Activity budgets and the relationship between feeding and stereotypic behaviors in Asian elephants (Elephas maximus) in a Zoo

Activity budgets were studied in eight Asian elephants (Elephas maximus) at Chester Zoo (UK) for 35 days, between January and November 1999. Recordings were made between 10:00 and 16:00 hr (with most behavior frequencies calculated between 10:00 and 14:00 hr). The elephants exhibited variation in activity depending on their age, sex, the time of day and the time of year. Only the five adult cows exhibited stereotypic behavior, with frequencies ranging from 3.9 to 29.4% of all observations. These elephants exhibited individual, diurnal and seasonal variation in stereotypic behavior. This has implications for studies that use short sampling periods and may make comparisons of data collected at different times of the day or year invalid. The six adult elephants spent 27.4–41.4% of the time feeding (between 10:00 and 14:00 hr), 22.9–42.0% standing still, 6.1–19.2% walking and 3.9–9.6% dusting. The hypothesis that the frequency of stereotypic behavior in adult cow elephants was negatively correlated with the frequency of feeding behavior was tested and was found to be true. Stereotypic behavior increased in frequency toward the end of the day—while waiting to return to the elephant house for food—and elephants spent more time stereotyping during the winter months than during the summer months. Elephants were inactive (i.e. exhibited behaviors other than locomotion) for between 70.1 and 93.9% of the time. Creating more opportunities for elephants to exhibit foraging behavior and the introduction of greater unpredictability into management regimes, especially feeding times, may reduce the frequency of stereotypic behavior and increase general activity levels. Zoo Biol 28:79–97, 2009. © 2008 Wiley-Liss, Inc.     

Effects of rubberized flooring on Asian elephant behavior in captivity

Six Asian elephants at the Oregon Zoo were observed to determine the effects of a poured rubber flooring substrate on captive Asian elephant behavior. Room utilization also was evaluated in seven rooms used for indoor housing, including Front and Back observation areas. Data were collected in three phases. Phase I (Baseline Phase) examined elephant behavior on old concrete floors. In Phase II (Choice Phase), elephant behavior was observed in the Back observation area where room sizes were comparable and when a choice of flooring substrates was available. Phase III (Final Phase) examined elephant behavior when all rooms in both observation areas, Front and Back, were converted to rubberized flooring. Room use in both observation areas remained stable throughout the study, suggesting that flooring substrate did not affect room use choice. However, there was a clear pattern of decreased discomfort behaviors on the new rubber flooring. Normal locomotion as well as stereotypic locomotion increased on the new rubber flooring. In addition, resting behavior changed to more closely reflect the resting behavior of wild elephants, which typically sleep standing up, and spend very little time in lateral recumbence. Overall, these findings suggest that the rubber flooring may have provided a more comfortable surface for locomotion as well as standing resting behavior. It is suggested that poured rubber flooring may be a beneficial addition to similar animal facilities. Zoo Biol 0:1–11, 2007. © 2007 Wiley‐Liss, Inc.     

Restrained Eating Behavior and the Metabolic Response to Dietary Energy Restriction in Women

Objective: To determine whether prior eating behavior characterized by dietary restraint alters responses in energy expenditure and substrate oxidation associated with a short‐term, energy‐restricted diet. Research Methods and Procedures: A repeated‐measures, 3‐day diet‐intervention study of adequate (125 kJ/kg of body weight) or restricted (62.5 kJ/kg) energy intake was conducted with 30 women, 20 to 46 years, BMI 25 to 45 kg/m², whose prior eating behavior was “restrained” or “unrestrained.” The Eating Inventory (cognitive restraint subscale) was used to measure restrained eating behavior. Energy expenditure and substrate oxidation were measured after a 12‐hour fast and during the first and fourth hours after a standard meal. Plasma glucose, nonesterified fatty acids, and insulin were measured at corresponding times. Body composition was determined by total body electrical conductivity. Results: Resting energy expenditure was not affected by 3 days of energy restriction. Short‐term energy restriction resulted in lower respiratory‐exchange ratios, higher rates of fat oxidation, and lower rates of carbohydrate oxidation. Subjects classified as restrained eaters had higher postprandial respiratory‐exchange ratios and carbohydrate‐oxidation rates compared with unrestrained eaters. Fasting insulin concentrations were lower in restrained eaters. These effects associated with prior eating behavior were independent of the diet intervention. Discussion: Metabolic outcomes associated with a 3‐day energy‐restricted diet (i.e., increased fat oxidation and decreased carbohydrate oxidation) were not affected by prior restrained eating behavior. However, restrained eating behavior was associated with increased carbohydrate oxidation after a mixed meal. This effect of restrained eating behavior may be attributable to increased insulin sensitivity.     

