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The tremble dance of honey bees can be caused by hive-external foraging experience.

The tremble dance of honey bee nectar foragers is part of the communication system that regulates a colony's foraging efficiency. A forager that returns to the hive with nectar, but then experiences a long unloading delay because she has difficulty finding a nectar receiver bee, will perform a tremble dance to recruit additional nectar receiver bees. A forager that experiences a short unloading delay will perform a waggle dance to recruit more nectar foragers. A long unloading delay was until now the only known cause of tremble dancing. However, several studies suggested that factors at the food source may also cause tremble dancing. Here I test whether one of these factors, crowding of nectar foragers at the food source, stimulates tremble dancing because it causes long unloading delays. To do so, I increased the density of nectar foragers at a food source by suddenly reducing the size of an artificial feeder, and recorded the unloading delay experienced by each forager, as well as the dance she performed, if any. A forager's unloading delay was measured as the time interval between entering the hive and either (1) the first unloading contact with a nectar receiver bee, or (2) the start of the first dance, if dancing began before the first unloading contact. I also recorded the unloading delays and dances of nectar foragers that returned from natural food sources. The results show that crowding of nectar foragers at the food source increases the probability of tremble dancing, but does not cause long unloading delays, and that tremble dancers that foraged at natural food sources also often have short unloading delays. When the cause of the tremble dance is not a low supply of nectar receiver bees, the tremble dance may have a function in addition to the recruitment of nectar receiver bees.

Animal Communication↗

Influence of milk production potential on forage dry matter intake by multiparous and primiparous Brangus females.

Brangus cows (n = 29) were used in three experiments to evaluate the effects of parity (multiparous vs. primiparous) and potential genetic merit for milk production (high vs. low) on forage intake during late gestation, early lactation, and late lactation. Cows were selected for milk production based on their sire's EPD for milk production (MEPD). Cows had ad libitum access to (130% of previous 2-d average intake) low-quality hay (5.3% CP and 76% NDF), and cottonseed meal was supplemented to ensure adequate degradable intake protein. All females were adapted to diets for at least 7 d, and individual intake data were collected for 9 d. During the lactation trials, actual milk production was determined using a portable milking machine following a 12-h separation from calves. During late gestation, multiparous cows consumed 24% more (P = 0.01) forage DM (kg/d) than primiparous cows; however, parity class did not influence forage intake when intake was expressed relative to BW. Furthermore, MEPD did not influence forage intake during late gestation. During early lactation, multiparous cows produced 66% more (P < 0.001) milk than primiparous cows, and high MEPD tended (P = 0.10) to produce more milk than low MEPD. Multiparous cows consumed 19% more (P < 0.0001) forage DM than did primiparous cows when expressed on an absolute basis, but not when expressed on a BW basis. High-MEPD cows consumed 8% more (P < 0.05) forage DM than did low-MEPD cows. During late lactation, multiparous cows produced 84% more milk than primiparous cows, although MEPD did not influence (P = 0.40) milk yield. In addition, multiparous cows consumed 17% more (P < 0.01) forage DM per day than primiparous cows, but when intake was expressed relative to BW, neither parity nor MEPD influenced forage DMI during late lactation. Milk yield and BW explained significant proportions of the variation in forage DMI during early and late lactation. Each kilogram increase in milk yield was associated with a 0.33- and 0.37-kg increase in forage DMI for early and late lactation, respectively. Results suggest that multiand primiparous cows consume similar amounts of low-quality forage DM, expressed per unit of BW, during late gestation and lactation. Selecting beef cows for increased genetic merit for milk production increases forage DMI during early lactation.

Animal Feed↗

Foraging enrichment for stabled horses: effects on behaviour and selection.

