenological window for leaf beetles by extending the willows’ leaf flush periods.     

The effect of bark phenols upon mountain hare barking of winter-dormant Scots pine

Due to high numbers of mountain hares in recent winters in northern Finland the barking of winter dormant Scots pines was widespread. The hares fed selectively upon upper crown bark of grafted pines, nitrogen fertilized trees and trees in poor condition. Bark of preferred upper crown branches and physiologically mature scions contained less total phenols than lower parts. However, there were no differences between total phenols concentrations of graft bark preferred by hares and nearby unbarked juvenile phase Scots pine. Neither decrease in resistance after nitrogen fertilization n- or stress was correlated with bark total phenolic concentration. Thus, total phenolic concentration is not a reliable predictor of the susceptibility of Scots pine to winter barking by the mountain hare.     

Defense of winter-dormant Alaska paper birch against snowshoe hares

Summary Mature growth-phase internodes of Alaska paper birch (Betula resinifera) are preferred by the snowshoe hare (Lepus americanus) over juvenile growth-phase internodes due to the low food value of the latter. While the mature over juvenile preferencec cannot be explained by the levels of inorganic nutrients or gross chemical fractions (resins or phenols), it can be explained by the striking differences in secondary metabolites of the two growth phases. The principle compound which renders the juvenile phase internodes unpalatable is papyriferic acid, a triterpene which is a demonstrated feeding deterrent to snowshoe hares and which is present in juvenile internodes at concentrations 25 times greater than those in mature internodes.     

Chlorogenic acid as an antiherbivore defence of willows against leaf beetles

In the leaves of 13 Finnish willow species, the content of a phenolic, chlorogenic acid, was found to vary from 0 up to 18 mg g^–1 D.W. Effects of pure chlorogenic acid on insect feeding behaviour were tested using four common leaf beetle species which are in the field mainly found on willows with low-chlorogenic acid leaves. One species, Lochmaea capreae L., was invariably deterred by pure chlorogenic acid applied in naturally occurring concentrations on the willow leaves. Accordingly, in 2-choice laboratory feeding trials L. capreae was found to prefer low-chlorogenic acid leaves of four willow species over high-chlorogenic acid leaves of Salix pentandra L. and S. myrsinifolia Salisb. When presented on the leaves of S. phylicifolia L, pure chlorogenic acid inhibited also the feeding by Phratora polaris Sp.-Schn. Instead, chlorogenic acid had no significant effect on Ph. polaris when it was presented on the leaves of another willow S. cinerea L. In laboratory, Ph. polaris did not show general preference for willow species with low chlorogenic acid content in their leaves. Thus, the response of Ph. polaris to chlorogenic acid seems to depend on the plant species. Apparently variation in other traits such as leaf hairyness may easily override the potential effect of chlorogenic acid content on Ph. polaris. To two other leaf beetle species, Galerucella lineola F. and Plagiodera versicolora Laich., chlorogenic acid is an ineffective deterrent even at unnaturally high concentrations. In laboratory, G. lineola and P. versicolora did not prefer willows with low chlorogenic acid content in their leaves. Thus, among four studied leaf beetle species, only L. capreae seems to be clearly affected by this phenolic. Therefore, overall importance of chlorogenic acid as a defence against willow-feeding leaf beetles appears to be very limited.     

Beetle assemblages on willow trees: do phenolic glycosides matter?