Nocturnal behavior in a group of female African elephants

The nocturnal behavior of a stable group of female, African elephants (Loxodonta africana africana) was studied to: (1) examine their behavior as a function of hour of night; (2) qualitatively compare the elephants' activity budgets to those reported in a previous study; and (3) document the presence of aggressive and stereotypic behaviors that might necessitate a change in their management. The elephants were systematically observed at least five times per week for 10 weeks between 17:00 and 08:00. Instantaneous focal samples of behavior, location, and proximity were taken every minute on a rotating basis, and all observed occurrences of social behavior were recorded. The hour of night affected elephant activity: significant relationships were found between hour of night and percent of time they spent feeding, lying, and standing. The overall activity budgets of the elephants were similar to the activity budgets reported in a previous study, although differences were evident in lying, stereotypic, and social behaviors. These differences might be a function of age. Affiliative behaviors accounted for 57% of the elephants' social behaviors, and agonistic behaviors among the elephants occurred infrequently and caused no injuries. Additionally, the elephants used all areas to which they had access. These findings provide compelling evidence that unrestricted social access during the night is the appropriate management strategy for these elephants. The results from the present study also highlight the importance of replicating existing studies and using multiple behavioral measures to make decisions regarding the welfare and management of stable groups of captive elephants. Zoo Biol 25:173–186, 2006. © 2006 Wiley‐Liss, Inc.     

Salient features in the locomotion of proboscideans revealed via the differential scaling of limb long bones

The standard differential scaling of proportions in limb long bones (length against circumference) was applied to a phylogenetically wide sample of the Proboscidea, Elephantidae and the Asian (Elephas maximus) and African (Loxodonta africana) elephants. In order to investigate allometric patterns in proboscideans and terrestrial mammals with parasagittal limb kinematics, the computed slopes between long bone lengths and circumferences (slenderness exponents) were compared with published values for mammals, and studied within a framework of the theoretical models of long bone scaling under gravity and muscle forces. Limb bone allometry in E. maximus and the Elephantidae is congruent with adaptation to bending and/or torsion induced by muscular forces during fast locomotion, as in other mammals, whereas the limb bones in L. africana appear to be adapted for coping with the compressive forces of gravity. Hindlimb bones are therefore more compliant than forelimb bones, and the resultant limb compliance gradient in extinct and extant elephants, contrasting in sign to that of other mammals, is shown to be a new important locomotory constraint preventing elephants from achieving a full‐body aerial phase during fast locomotion. Moreover, the limb bone pattern of African elephants, indicating a noncritical bone stress not increasing with increments in body weight, explains why their mean and maximal body masses are usually above those for Asian elephants. Differences in ecology may be responsible for the subtle differences observed in vivo between African and Asian elephants, but they appear to be more pronounced when revealed via mechanical patterns dictated by limb bone allometry. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 16–29.     

How selective are elephants as agents of forest tree damage in Bwindi Impenetrable National Park,Uganda?

Elephants are locally concentrated in Bwindi Impenetrable National Park. Vegetation damage attributable to elephants appears to be increasing and may result in the modification of the forest. We examined the implied selectivity of stem damage due to elephants. We followed 26.84 km of recent elephant trails and used 122 plots to document tree damage in relation to species, stem sizes and locations. Of 897 trees (DBH ≥2 cm), 542 (60.4%) were intact, 22 (2.5%) debarked, 274 (30.5%) toppled and 172 (19.2%) had broken branches. Small trees were more likely to be pushed over or have their branches broken, whereas large trees were more commonly debarked. The species most frequently selected for damage included mid‐successional species such as Newtonia buchananii, Myrianthus holstii and Chrysophyllum albidum. These species may be vulnerable to increasing elephant numbers. Our analyses using general linear models indicate that elephants are selective concerning where, how and what tree stems they damage. We found a higher incidence of elephant damage per‐tree stem in open areas than in more closed areas, suggesting feedback in which elephants maintain open habitats that may be conducive for other species such as mountain gorillas. More work is needed to better determine how changing elephant numbers may influence Bwindi's conservation values.     