The restricted access to pasture experienced by many competition horses has been linked to the exhibition of stereotypic and redirected behaviour patterns. It has been suggested that racehorses provided with more than one source of forage are less likely to perform these patterns; however, the reasons for this are currently unclear. To investigate this in 4 replicated trials, up to 12 horses were introduced into each of 2 identical stables containing a single forage, or 6 forages for 5 min. To detect novelty effects, in the first and third trials the single forage was hay. In the second and fourth, it was the preferred forage from the preceding trial. Trials were videotaped and 12 mutually exclusive behaviour patterns compared. When hay was presented as the single forage (Trials 1 and 3), all recorded behaviour patterns were significantly different between stables; e.g. during Trial 3 in the 'Single' stable, horses looked over the stable door more frequently (P<0.001), moved for longer (P<0.001), foraged on straw bedding longer (P<0.001), and exhibited behaviour indicative of motivation to search for alternative resources (P<0.001) more frequently. When a previously preferred forage was presented as the single forage (Trials 2 and 4) behaviour was also significantly different between stables, e.g in Trial 4 horses looked out over the stable door more frequently (P<0.005) and foraged for longer in their straw bedding (P<0.005). Further study is required to determine whether these effects persist over longer periods. However, these trials indicate that enrichment of the stable environment through provision of multiple forages may have welfare benefits for horses, in reducing straw consumption and facilitating the expression of highly motivated foraging behaviour.

Animal Feed↗

Effects of forage neutral detergent fiber and yeast culture on performance of cows during early lactation.

Sixty Holstein cows were assigned to two treatments at 21 d before calving and were group-fed a prepartum diet with or without yeast culture. After parturition, cows were individually fed one of five treatments for 140 d: 1) 21% forage neutral detergent fiber (NDF) without yeast culture, 2) 21% forage NDF with yeast culture, 3) 17% forage NDF without yeast culture, 4) 17% forage NDF with yeast culture, and 5) 25% forage NDF with yeast culture for 30 d and then switched to diet 4 for 110 d. Cows fed yeast culture prepartum were also fed yeast culture postpartum (60 g/d). A quadratic increase to 25, 21, and 17% forage NDF occurred during the first 30 d in milk (DIM) for dry matter intake, milk yield, and milk protein yield. No differences were observed for yeast culture or interaction of yeast culture and forage NDF for the first 30 DIM. Feeding 17 versus 21% forage NDF increased milk protein percentage and tended to increase dry matter intake as a percentage of body weight from 31 to 140 DIM. During this period, yeast culture tended to increase milk fat percentage and appeared to have positive effects on dry matter intake, milk yield, and milk fat yield when supplemented to diets with 21% forage NDF but not with 17% forage NDF. Feeding 17% forage NDF may be too low for the first 30 DIM but may improve animal performance after 30 DIM compared to 21% forage NDF.

Animals↗

Dissecting the genetics of forage quality traits in soft red winter wheat in the U.S. southeast region.

Winter wheat plays a viable role in agriculture, not only as a primary grain crop but also as a valuable forage source that bridges fall-spring forage gaps in many regions, including the southeastern (SE) U.S. Despite its nutritive potential, genetic basis of forage-quality traits remains insufficiently understood, limiting breeding efforts for dual-purpose cultivars. This study aimed to dissect the genetic architecture of forage quality in 182 soft red winter wheat (SRWW) genotypes adapted to the SE U.S. using genome-wide association study (GWAS). Field experiments were carried out in randomized complete block design across three Georgia locations over two growing seasons (2023-2025), with forage sampled at the end of tillering and evaluated using near-infrared reflectance spectroscopy. Significant phenotypic variation was observed for dry matter (DM), crude protein (CP), acid and neutral detergent fiber (ADF, NDF), acid detergent lignin (ADL), total digestible nutrients (TDN), sugars (SUG), and relative forage quality (RFQ). Heritability estimates ranged from low-to-moderate in combined environments and from low-to-high within individual locations. Correlation analysis revealed strong positive associations among fiber-related traits and negative associations with TDN, RFQ, and SUG, while CP declined with increasing fiber. Genome-wide association analysis identified 282 significant marker-trait associations (P&#x2009;<&#x2009;1&#xd7;10-4) across 19 chromosomes, which were consolidated into 121 QTLs, including 27 major-effect QTLs. Three QTLs QRfq.uga-3B.1, QRfq.uga-3B.2 (RFQ) and QDm/Sug.uga-7A (DM, SUG) were stable across locations while QAdf/Adl.uga-2A (ADF, ADL) and QDm/Sug.uga-7A (DM, SUG) indicated multi-trait control. Notably, 25 of the 27 major QTLs were putatively novel, highlighting substantial untapped allelic diversity for forage-quality improvement in SE SRWW. Favorable allele accumulation resulted in an overall improvement in forage quality, increasing desirable nutritive traits (DM, RFQ, SUG, CP) while reducing undesirable traits (ADF, ADL). Candidate gene analysis linked six major QTLs with genes implicated in abiotic stress response, plant development, and metabolic regulation, supporting their functional relevance in forage-quality determination. Incorporating these loci into breeding programs provides a robust genetic framework for marker-assisted selection, enabling the development of dual-purpose wheat cultivars with enhanced forage quality, thereby strengthening wheat's utility as a reliable forage resource during periods of seasonal feed scarcity in SE production systems.

