Optimal central-place foraging by beavers: Tree-size selection in relation to defensive chemicals of quaking aspen

Summary At a newly occupied pond, beavers preferentially felled aspen smaller than 7.5 cm in diameter and selected against larger size classes. After one year of cutting, 10% of the aspen had been cut and 14% of the living aspen exhibited the juvenile growth form. A phenolic compound which may act as a deterrent to beavers was found in low concentrations in aspen bark, and there was no significant regression of relative concentration of this compound on tree diameter. At a pond which had been intermittently occupied by beavers for over 20 years, beavers selected against aspen smaller than 4.5 cm in diameter, and selected in favor of aspen larger than 19.5 cm in diameter. After more than 28 years of cutting at this site, 51% of the aspen had been cut and 49% of the living aspen were juvenileform. The phenolic compound was found in significantly higher concentrations in aspen bark than at the newly occupied site, and there was a significant negative regression of relative concentration on tree diameter. The results of this study show that responses to browsing by trees place constraints on the predictive value of standard energy-based optimal foraging models, and limitations on the use of such models. Future models should attempt to account for inducible responses of plants to damage and increases in concentrations of secondary metabolites through time.     

Seasonal and year-to-year differences in food selection by beavers

Summary Selection of trees for food by a colony of beavers in central Massachusetts was studied from September 1972 through April 1974. The beavers exhibited both seasonal and year-to-year differences in preference for certain genera. Pine was selected against during fall but not spring, and there was a switch in preference from birch during fall 1972 to oak and witch hazel in fall 1973. These differences may partly reflect greater seasonal stability in concentrations of stored nutrients in coniferous tree bark than in deciduous tree bark, and greater year-to-year stability in bark concentrations of stored nutrients in non-mast-seeding species than in mast-seeding species (1972 was a mast year for oaks in central Massachusetts).     

Tree Regeneration and Species Diversity Following Conventional and Uniform Spacing Methods of Selective Cutting in a Subtropical Humid Forest Reserve1

There are a variety of ways to diminish the negative impacts of forest management for timber on biodiversity. A pilot project using a uniform spacing method of selective cutting was recently implemented in the 5340 ha Guarani Reserve, Misiones, Argentina, to design adequate management schemes for the subtropical forests of the region. Uniform spacing involves moderate timber harvesting and careful selection of remnant trees. In this article we compare tree regeneration in forests experimentally cut by diameter limit and uniform spacing methods. Seedlings of five size classes from 10 cm to 3 m in height were sampled using rectangular nested plots. Three years after cutting, an average of 54,330 and 22,270 seedling/ha (all height classes combined) of commercial and non-commercial species were found in the forest cut by uniform spacing and by minimum diameter, respectively. In an adjacent uncut forest there were 34,900 seedlings/ha, and in another forest cut by minimum diameters 30 years ago there were 50,000 seedlings/ha, The forest cut by uniform spacing had the highest number of commercial seedlings/ha, with three times as many as the forest cut by minimum diameter three years ago and twice as many as the forest treated by minimum diameters 30 years ago. The forest cut by uniform spacing and the forest treated by minimum diameters 30 years ago had the highest diversity of understory plants other than trees, as well as heterogeneous canopy cover, while the other two forests had a predominance of bamboo in the understory and more open canopy conditions. Although lack of site replication limits interpretation of the results, these early findings suggest that the uniform spacing method can be an ecologically sound forest management option for the region.     