Herbivorous beetles were sampled in central Slovakia and in the Tatra Mountains of northern Slovakia from seven different Salix species which are partly characterized by smooth leaves containing phenolic glycosides and partly by hairy leaves containing tannins. The collection included about 8000 individuals representing 129 species. Of the 129 species, 77 species are able to use the willows as hosts; the remaining 52 ‘tourist’ species comprise less than 3% of the collected individuals. The data on species richness and abundance did not generally support the ‘feeding specialization’ hypothesis. The proportion of specialized (= monophagous and oligophagous) beetles feeding on willows of both morphological and biochemical groups was in the same range. Host plants of the two groups could support high diversity and high evenness values, even when leaf characteristics and plant chemicals largely influenced species assemblage. The region in which the willow trees grow had a considerable impact on host plant use. Generalist beetle species predominated in central Slovakia. By contrast, in the Tatra Mountains, specialist feeders which are able to use phenolic glycosides to their advantage were predominant. The number of species and the total density of individuals collected from willows containing phenolic glycosides (S. fragilis and S. purpurea) did not usually vary between the two regions. In contrast, fewer species and individuals were found in the Tatra Mountains when they settled on willow species containing tannins (S. caprea and S. cinerea). Also, the phylogenetic status of host plants affected species assemblages. In central Slovakia willow species of the subgenus Vetrix (S. purpurea, S. caprea and S. cinerea) generally showed a higher beetle diversity (H_s) than species of the subgenus Salix (S. fragilis, S. alba and S. triandra), although both subgenera comprise species of both morphological and biochemical groups. Furthermore, when the analysis was restricted to beetles of central Slovakia, which should be most adapted to their host plants (i.e. catkin feeders and phyllophages in the adult and larval stage), the phylogenetic status was found to be more important than any single leaf character measured.     

Morphological and biomechanical responses of floodplain willows to tidal flooding and salinity

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Food competition between a large ruminant and a small hindgut fermentor: the case of the roe deer and mountain hare

In this study, we demonstrate that the mountain hare and roe deer compete with each other. This was determined using "natural experiments" of populations found in sympatry and allopatry on the islands along the west coast of Norway. We demonstrate that both species occupy the same habitats, share the same food resources and that food availability is limited. Two browsing species as different in size as roe deer and mountain hare might be expected to partition the available vegetation (e.g. woody scrub) in terms of height above ground level. However, from the evidence collected, the feeding-height-separation hypothesis must be rejected as an explanation for ecological separation between roe deer and mountain hares because there was extensive height overlap in resource utilisation by both species and neither species changed its feeding height in response to the presence of the other. Total browse utilisation did not increase when both species were together; rather, species-specific browse utilisation declined when the other species was present. However, the foraging behaviour of each herbivore varied significantly between the allopatric and sympatric sites. When both herbivores were present, the clip diameter of shoots browsed by mountain hares declined to match those selected by roe deer, while roe deer switched from a browse-dominated diet to a diet dominated by winter-green gramineae. The change to smaller-diameter shoots likely resulted in the hare increasing its intake of digestion-inhibiting or toxic secondary metabolites, while the alternative choice of digging through the snow like roe deer to reach the winter-green gramineae is a practice considered energetically too costly for hares. On this basis, we conclude that the enforced switch to a nutritionally inferior diet by mountain hares at the sympatric sites may result in changes to growth rate and body size which consequently impact on mortality and may explain the competitive superiority of the roe deer.     

Age-related changes in defensive traits of Acacia tortilis Hayne

The theory of plant defences proposes that investments in physical and chemical defences are driven by the risk of herbivore damage, and limited by the cost of producing the particular defensive trait in terms of resources that could be directed to other sinks, such as growth and reproduction. We sampled twigs of 18 mature Acacia tortilis trees and their cohort of juveniles to test some predictions of this hypothesis. We expected a higher allocation of defensive traits to leaves and twigs in the young plants than in the mature ones as a result of a higher risk of damage by ungulates at the juvenile stage. Our results show that the juvenile plants produce more spines along their twigs, but have lower concentrations of phenolic compounds in their leaves than in the mature ones. We also expected a negative relation between the concentration of foliar nutrients and phenolic compounds, as predicted by the carbon/nutrient hypothesis. Only mature plants showed this pattern. Reproduction (in mature plants) and water stress (in juvenile plants) did not relate to allocation to secondary compounds as predicted by current hypotheses of plant defence.     