GWAS↗

Neandertal energetics and foraging efficiency.

Mechanical interpretations of Neandertal skeletal robusticity suggest extremely high activity levels compared to modern humans. Such activity patterns imply high energy requirements; yet it has been argued that Neandertals were also inefficient foragers. The present study addresses this apparent conflict by estimating energy needs in Neandertals and then evaluating those estimates in the context of energetic and foraging data compiled for contemporary human foragers and nonhuman primates. Energy demands for Neandertals were determined by first predicting basal metabolic rates (BMR) from body weight estimates using human standards developed by the World Health Organization [FAO/WHO/UNU (1985) Energy and Protein Requirements. Report of the Joint FAO/WHO/UNU Export Committee, Geneva: WHO]. Total daily energy expenditure (kcal/day) was then estimated assuming high levels of physical activity (i.e., 2--3 x BMR), comparable to those observed among subsistence-level populations today. These estimates of energy requirements (ranging from 3000--5500 kcal/day) were then used to determine Neandertal foraging efficiency assuming (1) minimal survival-level foraging returns, and (2) daily foraging times longer than those observed among any contemporary foraging group and comparable to a nonhuman primate. Even with these extremely conservative parameters, estimates of Neandertal foraging efficiency (approximately 800--1150 kcal/h foraged) were comparable to those observed among living hunter-gatherers. These results indicate that if Neandertals did have heavy activity levels, as implied by their skeletal robusticity, they would have required foraging efficiencies within the range observed among modern groups. Thus, Neandertals could have been either highly active or poor foragers, but they could not have been both.

Adult↗

The effect of genotype, age, sex, and caste on response thresholds to sucrose and foraging behavior of honey bees (Apis mellifera L.).

Bees derived from artificially selected high- and low-pollen-hoarding strains were tested for their proboscis extension reflex response to water and varying sucrose concentrations. High-strain bees had a lower response threshold to sucrose than low-strain bees among pre-foragers, foragers, queens and drones. Pre-foraging low-strain workers showed ontogenetic changes in their response threshold to sucrose which was inversely related to age. High-strain foragers were more likely to return with loads of water compared to low-strain foragers. Whereas low-strain foragers were more likely to return with loads of nectar. Low-strain nectar foragers collected nectar with significantly higher sucrose concentrations than did the high-strain nectar foragers. Alternatively, low-strain foragers were more likely to return empty compared to high-strain foragers. These studies demonstrate how a genotypically varied sensory-physiological process, the perception of sucrose, are associated with a division of labor for foraging.

Aging↗

Foraging behavior and physiological changes in precocial quail chicks in response to low temperatures.