Foraging decisions of North American beaver (<Emphasis Type="Italic">Castor canadensis</Emphasis>) are shaped by energy constraints and predation risk

Foraging herbivores have to trade-off between energy requirements and predator avoidance. We aimed to study the relative roles of these factors in beavers (Castor canadensis) when foraging on land. We hypothesized that beavers were able to assess the risk of predation by using two main cues: the distance from the water and the presence or absence of predator odors. First, we studied the food selection of beavers in relation to distance from the water in natural settings. The transects were made at beaver ponds, and the diameter, species, and distance from the shore of intact and beaver-cut trees were recorded. Secondly, we placed rows of aspen sticks (Populus tremula) perpendicular to the shore around beaver ponds, and treated each row with a neutral, alien, or wolf odor. We found that aspen, downy birch (Betula pubescens), and speckled alder (Alnus incana) were the preferred tree species. More of these species were cut close to the shore, and cut trees were smaller further away from the shore, except in the case of aspen. In the experiment, most of the aspen sticks were taken close to the shore. As predicted, beavers took less aspen sticks in rows treated with wolf odor than water. As the predator odor did not affect the foraging distance from the shore, it is likely that our observation that foraging was the most intense close to shore is due to energetic constraints. However, predation risk probably affects the decision whether to forage on the land in the first place.     

The size at reproduction of canopy tree species in central Africa

Size at reproduction is a key aspect of species life history that is relatively understudied for long‐lived tropical trees. Here, we quantified reproductive diameter for 31 major timber species across 11 sites in Cameroon, Congo, and Central African Republic. Specifically, we examined whether (1) between‐species variability is correlated with other species traits; (2) reproductive diameter varies within‐species among sites; (3) reproductive status varies with crown exposure; and (4) the minimum cutting diameter limits (MCDL) imposed by national forest regulations enable seed trees to persist after logging operations. Consistent with studies conducted elsewhere in the tropics, we found great variability in diameter at reproduction among species, which correlated with adult stature (maximum diameter and height). For some species, reproductive diameter thresholds substantially varied between sites, and crown exposure had a significant positive effect on reproductive status. Most MCDLs were found to be suitable, with trees having a high probability of being seed trees at MCDL. Our findings have implications for the sustainable management of production forests, and they highlight questionable MCDLs for some species and between‐site variation in reproductive diameter. The study also highlights the need for long‐term phenological monitoring of tree species spanning a large range of ecological strategies to address both theoretical (species life history, allocation trade‐offs) and practical questions (MCDL).     

Trunk cutting initiates bottom-up cascades in a tri-trophic system: sprouting increases biodiversity of herbivorous and predaceous arthropods on willows

We examined the effects of trunk cutting on the regrowth responses of two dominant willow species, Salix eriocarpa and S. gilgiana , and the subsequent effects on the community structure (abundance and species richness) of herbivorous and predaceous arthropods. We studied ten randomly selected pairs of cut and uncut (control) trees of each willow species. Field observations showed that when the trunks were cut, shoots sprouted from the base and developed rapidly. These shoots continued to grow until July, producing lateral shoots and leaves, whereas current-year shoots on uncut trees essentially stopped growing by April. In July, the upper leaves of cut trees were less tough and had a greater water and nitrogen content than leaves of uncut trees. Leaf consumption and abundance and species richness of both herbivorous and predaceous arthropods were significantly greater on cut trees than on uncut trees in both willow species. Overall, trunk cutting resulted in at least a two-fold increase in both relative abundance and species richness within the arthropod community on the willow species. We concluded that the severe physical damage caused by trunk cutting greatly increased the biodiversity of herbivorous and predaceous arthropods through bottom-up cascading effects.     

Selective foraging on woody plant species by the Eurasian beaver (Castor fiber) in Telemark, Norway

Beavers Castor spp. are generalist herbivores, feeding on the bark, shoots and leaves of woody plants, terrestrial herbs and forbs, ferns and aquatic vegetation. As central-place foragers, beavers move out from water to select and cut trees and vegetation, and then transport it back to their refuge. These terrestrial forays are energetically costly; therefore, beavers should concentrate their foraging activity near the central place and increase the degree of selectivity for specific sizes or species of food with increasing distance from the water. The aim of this study was to test the predictions of the central place and the optimal foraging theories for the food selection of the Eurasian beaver Castor fiber , and show the foraging preferences of the focal species in the boreal conifer forest zone of Europe. Foraging intensity by beavers and the abundance of woody species were surveyed in transects positioned randomly at seven beaver territories. In compliance with the central-place foraging theory, the foraging intensity declined with increasing distance from the river. Beavers fed preferentially on willows ( Salix ), rowan ( Sorbus ) and birches ( Betula ), although alders ( Alnus ) dominated their diet. Size selectivity showed similar patterns to previous North American studies, which were also carried out in habitats with predominantly small saplings. The probability of selection of small saplings dropped as distance increased, which is consistent with the predictions of optimal foraging models that larger prey items are more likely to be favoured with increasing provisioning distance.     