Graph theory illustrates spatial and temporal features that structure elephant rest locations and reflect risk perception

Understanding the spatial structuring of animal behaviors and how they link landscapes can be critical for conservation management. This emerging field has been greatly facilitated by technologically advanced acquisition and analysis of data on animal movements. The framework of graph theory, which directly quantifies network connectivity properties, provides a useful addition to this tool set. Using a novel application of graph theory, we investigate the structure and patterning of African elephant Loxodonta africana rest sites, a potentially critical feature structuring spatial properties of animal populations. Elephants in the study rested intermittently and for short durations (1–3 rests d^–1, lasting 3–5 h total). They switched circadian rest patterns according to landscape attributes, resting more during the day and further from permanent water in areas with high human density outside protected areas. Within protected areas and during the dry season, elephants showed clustering and sequential use of rest nodes (repeated motifs). Repeated use of specific rest nodes (self‐looping) was more frequent than expected if rest nodes were chosen at random, particularly when outside protected areas further from water, indicating the importance of preferred rest sites. Our results suggest that elephants adjust resting behavior when in human‐dominated areas, using preferred resting sites presumably in locations that reduce the risk of interactions. This study demonstrates how graph theory may be used practically to gain novel insight into behaviours, such as resting, that are discrete in time and space. Furthermore, analysis of the spatial and network properties of rest sites, given an individual's susceptibility when engaged in rest behavior, allowed characterization of spatio‐temporal risk perception, providing a powerful behavioral based means to quantify the landscape of fear.     

PHYLOGEOGRAPHY OF THE ASIAN ELEPHANT (ELEPHAS MAXIMUS) BASED ON MITOCHONDRIAL DNA

Abstract Populations of the Asian elephant (Elephas maximus) have been reduced in size and become highly fragmented during the past 3000 to 4000 years. Historical records reveal elephant dispersal by humans via trade and war. How have these anthropogenic impacts affected genetic variation and structure of Asian elephant populations? We sequenced mitochondrial DNA (mtDNA) to assay genetic variation and phylogeography across much of the Asian elephant's range. Initially we compare cytochrome b sequences (cyt b) between nine Asian and five African elephants and use the fossil‐based age of their separation (～5 million years ago) to obtain a rate of about 0.013 (95% CI = 0.011–0.018) corrected sequence divergence per million years. We also assess variation in part of the mtDNA control region (CR) and adjacent tRNA genes in 57 Asian elephants from seven countries (Sri Lanka, India, Nepal, Myanmar, Thailand, Malaysia, and Indonesia). Asian elephants have typical levels of mtDNA variation, and coalescence analyses suggest their populations were growing in the late Pleistocene. Reconstructed phylogenies reveal two major clades (A and B) differing on average by HKY85/Γ‐corrected distances of 0.020 for cyt b and 0.050 for the CR segment (corresponding to a coalescence time based on our cyt b rate of ～1.2 million years). Individuals of both major clades exist in all locations but Indonesia and Malaysia. Most elephants from Malaysia and all from Indonesia are in well‐supported, basal clades within clade A, thus supporting their status as evolutionarily significant units (ESUs). The proportion of clade A individuals decreases to the north, which could result from retention and subsequent loss of ancient lineages in long‐term stable populations or, perhaps more likely, via recent mixing of two expanding populations that were isolated in the mid‐Pleistocene. The distribution of clade A individuals appears to have been impacted by human trade in elephants among Myanmar, Sri Lanka, and India, and the subspecies and ESU statuses of Sri Lankan elephants are not supported by molecular data.     