We examined whether low ambient temperatures influence foraging behavior of precocial Japanese quail chicks and alter the balance between investment in growth and thermogenic function. To test this, one group of chicks was exposed to 7 degrees C and one group to 24 degrees C during foraging throughout the developmental stage. Chicks adapted well to the temperatures through a high flexibility in behavior and physiological development. In response to cold, chicks shortened foraging bout lengths two- to sixfold, and increased cycle lengths (i.e. length of a brooding bout plus subsequent foraging bout), resulting in a decrease in total foraging time. Body temperature during foraging was lower in cold-exposed chicks but did not drop below 37.8 degrees C, suggesting that the direct cause to end a foraging bout was not body temperature. The metabolic rate of cold-exposed chicks was reduced by 48% to 60% when switching from foraging to brooding, which may be an important factor in determining foraging behavior of precocial chicks. Mass-specific food intake of the cold-exposed chicks exceeded that of warm chicks by 15%. Cold-exposed chicks doubled foraging efficiency to reach this intake during their reduced foraging time. Metabolic rates initially were similar in both groups, but increased rapidly and were elevated in cold-exposed chicks from 15 days of age onward. Growth rate initially was reduced in cold-exposed chicks, and exceeded growth of warm chicks only after 21 days of age. These results suggest that in response to cold, a shift occurs in the balance between growth rate and thermoregulatory function in favor of thermoregulatory function.

Animals↗

Lambs fed protein or energy imbalanced diets forage in locations and on foods that rectify imbalances.

Ruminants eat a variety of foods from different locations in the environment. While water, cover, social interactions, and predators are all likely to influence choice of foraging location, differences in macronutrient content among forages may also cause ruminants to forage in different locations even during a meal. We hypothesized that lambs forage at locations containing foods that complement their basal diet and meet their nutritional needs. Based on this hypothesis, we predicted that lambs (n=12) fed a basal diet low in protein and high in energy would forage where a high-protein food (Food P) was located, and that lambs (n=12) fed a basal diet low in energy and high in protein would forage where a high-energy food (Food E) was located. Food P was a ground mixture of blood meal (50%), grape pomace (30%), and alfalfa (20%) that contained 47% crude protein (CP) and 2.211 Mcal/kg digestible energy (DE). Food E was a ground mixture of cornstarch (50%), grape pomace (30%), and rolled barley (20%) that contained 6% CP and 3.07 Mcal/kg DE. Food P provided 212 g CP/Mcal DE, whereas Food E provided 20 g CP/Mcal DE. Lambs growing at a moderate rate require 179 g CP and 3.95 Mcal DE. During Trial 1, we determined if lambs foraged to correct a nutrient imbalance, and if they preferred a variety of foods (Foods P and E) to only one food at a location (Food P or E). During Trial 2, we determined if nutrient-imbalanced lambs foraged in the location with the food that corrected the imbalance when the location of the foods changed daily. During Trial 3, lambs were offered familiar foods (Foods P and E) at the location furthest - and novel foods (wheat and soybean meal) at the location nearest - the shelter of their pen. During all three trials, lambs foraged most at the location with the food that contained the highest concentration of the macronutrient lacking in their basal diet, but they always ate some of both foods. Lambs did not feed exclusively at the location with a variety of foods (P and E). Rather, they fed at the location nearest the shelter that contained the macronutrient lacking in their diet. As availability of the food with the needed macronutrient declined in one location, lambs moved to the nearest location that had food with the needed macronutrient. When food that complemented their basal diet was moved to a different location, lambs foraged in the new location. Collectively, these results show that lambs challenged by imbalances in energy or protein selected foods and foraging locations that complemented the nutrient content of their macronutrient imbalanced basal diets.

Journal Article↗

Foraging methods can affect patch choice: an experimental study in Mallard (Anas platyrhynchos).

Animals can adapt to changes in feeding conditions by switching between foraging methods. Dabbling ducks use different foraging methods, including dabbling in deep water with the head and neck submerged, and grubbing in the mud (or shallow water) where the eyes are above the surface, so the bird can visually monitor its environment while foraging. Deep foraging is considered to provide lower intake rates and to have high associated costs, such as predation risk, compared to shallow foraging. Ducks should thus prefer shallow foraging and switch to deeper methods when feeding conditions deteriorate. We conducted a set of experiments with Mallard to assess the importance of intake rate as a cue to choose between patches associated with different foraging methods, and evaluate the influence of food depletion on the decision to switch between methods. When 50 g of wheat were presented in two patches, one at a depth of 5 cm and one at 35 cm, most of the foraging was in the shallow area. Reducing food abundance to 10 g in the shallow area led to an increase in deep foraging, although the birds still preferred the shallow area at the beginning of the tests despite the fact that it did not provide a higher intake rate. This area was used until complete depletion, and birds did not turn to deep foraging before ensuring that the shallow patch was empty. These results show that food depletion affects the choice between feeding patches hence foraging method. However the value of intake rate is not the main cue for decision, rather the birds appear to choose between patches with different methods on account of their respective costs.