Predicting tree diameter and height from above-ground biomass for four eucalypt species

Summary Previous work has suggested that tree stems are structured dimensionally to resist the forces to which they are subjected by the weight of the crown and the action of wind, snow and other loads on the crown. This proposition has been used to develop allometric relationships relating diameter at breast height or height of individual trees growing in even-aged monoculture to their above-ground fresh biomass. These models have practical application as estimators of tree diameters or heights from tree biomass as extensions of mechanistically based models of forest tree growth which predict tree biomasses. The present work applied these models to Eucalyptus regnans F. Muell, E. delegatensis R. Baker, E. nitens (Deane: Maiden) Maiden and E. grandis Hill ex Maiden trees, growing in plantation or regrowth stands, aged between 1.5 and 20 years, at eight geographically diverse sites extending from temperate to sub-tropical regions of Australia. While the models held for the various species at the various sites, their parameter values differed significantly between sites and/or species. This suggested there may be some inadequacy in the models. However, the differences were small and it was found reasonable to fit single models across all species and sites for practical use in estimating diameter or height. The errors about predicted values of height and diameter from these models were quantified. The models were also found to estimate diameter or height with little loss of precision when dry biomass was used in place of fresh biomass.     

Look before leaping: foraging selectivity of capuchin monkeys on acacia trees in Costa Rica

Acacia trees in Costa Rica have an obligate mutualism with three species of Pseudomyrmex ants, which vigorously defend their host tree from insect and mammalian herbivores. Depending on the size and species of ant colony, individual acacia trees may be differentially protected. For animals able to discern between weakly and highly aggressive ant colonies, costs of ant stings from less active colonies might be offset by nutritional value acquired from feeding on acacia fruit or ant larvae in swollen thorns. We examined foraging selectivity of capuchin monkeys on acacia trees in Santa Rosa National Park, Costa Rica. We measured four characteristics of the acacia trees from which capuchins fed and of acacias immediately adjacent to those in which the monkeys fed: diameter at breast height (DBH), accessibility, species of closest tree and ant species present. We found that capuchins prefer to forage in acacias that are large and accessible. We also made two measurements of ant colony activity on each tree, one before and one after disturbing the ant colony. We found that the three species of mutualistic ants differ in baseline activity levels and that mutualistic ants are more active than non-mutualistic ant species found in acacia trees. We also found that capuchins foraged more frequently in trees colonized by non-mutualistic ants, but the explanatory value (r ²) of this model was low. Furthermore, monkeys did not discriminate between acacias on the basis of baseline ant activity or the ant colony’s response to disturbance. We conclude that these monkeys select acacia trees in which to forage based on characteristics of the trees rather than the ants. In addition, our study suggests that white-faced capuchins act as predators on the acacia ants but they probably benefit the dispersal and reproductive success of acacia trees. Capuchins may in fact function as an additional mutualistic partner for acacia trees via seed dispersal, but they must overcome the ants’ defense of the trees to do so.     