Long‐term administration of α‐tocopherol in captive Asian elephants (Elephas maximus)

After the loss of an African elephant (Loxodonta africana) in February 1989 at the Denver Zoological Gardens (DZG) with very low circulating serum α‐tocopherol, a long‐term study was initiated with three Asian elephants (Elephas maximus) to evaluate the effect of an oral micellized, water‐soluble, natural source d‐α‐tocopherol supplement. Baseline α‐tocopherol levels were evaluated and found to be approximately 3.75‐fold less than those reported for semi‐free‐ranging Asian Nepalese work camp and free‐ranging African elephants. The DZG elephants were then administered a liquid d‐α‐tocopherol (Emcelle^®) at 2.2 IU/kg body weight orally once daily. Serum samples were obtained and analyzed at 1, 2, 8, and 12 months and then annually for 96 months. The oral vitamin E supplement significantly elevated serum levels above baseline and were found to be comparable with levels reported for semi–free‐ranging and free‐ranging elephants. Zoo Biol 20:245–250, 2001. © 2001 Wiley‐Liss, Inc.     

The Eating Inventory and Body Adiposity from Leanness to Massive Obesity: a Study of 2509 Adults

Objective: To examine how chronic dietary restraint, disinhibition, and hunger, as assessed by the Eating Inventory, vary over a broad range of BMI values from leanness to massive obesity, in subjects with family obesity. Eating Inventory factors were also studied as a function of personal weight history. Research Methods and Procedures: Subjects were 2509 participants in a genetic study of obesity. BMIs ranged from 15 to 87 kg/m². Six BMI groups were formed (<27, 27 to 30, 30 to 35, 35 to 40, 40 to 45, and >45). Results: Multivariate analyses showed that restraint and disinhibition were significantly associated with BMI in men, whereas only disinhibition was in women. Disinhibition scores correlated strongly with hunger scores in both genders in all BMI categories; dietary restraint tended to correlate with the other two factors positively in leaner subjects and negatively in the highest BMI categories. Highly restrained normal‐weight subjects were likely to exhibit disinhibition and hunger, whereas massively obese persons with very high disinhibition scores showed high hunger but little restraint. The highest restraint scores were observed in nonobese adult women with previous obesity in childhood and/or adolescence. Discussion: The factor most strongly associated with BMI in this large population was disinhibition, suggesting that obesity treatment should target behaviors associated with disinhibition, especially in individuals showing a low level of dietary restraint. High restraint scores in formerly obese normal‐weight women suggest that dietary restraint may exert a beneficial influence on body weight control under conditions that deserve further investigation.     

On the Relation of Attempting to Lose Weight,Restraint, and Binge Eating in Outpatients with Binge Eating Disorder

Objective: To examine the relationship among attempts to lose weight, restraint, and eating behavior in outpatients with binge eating disorder (BED). Research Methods and Procedures: Participants were 93 consecutive outpatients evaluated for a clinical trial who met Diagnostic and Statistical Manual, Fourth edition criteria for BED. The Eating Disorder Examination Interview was administered to assess attempts at weight loss, restraint, different forms of overeating, and the attitudinal psychopathology of eating disorders (i.e., concerns regarding eating, shape, and weight). In addition, the Three‐Factor Eating Questionnaire was used to assess cognitive restraint, hunger, and disinhibition. Psychometrically established measures were given to assess body dissatisfaction, depression, and self‐esteem. Results: The majority of participants (75.3%; N = 70) reported attempting to lose weight, but only 37.6% (N = 35) reported dietary restraint on at least half the days of the month. Dietary restraint and cognitive restraint were not associated with any form of binge eating or overeating. Dietary restraint and cognitive restraint were positively correlated with weight concern, shape concern, and body dissatisfaction, and negatively correlated with body mass index. To further examine the interplay between attempting to lose weight and restraint, three study groups were created: unrestrained nonattempters (21.5%, N = 20), unrestrained attempters (40.9%; N = 38), and restrained attempters (34.4%; N = 32). The three groups did not differ significantly on binge eating or other eating behaviors; however, significant differences were observed for weight concern, shape concern, and body dissatisfaction. Discussion: Attempts to lose weight and restraint are not synonymous for patients with BED. Although 75.3% of BED patients reported that they were attempting to lose weight, only 37.6% reported dietary restraint on at least half the days of the previous month. While restraint was negatively associated with body mass index, it was not related to binge eating or overeating. Our findings raise questions about prevailing models that posit restraint as a predominant factor in the maintenance of binge eating in BED.     