Journal Article↗

Influence of moisture content of forage diets on intake and digestion by sheep.

To determine the effect of forage moisture content on intake and digestion kinetics in sheep, a metabolism trial was conducted using 16 mature wethers (44 kg BW) in a completely randomized design. Forage was harvested at two maturities in early spring from a naturalized pasture composed of temperate grass and legume species. Herbage was harvested at either 8 (early) or 16 cm (late) in height and fed after freezing (high-moisture) or as a dried hay. Intakes of DM (grams/[kilogram BW.75.day]), NDF, ADF, and CP (grams/day) (P < .05) and coefficients for DM, NDF, and ADF digestibilities were greater (P < .01) for hay than for high-moisture forage. Mean particulate retention times (MRT) were shorter (P < .05) for high-moisture forage (23.3 h) than for hay (30.7 h) diets. Early-harvested forages had shorter (P < .05) MRT values (23.9 h) than late-cut forages (30.1 h). Fractional passage rates of 1-mm nylon particles of specific gravity (SG) .90, 1.14, and 1.32 through the alimentary tract were influenced by moisture content of the forage (P < .10) and were faster for frozen forages and increased (P < .01) with an increase in SG. Nitrogen retention was greater (P < .01) for the hay than for the high-moisture forage. In situ DM digestion rates, determined using four ruminally fistulated wethers, showed no differences (P = .67) among forages. The results of this study indicate that differences in digestibilities between hays and high-moisture forages are most likely due to differences in digesta passage rates.

Animal Feed↗

Effect of particle size upon digestible cell wall and rate of in vitro digestion of alfalfa and orchard grass forages.

Alfalfa and orchardgrass forages chopped through a 13-mm screen were larger in particle size than were rumen ingesta samples from sheep fed the same forages. Samples of the chopped forages were ground through 1-, 4-, 8-, and 12-mm screens; particle sizes of the 8- and 12-mm ground forages were not significantly different from those of rumen ingesta samples or chopped forages. The 1- and 4-mm ground forages were significantly smaller in particle size than the 8- and 12-mm ground forages, rumen ingesta and chopped forage. Alfalfa contained less cell wall at the beginning of in vivo digestion, more cell wall residue at the end of digestion and less potentially digestible cell wall than did orchardgrass. The cell wall of alfalfa was 44% digestible, whereas that of orchardgrass was 61% digestible; particle size did not affect the amount of digestible fiber. Alfalfa ground through 1- and 4-mm screen openings had digestion rate constants that were about twice those of the 8- and 12-mm granulations (.10 less than P less than .20). Particle size did not affect rate constants of orchardgrass. These data suggest that in vitro kinetic studies based on examination of finely ground forages (1) may not reflect in vivo conditions, and (2) may depend upon the type of forage. Differences between legumes and grasses in (1) amount of digestible fiber and (2) digestion rate constants and the interaction of the above two factors with particle size and passage rate have important implications for digestibility of forage fiber.

Animal Feed↗

Effects of supplemental silage on forage intake and utilization by steers grazing wheat pasture or bermudagrass.