Load-size determination in the leaf-cutting ant, Atta cephalotes

I examined load-size determination by a highly polymorphic leaf-cuttingant, Atta cephalotes, cutting leaves of artificial trees (branchesplaced in the top of plastic tubes). I compared load size forants cutting thin leaves (starfruit, Averrhoa carambola) andthick leaves (grapefruit, Citrus parodist). At each source,larger ants cut larger fragments. Distance from the nest hadno effect on load size. The mass of fragments cut by an antof a given size was significantly greater when cutting grapefruitleaves. The leaf area cut, however, showed no significant differencebetween the two leaf types. Leaf area increased approximatelyin proportion to ant body mass to the 0.6 power. As a resultof their method of load-size determination, ants of a givensize cut heavier loads when cutting the thicker leaves. Thisdifference, however, was counteracted at the colony level byrecruitment of larger ants, which cut smaller area fragmentsrelative to their body mass, to cut at thicker leaf sources.     

Dynamics of food resources for beavers in settlements colonized and abandoned several times

The state of food resources for Eurasian beavers (Castor fiber Linnaeus, 1758) after several cycles of habitat colonization and abandonment has been analyzed in three beaver settlements on small rivers of Novgorod oblast. It has been shown that tree stands surrounding the settlements differ in composition and degree of damage but are similar in that they contain no forage for beavers: aspen is absent, and birch trees are large and require great efforts to cut them down and utilize. The undergrowth and understory also contain no aspen, while abundant willows and birch are represented by young, thin-stemmed plants unevenly distributed over the area. Tree species prevailing in abandoned settlements are not favored by beavers, implying that the animals must have expended extra energy for foraging. Supposedly, populations of beavers on small rivers in the southern taiga subzone, in habitats modified by their activities, will maintain their abundance far below the maximum possible level, depending on the rate of forage plant regrowth against the background of deteriorating habitat quality.     

Colonization, riparian habitat selection and home range size in a reintroduced population of European beavers in the Loire

1. Colonization in a reintroduced population of European beavers in the Loire Valley was studied between 1974 and 1999. It followed a discontinuous remoteness model and a scattered distribution, beavers occupying only 25% of the river system over the 2800 km explored.
2. After 5 years, the colonization rate reached 104.2% year^–1 of new sites occupied (SD 75% year^–1), before dropping over the next 20 years. Nevertheless, the number of new colonies per km (0.125) remained stable throughout the years.
3.  Populus nigra , Salix alba and Fraxinus angustifolia were the dominant woody species in beaver sites, often associated with some herbaceous species.
4. The length of willow grove dominated by S. alba and P. nigra ( x ) was the best predictor of beaver home range ( y ), fitting the equation y =−0.742 x  + 5.9. Long-term maintenance of the population requires a minimum of 1.79-km of willows per colony.
5. In cutting tree trunks, beavers stimulate shoot development from the remaining stumps. They rejuvenate riparian forests, increase the number of tree stems and help stabilize the banks. Their effect on woody plant morphogenesis may have consequences for the helophyte communities used as food or habitat by other aquatic species.     

Tree allometry of Douglas fir and Norway spruce on a nutrient-poor and a nutrient-rich site

Research related to the allometric relationships of tree height and projected tree crown area to diameter at breast height was conducted to look at the biological suitability and timber production potential of Douglas fir under the conditions present in central Europe. The dependence of allometric relationships on soil nutrient conditions were described in forest stands of Douglas fir and Norway spruce. The studied sites were climatically similar but differed in soil nutrient availability. A significant difference was found in the allometric relationships of Norway spruce trees from the nutrient poor and nutrient rich site. In contrast to the Norway spruce, there was no significant effect of site fertility on allometric relationships for Douglas fir suggesting that its allocation patterns were less sensitive to site nutrient conditions. Stem growth increment, which was measured weekly during two consecutive seasons for both species, was related to the weather conditions and available soil moisture. Stem growth of Douglas fir began about 2 weeks earlier than in the Norway spruce at both sites. At the nutrient rich site, most of the stem growth of both species occurred at the beginning of the season, while growth at the other site was more evenly distributed throughout the season. Data obtained in this study will be useful for modeling stem growth and analysis of water use efficiency of these two tree species.     