Features and Trajectories of Eating Behavior in Weight‐Loss Maintenance: Results from the German Weight Control Registry

Objective

Eating behaviors such as dietary restraint and disinhibition caused by emotional and external cues play a relevant role in weight‐loss maintenance.

Methods

Four hundred forty individuals with successful weight‐loss maintenance included in the prospective German Weight Control Registry completed the Dutch Eating Behavior Questionnaire. Participants were categorized into the following two weight groups: stable weight trajectory (WS) (n = 280) and unstable weight trajectory (WUS) (n = 160) over the 2‐year assessment period.

Results

Those with successful weight‐loss maintenance had significantly higher scores on the restrained and emotional eating subscales compared with a general population (GP) sample. At baseline, the WS individuals had lower restrained, emotional, and external eating scores compared with the WUS individuals. Over the 2‐year follow‐up period, the trajectories of the restraint scores decreased in both groups but stayed elevated compared with the GP sample. Scores of the emotional and external eating subscales remained stable in the WS group but increased in the WUS group.

Conclusions

A certain degree of restraint seems to be necessary for successful weight‐loss maintenance; however, high emotional and external eating may counteract this effect, resulting in weight regain in the long run.     

Why inter‐country loans will not help sumatra's elephants

Asian elephants (Elephas maximus) in western zoos are likely to become extinct unless elephants are regrouped into breeding units or additional elephants are imported from range States. There have been proposals for the export of elephants from elephant camps in Sumatra, Indonesia. In exchange, zoos would be expected to provide funds or support ‘in kind’ for the camps or for the conservation of wild elephants. Most of the elephants in the Sumatran camps were captured because of crop‐raiding problems around protected areas or because elephant habitat has been and continues to be lost to development schemes and illegal conversion of protected areas to agriculture. Capture‐related mortality rates are high and conditions in the camps are poor, with low standards of veterinary care and husbandry. This is partly due to over‐crowding and inadequate budgets. It might seem, therefore, that the loan of elephants to western zoos would improve the lot of these elephants and reduce the pressures on the camps. However, we show that both total and annual demand for Asian elephants, and particularly Sumatran elephants (E. m. sumatranus), by western zoos are low, and consequently the resources generated by any loan scheme would be limited. Elephant loan schemes are unlikely to have significant beneficial impact on either the conservation or welfare of elephants in Sumatra. More importantly, a credible loan scheme would require a permanent moratorium on the capture of wild elephants in Sumatra. Such a moratorium is needed to prevent illicit captures for sale or loan. At present, wild elephants are caught to replace those that die at the camps or are moved to other facilities. Without a moratorium, the loan of elephants to overseas zoos would contribute directly to reductions in wild elephant populations in Sumatra. However, a moratorium is likely to prove impossible to enforce, and this alone should call into question the desirability of any loan scheme. Zoo Biol 25:235–246, 2006. © 2006 Wiley‐Liss, Inc.     

A TECHNIQUE TO ‘TAG’ SPECIFIC VOCALIZATIONS IN TAPE-RECORDINGS OF COMPLEX VOCAL INTERACTIONS

ABSTRACT

It has been suggested that African savanna elephants Loxodonta africana produce 31 different call types (Langbauer 2000). Various researchers have described these calls by associating them with specific behavioural contexts. More recently Leong et al. (2003) have attempted to classify elephant call types based on their physical properties. They classified 8 acoustically distinct call types from a population of captive elephants. This study focuses on one of these call types, the rumble, in a wild population of elephants in Kruger National Park, South Africa. A single family group of elephants was followed to record group behaviours and vocalizations from January through August 2001. By measuring the physical properties of 663 rumbles and subjecting these to cluster analysis, we present evidence that shows that rumbles can be categorized by their physical properties and that the resulting rumble types are associated with specific group behaviours. We characterize three types of rumbles that differ significantly by ten acoustic parameters. Two rumble types were associated with the elephant group feeding and resting, while the third was associated with socializing and agitation.     