Willow bud and shoot foraging by ptarmigan in relation to snow level in NW Finnish Lapland

Willow buds and twigs compose the main part of the willow ptarmigan’s (Lagopus lagopus) diet during winter. This study evaluated how position of willow ramets in relation to snow level affects the browsing of willow ptarmigan in NW Finland. Ramet position in relation to snow level affected the browsing intensity of willow ptarmigan: 5% of shoots were cut and more than 8% of buds were eaten by willow ptarmigan when ramet was staked at the maximum snow level, whereas ramets bent down were unbrowsed and browsing of ramets staked upright was negligible. Browsing significantly decreased the number of vegetative buds and catkins. The results show that snow level and ramet characteristics are important determinants of willow ptarmigan foraging.     

Host‐plant quality adaptively affects the diapause threshold: evidence from leaf beetles in willow plantations

1. Voltinism of herbivorous insects can vary depending on environmental conditions. The leaf beetle Phratora vulgatissima L. is univoltine in Sweden but will sometimes initiate a second generation in short‐rotation coppice (SRC) willow plantations. 2. The study investigated whether increased voltinism by P. vulgatissima in plantations can be explained by (i) rapid life‐cycle development allowing two generations, or (ii) postponed diapause induction on coppiced willows. 3. In the field, no difference was found in the phenology or development of first‐generation broods between plantations (S. viminalis) and natural willow habitats (S. cinerea). However, the induction of diapause occurred 1–2 weeks later in SRC willow plantations. 4. Laboratory experiments indicated no genetic difference in the critical day‐length for diapause induction between beetles originating from plantations and natural habitats. Development time was unaffected by host‐plant quality but critical day‐length was prolonged by almost an hour when the beetles were reared on a non‐preferred willow species (S. phylicifolia). When reared on new leaves from re‐sprouting shoots of recently coppiced willow plants, diapause incidence was significantly less than when the beetles were reared on mature leaves from uncoppiced plants. 5. The study suggests that P. vulgatissima has a plastic diapause threshold influenced by host‐plant quality. The use of host‐plant quality as a diapause‐inducing stimulus is likely to be adaptive in cases where food resources are unpredictable, such as when new host‐plant tissue is produced after a disturbance. SRC willows may allow two beetle generations due to longer growing seasons of coppiced plants that grow vigorously.     

Mortality factors and resource use of the bud-galling sawfly, Euura mucronata (Hartig), on willows (Salix spp.) in arctic Eurasia

We studied egg and larval mortality factors in arctic populations of the bud-galling sawfly, Euura mucronata (Hartig) on three willow species (Salix glauca L., S. phylicifolia L., and S. lapponum L.) and the quality of resources (shoot length of willows) required for egg-laying. The survival was independent of latitude. There was a positive correlation in survival on different willow species among sampling sites, indicating that similar, locally operating factors affected survival. Host plant-based mortality factors were dominant and caused 17.9–48.0% mortality in eggs and 6.6–44.1% mortality in larvae. Parasitoids and inquilines caused relatively low and variable rates of mortality. Parasitoids were absent from several of the northernmost populations, but caused up to 11.0% mortality at southern sites. Mortality caused by inquilines was minor in other areas except in some sites in the Taymyr Peninsula, where it varied from 0 to 23.1%. E. mucronata laid eggs on the longest shoots available. Plant vigour as measured by shoot length decreased toward the north, and densities of galls were positively correlated with plant vigour. The difference in length between galled and ungalled shoots was 2.0- to 4.6-fold. Absence of E. mucronata in the most northern populations of willows was apparently caused by insufficient shoot growth. Short shoots failed to provide sufficient resources for successful development of galls. Our results suggest that the quality of host plants is the main factor determining abundance and distribution of E. mucronata in arctic areas. Received: 10 February 1997 / Accepted: 2 May 1997     

The importance of willow thickets for ptarmigan and hares in shrub tundra: the more the better?