Studies of Aquatic Fungi. XXVI. Hyphomycetes in the River Supraśi in Various Seasons of the Year with Reference to Environmental Conditions

The authors investigated the aquatic mycoflora and the environmental factors at two sites of the River Supraśl on the occurrence of various aquatic Hyphomycetes. At the stations investigated the presence of 56 aquatic Hyphomycetes species was noted. The foam of both site I and site II contained more species than did the water collected at these sites. Most of the aquatic Hyphomycetes found at the sites studied are common fungi quite frequently found in different kinds of water bodies. On the other hand 17 species have been comparatively seldom encountered up to date.     

An evaluation of hypotheses to explain the pattern of sap feeding by the yellow-bellied glider,Petaurus australis

Yellow-bellied gliders on occasion extensively incise the trunks of eucalypts to feed on sap but in doing so show strong preference for the species and number of trees utilized. Three hypotheses that may account for aspects of this behaviour were examined. They were: (i) sap feeding is related to the incidence of rain; (ii) sap feeding is influenced by the sap flow idiosyncracies of trees; and (iii) sap feeding is influenced by the availability of alternative food resources. Sap feeding at two sites occurred independent of ‘wet’ or ‘dry’ nights. Sap flow (measured as an index) in two species of eucalypt was highly variable among trees and among sample periods. This is the only hypothesis that accounts for the selectivity of the trees incised. Sap feeding at one site coincided with periods of high sap flow. However, at another site, sap feeding did not occur on measured trees despite their often high sap flow. At such times, flowering trees were abundant and gliders fed extensively on nectar. Thus, elevated sap flow may provide the necessary precondition conducive for sap feeding but may be ignored when other food types are available.     

Epiphytic Ferns for Sale: Influence of Commercial Plant Collection on the Frequency of Platycerium stemaria (Polypodiaceae) in Coconut Plantations on the Southeastern Ivory Coast

Abstract: On the southeastern Ivory Coast, two coconut plantations were compared with regard to the frequency of the epiphytic fern Platycerium stemaria on coconut trees as influenced by plant collecting for commercial purposes. At each of the two sites, 250 coconut trees were investigated for the presence of P. stemaria and signs of collecting activity. General species richness of vascular epiphytes at the study sites was low, with only three other epiphytic ferns ( Asplenium sp., Microgramma owariensis, Phymatodes scolopendria ) being present. At study site no. 1, which was located directly behind several flower stalls offering P. stemaria for sale, the number of coconuts bearing living P. stemaria was significantly lower than at site no. 2 with no flower stalls in the immediate surroundings. At site no. 1, the frequency of P. stemaria was lowest in the neighbourhood of the flower stalls and increased with distance. Within a distance of less than 50 m from the flower stalls, ca. 70 % of the coconuts showed signs of P. stemaria collecting activity. Compared with site no. 1, the number of palm trees with living P. stemaria at site no. 2 was considerably higher ( ca. 20 %) and only three coconut trees showed signs of Platycerium collection. Over much of the Ivorian rain forest region where the forest has been cut or converted into agricultural land, plantation trees are important phorophytes (in particular cocoa trees). In many West African regions plantation trees form the last remaining sites for vascular epiphytes, which often occur with considerable abundance. In the Ivory Coast, the collecting of P. stemaria in coconut plantations causes locally a drastic decline in the population size. However, much more threatening to the long-term maintenance of epiphyte communities in plantations is their conversion into non-tree plantations (e.g., pineapples, bananas) or their replacement by urban structures and tourist facilities.     