The behavioral effects of exhibit size versus complexity in African elephants: A potential solution for smaller spaces

Population‐level analyses suggest that habitat complexity, but not necessarily space availability, has important welfare outcomes for elephants in human care. At the Dallas Zoo, the opening of a new exhibit complex allowed us to measure the behavior of two female African elephants across three treatments to evaluate the independent effects of complexity and space. Preoccupancy observations were conducted in the elephants’ older exhibit, which consisted of a smaller, more simple yard (630 m²). Subsequent postoccupancy observations measured behavior in two different spaces in the new exhibit: a larger, complex yard (15,000 m²), and a smaller, but complex yard (1,520 m²). The elephants’ overall activity levels were greater in complex habitats, regardless of their size. Similar effects of habitat complexity oversize were observed with greater rates of foraging and lower rates of being stationary. Furthermore, elephants were out of view of visitors significantly more in the small, simple yard compared to either of the more complex habitats. However, exhibit size affected the incidence of stereotypic behavior (with lower rates of stereotypy in the larger exhibit compared to the smaller yards) and investigatory behavior (elephants investigated their environments more with increasing size and complexity). Behavioral diversity also increased with exhibit size and complexity. These results indicate that space availability alone is not sufficient to enhance the behavioral welfare of zoo elephants. Therefore, facilities with limited space can still encourage species‐appropriate behaviors and improved welfare for the elephants in their care by converting a small, simple area into a more complex habitat.     

Birth seasonality and calf mortality in a large population of Asian elephants

In seasonal environments, many species concentrate their reproduction in the time of year most likely to maximize offspring survival. Asian elephants (Elephas maximus) inhabit regions with seasonal climate, but females can still experience 16‐week reproductive cycles throughout the year. Whether female elephants nevertheless concentrate births on periods with maximum offspring survival prospects remains unknown. We investigated the seasonal timing of births, and effects of birth month on short‐ and long‐term mortality of Asian elephants, using a unique demographic data set of 2350 semicaptive, longitudinally monitored logging elephants from Myanmar experiencing seasonal variation in both workload and environmental conditions. Our results show variation in birth rate across the year, with 41% of births occurring between December and March. This corresponds to the cool, dry period and the beginning of the hot season, and to conceptions occurring during the resting, nonlogging period between February and June. Giving birth during the peak December to March period improves offspring survival, as the odds for survival between age 1 and 5 years are 44% higher for individuals born during the high birth rate period than those conceived during working months. Our results suggest that seasonal conditions, most likely maternal workload and/or climate, limit conception rate and calf survival in this population through effects on maternal stress, estrus cycles, or access to mates. This has implications for improving the birth rate and infant survival in captive populations by limiting workload of females of reproductive age. As working populations are currently unsustainable and supplemented through the capture of wild elephants, it is imperative to the conservation of Asian elephants to understand and alleviate the effects of seasonal conditions on vital rates in the working population in order to reduce the pressure for further capture from the wild.     

From left to right all through the night: Characteristics of lying rest in zoo elephants

Despite increased research during the past years, many characteristics of resting behavior in elephants are still unknown. For example, there is only limited data suggesting elephants express longer lying bouts and increased total nightly lying durations on soft substrates as compared to hard surfaces. Additionally, it has not been investigated how frequently elephants change body sides between lying bouts. Here we present these characteristics based on observations of nighttime lying behavior in 10 zoo elephants (5 African Loxodonta africana and 5 Asian Elephas maximus elephants) living in five different European facilities. We found that elephants housed on soft substrates have significantly increased total lying durations per night and longer average lying bouts. Furthermore, at 70%−85% of all bouts, a consistently higher frequency of side change between lying bouts occurred on soft substrates, leading to an overall equal laterality in resting behavior. Deviations from this pattern became evident in elephants living on nonsand flooring or/and in nondominant individuals of nonfamily groups, respectively. Based on our findings, we consider elephants to normally have several lying bouts per night with frequent side changes, given an appropriate substrate and healthy social environment. We encourage elephant-keeping facilities to monitor these characteristics in their elephants' nighttime behavior to determine opportunities for further improvements and detect alterations putatively indicating social or health problems in individual elephants at an early stage.     