Seven trials were conducted to determine the effects of increasing amounts of silage (corn, wheat or sorghum) on forage intake, gastrointestinal tract fill, fecal output and ruminal flow and degradation of forage by cattle grazing wheat pasture or bermudagrass. In each of 3 yr, 24 steers grazed a common wheat or bermudagrass pasture and were randomly allocated to four treatments (0, .35, .70 or 1.05 kg silage DM.100 kg body weight-1.d-1). Intake and ruminal flow of forage were measured by feeding a single pulse dose of Yb-labeled forage followed by collection of fecal samples for 4 to 5 d and fitting Yb concentrations to a one-compartment, age-dependent model. Ruminal digestion kinetics of wheat forage were estimated in situ using eight ruminally cannulated steers fed 0 or .55 kg sorghum silage DM.100 kg body weight-1.d-1. Supplemental silage decreased wheat forage (P less than .10) and bermudagrass (P less than .01) intake linearly. However, total forage intake of bermudagrass plus silage increased linearly (P less than .05). Each kilogram of added silage DM decreased DM intake of wheat forage by .66 +/- .25 and of bermudagrass by .63 +/- .17 kg. Flow and turnover of wheat forage or bermudagrass were not altered (P greater than .15) by supplemental silage. Silage consumption increased extent of ruminal degradation of wheat forage DM (P less than .05; 63.1 vs 52.5%), indicating a positive associative effect of silage on wheat forage utilization.(ABSTRACT TRUNCATED AT 250 WORDS)

Animal Feed↗

Partial replacement of forage with nonforage fiber sources in lactating cow diets. II. Digestion and rumen function.

Replacement of forage with cereal byproducts may be a viable alternative for feeding dairy cows. The objective of this experiment was to evaluate total tract digestion and rumen fermentation profile when diets were formulated to contain low-forage neutral detergent fiber (NDF) (12.6% forage NDF, 18.8% total NDF), adequate NDF from forages (20% forage NDF, 24.4% total NDF) or low-forage NDF with high levels of NDF from cereal byproducts (12.7% forage NDF, 35.1% total NDF). Sodium bicarbonate (0.8% of dry matter) was factorialized over these diets. Total tract apparent digestibilities of organic matter (OM) and carbohydrates were determined in 73 Holsteins. Eight rumen-cannulated cows were used concurrently to evaluate rumen fermentation profile and in situ degradation of forages. Bicarbonate did not increase NDF or OM digestibility, but increased intake of digestible OM. Rumen fermentation parameters were determined by dietary alfalfa NDF content. Adding alfalfa NDF to the low-forage, high-starch diet increased in situ degradation of forage NDF more than adding byproduct NDF. However, increased ruminal forage NDF degradability was not reflected in greater total tract NDF digestibility. Replacement of dietary starch with NDF from byproducts decreased OM digestibility, but energy intake was similar across diets due to increased intake.

Animal Feed↗

Response of lactating dairy cows to a cellulase and xylanase enzyme mixture applied to forages at the time of feeding.

Fifty Holstein cows (20 primiparous and 30 multiparous) were used in an experiment to evaluate the dose response to a direct-fed cellulase and xylanase enzyme mixture applied to the forage portion (60% corn silage and 40% alfalfa hay) of a total mixed ration just prior to feeding. Cows were fed one of five treatment diets for 12 wk. Diets 1 through 4 were 55% forage: 45% concentrate and consisted of an untreated control and the control diet plus 0.7, 1.0, or 1.5 L of enzyme concentrate/tonne of forage dry matter, respectively. Diet 5 was an untreated 45% forage: 55% concentrate diet. Actual production of milk increased as much as 10.8% relative to the control diet with 1.5 L of enzyme, and production of fat and protein increased as much as 20 and 13%, respectively. The lowest concentration of enzyme treatment (0.7 L/tonne) accounted for approximately one-half of the production increases. The responses to enzyme-treated forages occurred 2 to 4 wk after the cows started to consume the treated forages, and the same responses were maintained throughout the remainder of the experiment. Cows that started to receive enzyme-treated forage during the first 100 d postpartum produced 9 to 15% more milk and 16 to 23% more energy-corrected milk than did cows fed the control diet. However, production was not increased when cows were in midlactation at the start of the experiment. Responses to enzyme-treated forages fed in 55: 45 forage to concentrate rations were similar to the response to increased proportions of concentrates fed with untreated forages at the 45:55 forage to concentrate ratio.

Animal Feed↗

Interruptions to foraging and learning in a changing environment.