Juvenile sprouting ability of the endangered maple, <Emphasis Type="Italic">Acer pycnanthum</Emphasis>

Japanese red maple, Acer pycnanthum, is an endangered species having a limited distribution of central Honshu, Japan. Sprouting is an important part of its natural history, and many remnant populations contain high number of multiple-stem clones. However, knowledge of sprout formation and growth is lacking, which has hampered understanding of the reestablishment process after major disturbance. My research objectives were to characterize the formation and growth of basal sprouts in the juvenile stage and determine the effects of light intensity and stump diameter on these characteristics. Twenty sprouting clones were investigated at a natural site in Nagano Prefecture during a four-year period after cutting. Japanese red maple exhibited abundant and vigorous sprouting from the root collar of cut trees. Many sprouts arose immediately after cutting. Average number of sprouts per stump was 12.2. Number of sprouts per stump was negatively correlated with relative light intensity. The two dominant sprouts (sprouts 1 and 2) of each clone were selected to analyze their growth. The average extension growth of sprout 1 (i.e., the tallest and most vigorous) was 173.3 cm at the end of first growing season. At the end of fourth growing season, average height and diameter at breast height (dbh) of sprout 1 were 377 and 2.1 cm, respectively. Relative light intensity and stump diameter significantly and positively affected D²H (diameter at breast height squared × height) of sprout 1. Because the growth rate of sprouts is much faster than that of seedlings, regeneration by sprouting is more significant than seedling regeneration to maintain populations after disturbances.     

Physical site characteristics of wetland ecosystems dominated by the endangered maple, <Emphasis Type="Italic">Acer pycnanthum</Emphasis>

Japanese red maple, Acer pycnanthum, is an endangered species having a limited distribution of central Honshu, Japan. Sprouting is an important part of its natural history, and many remnant populations contain high number of multiple-stem clones. However, knowledge of sprout formation and growth is lacking, which has hampered understanding of the reestablishment process after major disturbance. My research objectives were to characterize the formation and growth of basal sprouts in the juvenile stage and determine the effects of light intensity and stump diameter on these characteristics. Twenty sprouting clones were investigated at a natural site in Nagano Prefecture during a four-year period after cutting. Japanese red maple exhibited abundant and vigorous sprouting from the root collar of cut trees. Many sprouts arose immediately after cutting. Average number of sprouts per stump was 12.2. Number of sprouts per stump was negatively correlated with relative light intensity. The two dominant sprouts (sprouts 1 and 2) of each clone were selected to analyze their growth. The average extension growth of sprout 1 (i.e., the tallest and most vigorous) was 173.3 cm at the end of first growing season. At the end of fourth growing season, average height and diameter at breast height (dbh) of sprout 1 were 377 and 2.1 cm, respectively. Relative light intensity and stump diameter significantly and positively affected D²H (diameter at breast height squared × height) of sprout 1. Because the growth rate of sprouts is much faster than that of seedlings, regeneration by sprouting is more significant than seedling regeneration to maintain populations after disturbances.     

Beaver lodge location on the upstream Loire River

In the part of the Loire River recently colonized by Eurasian beavers, we compared habitat characteristics among sites with lodges, sites with cut trees and sites without beaver. The absence of sandbank and canopy cover (by 10-15-m tall trees, by tall Salicaceae, and by bushy Salicaceae) appeared as good predictors for lodge settling. Based on this model, the number of proper lodge sites was estimated for the next downstream 36 kilometers stretch. The number of favourable sites decreases as anthropization increases.     

Unequal diameters and their effects on time-varying voltages in branched neurons.

A theoretical method, developed in a previous paper, enables one to calculate analytical expressions for time-varying voltages at specific locations in branching dendritic systems in response to synaptic current inputs at other sites. Exact results were obtained for a number of dendritic trees that possessed certain symmetries: all branch lengths had to be integral multiples of one another, and all branch diameters had to be equal. Because the second of these conditions is unrealistic, the method has been generalized to treat dendritic trees whose branches differ in diameter. The method entails adding onto the symmetric results a sum of correction terms. It is found that the correction terms, as well as the symmetric results, can be expressed as combinations of two families of functions. These functions, generalizations of those found in our earlier paper, provide a precise formalism for analyzing how voltage transients depend on the geometrical structure of the dendritic tree. Examples are given that show how the correction terms affect the value of the voltage, and how variations in branch diameters alter the behavior of the propagated postsynaptic potential. The implications of these results for our understanding of neuronal functioning are discussed.