In patchy habitats, the relationship between animal abundance and cover of a preferred habitat may change with the availability of that habitat, resulting in a functional response in habitat use. Here, we investigate the relationship of two specialized herbivores, willow ptarmigan (Lagopus lagopus) and mountain hare (Lepus timidus), to willows (Salix spp.) in three regions of the shrub tundra zone-northern Norway, northern European Russia and western Siberia. Shrub tundra is a naturally patchy habitat where willow thickets represent a major structural element and are important for herbivores both as food and shelter. Habitat use was quantified using feces counts in a hierarchical spatial design and related to several measures of willow thicket configuration. We document a functional response in the use of willow thickets by ptarmigan, but not by hares. For hares, whose range extends into forested regions, occurrence increased overall with willow cover. The occurrence of willow ptarmigan showed a strong positive relationship to willow cover and a negative relationship to thicket fragmentation in the region with lowest willow cover at landscape scale, where willow growth may be limited by reindeer browsing. In regions with higher cover, in contrast, such relationships were not observed. Differences in predator communities among the regions may contribute to the observed pattern, enhancing the need for cover where willow thickets are scarce. Such region-specific relationships reflecting regional characteristics of the ecosystem highlight the importance of large-scale investigations to understand the relationships of habitat availability and use, which is a critical issue considering that habitat availability changes quickly with climate change and human impact.     

Plant sex and hare feeding preferences

Summary To evaluate the general extent to which sex-related differences in palatability occur in boreal dioecious woody plants, males and females of five dioecious woody plant species were presented to free-ranging mountain hares (Lepus timidus) during winter. Hares strongly preferred branches from male plants when feeding on Populus tremula and Salix caprea and weakly preferred male S. pentandra. However, they did not show any sex-related preference when feeding on the other two species studied (Myrica gale and Juniperus communis). Nitrogen concentration and, to some degree, digestibility showed strong relationships with hare food preferences. By contrast, the concentration of phenolics was only weakly related to feeding preference. Phenolics could, nevertheless, still be important if only one or a few specific compounds deter hare feeding. These results indicate that sex-related differences in plant palatability in the boreal forest might be more widespread than previously believed, particularly for species of the family Salicaceae. Thus, herbivores might be responsible for the female-biased sex ratios found in willow populations in northern Scandinavia (e.g. Elmqvist et al. 1988).     

Susceptibility of willow clones (Salix spp.) to herbivory by Phyllodecta vulgatissima (L.) and Galerucella lineola (Fab.) (Coleoptera, Chrysomelidae)

Levels of damage by mixed natural infestations of the leaf-feeding chrysomelid beetles, Phyllodecta vulgatissima (L.) (the blue willow beetle) and Galerucella lineola (Fab.) (the brown willow beetle), were determined in replicated field plots of 24 Salix clones at Long Ashton (Bristol, UK) during 1993–94. Over the same period, the host plant preferences of both chrysomelids were investigated in a standard multiple-choice laboratory procedure, where beetles were enclosed in Petri dishes with leaf discs cut from young pot-grown trees propagated from shoot cuttings taken from 20 of the 24 willow clones represented in the field study. The laboratory experiments indicated that P. vulgatissima and G. lineola had similar host plant preferences in the range of willows examined (r >0.85). In both field and laboratory, the least preferred Salix clones and hybrids were those of 5. eriocephala, followed by S. purpurea, S. burjatica, S. dasyclados and S. triandra. Clones of S. eriocephala and S. purpurea were frequently rejected altogether in laboratory tests. Most preferred were clones of S. viminalis and several hybrids of S. viminalis, S. aurita, S. caprea and S. cinerea. These results substantiate the reports that P. vulgatissima and G. lineola are deterred from feeding on willows which have relatively high concentrations of phenolic (salicylate) glucosides in the leaves. The least preferred willows, particularly S. eriocephala, S. purpurea and S. burjatica, could be of great potential value in plant breeding for resistance to these willow beetle pests.