Many resources are both stochastic and variable in their average profitability. Animals have to sample them to track their current states, but whether it is economic to attempt this depends on many factors. Furthermore, there are many interruptions and distractions from foraging (e.g. escape from predators, bad weather, displacement by competitors) which interfere with the acquisition of information. We present a dynamic model of foraging in a stochastic and varying environment, under the constant threat of interruption, to investigate this very general problem. A forager faces two foraging options, one of which provides a known and constant reward, the other providing a reward that is not only stochastic, but whose mean payoff varies in time. The forager has to learn which option has the highest current payoff by sampling. However, interruptions to foraging can occur at any time, the timing and duration of which are beyond the animal's control. When there is a small probability of foraging being interrupted, the forager should forage extensively on the unknown option, but as the probability of interruptions is increased, there is a sudden transition to foraging only on the known option. This occurs because interruptions affect both the level of information required to make exploitation of the unknown option profitable, and the ability to acquire and maintain that information. At what probability of being interrupted this threshold emerges is affected by the value of learning about the unknown option and the duration of interruptions. We discuss the generality of our results with reference to the pervasive problem of updating information in the face of different types of interruption. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

The Effects of a Limited Memory Capacity on Foraging Behavior

We introduce a simple mathematical model to describe the behavior of a forager with a limited memory capacity in the presence of two prey types that differ in their energetic values. The model is used to analyze the effect of memory capacity on foraging behavior and on foraging energetics and it contrasts with classic optimal foraging theory, which implicitly assumes that the forager has an infinite memory capacity. Classic optimal foraging theory dictates that the low value prey type should be invariably excluded from the diet when the high value type exceeds some critical relative frequency and invariably consumed otherwise. Our model forager behaves similarly except that, as its memory capacity declines, it is increasingly predisposed to consume the low value prey when it is suboptimal to do so. Nevertheless, our analysis indicates that the energetic efficiency of a forager with an infinite memory capacity can be approximated by a forager with a memory of modest capacity, perhaps one that contains information about 5-20 previously consumed prey items. In our model, memory constraints necessarily result in so-called "partial preferences" being exercised towards the low value prey type. Thus, limited memory capacity offers a possible explanation for observed violations of the "zero-one" rule of optimal diet composition. We also find a resemblance to a sigmoidal shape in the relationship between the representation of each prey type in the diet and its relative frequency. Thus, the memory-constrained forager exhibits behavior that resembles the type III functional response. Therefore, it is possible that memory constraints such as we envisage can contribute to the explanation of both partial preferences and functional responses. Copyright 1997 Academic Press

Journal Article↗

Simulating Secondary Succession of Elk Forage Values in a Managed Forest Landscape, Western Washington

Modern timber management practices often influence forage production for elk (Cervus elaphus) on broad temporal and spatial scales in forested landscapes. We incorporated site-specific information on postharvesting forest succession and forage characteristics in a simulation model to evaluate past and future influences of forest management practices on forage values for elk in a commercially managed Douglas fir (Pseudotsuga menziesii, PSME)-western hemlock (Tsuga heterophylla, TSHE) forest in western Washington. We evaluated future effects of: (1) clear-cut logging 0, 20, and 40% of harvestable stands every five years; (2) thinning 20-year-old Douglas fir forests; and (3) reducing the harvesting cycle from 60 to 45 years. Reconstruction of historical patterns of vegetation succession indicated that forage values peaked in the 1960s and declined from the 1970s to the present, but recent values still were higher than may have existed in the unmanaged landscape in 1945. Increased forest harvesting rates had little short-term influence on forage trends because harvestable stands were scarce. Simulations of forest thinning also produced negligible benefits because thinning did not improve forage productivity appreciably at the stand level. Simulations of reduced harvesting cycles shortened the duration of declining forage values from approximately 30 to 15 years. We concluded that simulation models are useful tools for examining landscape responses of forage production to forest management strategies, but the options examined provided little potential for improving elk forages in the immediate future.KEY WORDS:Cervus elaphus; Elk; Forage; Forest management; Modeling; Secondary succession

Journal Article↗