Behavioral effects of environmental enrichment on harbor seals (Phoca vitulina concolor) and gray seals (Halichoerus grypus)

Zoos and aquariums have been incorporating environmental enrichment into their animal care programs for the past 30 years to increase mental stimulation and promote natural behaviors. However, most attempts to document the effects of enrichment on animal behavior have focused on terrestrial mammals. Staff at the National Aquarium in Baltimore conducted an investigation of the behavioral effects of enrichment on the seven harbor seals and two gray seals housed in the aquarium's outdoor seal exhibit. We expected that enrichment would change the amount of time the animals spent engaged in specific behaviors. The behaviors recorded were: resting in water, resting hauled out, maintenance, breeding display, breeding behavior, aggression, pattern swimming, random swimming, exploration, and out of sight. Activity levels (random swimming and exploration) were expected to increase, while stereotypic behaviors (pattern swimming) were expected to decrease. The frequency and duration of behaviors were documented for 90 hr in both the control phase (without enrichment) and the experimental phase (with enrichment). Statistically significant differences (P<0.05) in the time spent in pattern swimming, random swimming, exploration, and out of sight were observed between the two phases. With enrichment, pattern swimming and out of sight decreased, while random swimming and exploration behavior increased. These findings demonstrate that enrichment can promote behaviors (random swimming and exploration) that are likely to be normal for phocids in the wild, and that may contribute to the behavioral complexity of these seals in captivity. Zoo Biol 21:375–387, 2002. © 2002 Wiley‐Liss, Inc.     

Modelling large herbivore movement decisions: Beyond food availability as a predictor of ranging patterns

The ability of animals to adapt to their changing environment will depend in part on shifts in their ranging patterns, but when and why individuals choose to move requires detailed understanding of their decision‐making processes. We develop a simple decision‐making model accounting for resource availability in habitually used ranges. We suggest that disparities between model predictions and animal tracking data indicate additional factors influencing movement decisions, which may be identified given detailed system‐specific knowledge. The model was evaluated using movement data from satellite‐tracked elephants (Loxodonta africana) inhabiting the Amboseli basin in Kenya, moving from savannah areas with low quality but constant resource availability, to areas with temporally constrained higher nutrient availability. Overall, the model fits the data well: There was a good correlation between predicted and observed locations for the combined data from all elephants, but variation between individuals in how well the model fits. For those elephants where model predictions were less successful, additional factors likely to affect movement decisions, including reproduction, anthropogenic threats, memory and perception are suggested. This protocol for building and testing decision‐making models should contribute to success in attempts to preserve sufficient space for large herbivores in their increasingly human‐dominated ecosystems.     

Variation in the bitter-taste receptor gene TAS2R38, and adiposity in a genetically isolated population in Southern Italy

Objective: Variation in the bitter‐taste receptor gene, TAS2R38 confers the ability to taste 6‐n‐propylthiouracil (PROP). The objective of this study was to relate TAS2R38 haplotypes and PROP‐tasting phenotypes to adiposity in a genetically isolated population. We hypothesized that the nontaster phenotype would be associated with higher BMI and waist circumference (WC) in females, and that dietary restraint would mediate this relationship. Methods and Procedures: Participants were 540 healthy inhabitants of the genetically isolated village of Carlantino in southern Italy who were 15–89 years of age at the time of the study. Haplotype analyses were performed and PROP tasting was assessed using a filter paper method. Height, weight, and WC were measured and restrained eating was assessed using a brief questionnaire. Results: Nontaster females had higher BMI and WC than females who were phenotypic tasters, and this relationship was specific to females with low dietary restraint. Regression analysis showed that BMI declined by 1.7 units across taster groups in females when the model included the PROP by restraint interaction. PROP phenotype was not significantly associated with WC in the regression models. Polymorphisms in TAS2R38 were not associated with BMI or WC in females. Neither TAS2R38 haplotype nor PROP phenotype was strongly related to BMI or WC in males. Discussion: These data support previous findings of a relation between the nontaster phenotype and higher BMI in females that is modified by dietary restraint. Assessment of PROP phenotypes might provide unique information about adiposity that is not captured by haplotype analysis alone.