Environmental influences on stereotypy and the activity budget of Indian leopards (Panthera pardus) in four zoos in Southern India

To eliminate abnormal behaviors in leopards (Panthera pardus), such as stereotypic pacing, by utilizing environmental enrichment techniques, a proper understanding of their behavior in captive environments is required. Hence there is a need for animal welfare studies in Indian zoos. The activity budgets of 16 leopards were recorded across four southern Indian zoos: Thiruvananthapuram Zoo, Arignar Anna Zoological Park, Shri Chamarajendra Zoological Gardens, and the Guindy Children's Park. Of the 16 study animals, 14 were studied on‐exhibit on zoo holidays as well as on days with visitors present, and all 16 individuals were studied off‐exhibit on other days with visitors present. The 11 behaviors recorded were categorized into active, resting, and stereotypic behaviors. Leopards exhibited higher levels of activity in the on‐exhibit enclosures on days with no visitors. Feeding time influenced the behavioral repertoire of all 14 leopards studied on‐exhibit. Lower proportions of resting were exhibited during the hours before feeding. The proportion of active behaviors differed significantly across zoos. Stereotypic pacing levels were not influenced by the presence of visitors or by feeding time, but was significantly influenced by enclosure features. Higher levels of stereotypic pacing were exhibited in off‐exhibit than on‐exhibit enclosures. Our study shows that the behavior of captive leopards is influenced by enclosure type, feeding regime, and the presence of visitors. Zoo Biol 21:585–595, 2002. © 2002 Wiley‐Liss, Inc.     

Serum and fecal steroid analysis of ovulation,pregnancy, and parturition in the black rhinoceros (Diceros bicornis)

Studies were conducted to determine: (1) if fecal hormone metabolite concentrations correlated with serum estrogen and progesterone concentrations, follicular activity and reproductive behavior in the black rhinoceros (Diceros bicornis) and (2) if threshold values of respective fecal metabolite concentrations correlated with pregnancy. Blood and fecal samples were collected, in conjunction with transrectal ultrasound and behavior observations, for an 18-month period from one black rhinoceros female. Subsequently, serial fecal samples were collected from 13 females in 10 zoos. Quantitative analysis of serum progesterone (P₄) and estradiol (E₂) was performed by radioimmunoassay (RIA): analysis of fecal estrogen metabolites (E) and fecal progesterone metabolites (P) were performed by enzyme immunoassay (EIA). Serum P₂ concentrations identified two luteal phase patterns and two nadirs which corresponded with behavioral estrus. Fecal E patterns indicated a sharp peak which corresponded with breeding. concentrations of fecal P illustrated identifiable nadirs and several peaks which corresponded to serum P₄ nadirs and luteal phases. Serum P₄ concentrations were not different between the luteal phase and pregnancy. Fecal P concentrations started to rise above luteal phase concentrations approximately 150 days postbreeding and remained elevated until immediately before parturition. Serum E₂ and fecal E concentrations rose and subsequently declined after parturition. In the fecal samples from seven pregnant females, fecal P concentrations were similarly elevated compared to six nonpregnant females. Results indicated that fecal steroid metabolites accurately reflected serum steroid hormone concentrations and that the measurement of P and E concentrations permitted the characterization of the estrous cycle, the diagnosis of pregnancy, and the onset of parturition. Zoo Biol 16:121–132, 1997. © 1997 Wiley-Liss, Inc.     

Urinary steroid evaluations to monitor ovarian function in exotic ungulates: V. Estrogen and pregnanediol-3-glucuronide excretion in the black rhinoceros (Diceros bicornis)

A study of female black rhinoceros (Diceros bicomis) urinary steroid and steroid metabolite excretion was performed to determine if techniques useful for monitoring reproductive events in the Indian rhinoceros (Rhinoceros unicornis) could be utilized to evaluate the black rhinoceros. Urine samples from 19 zoo-held black rhinoceros were analyzed for estrogen, estrone conjugates (EC), and pregnanediol-3-glucuronide (PDG) content by direct radioimmunoassays. Estrogen analysis revealed that >95% of the estrogens present in female black rhinoceros urine are conjugated, with estrone and estradiol accounting for virtually all of these estrogens. There is no observable difference in the amount of estrogen present in estrus; postestrus; and early-, mid-, and late-gestation urine samples. Analysis of serial urine samples for EC failed to reveal any discernible levels or patterns which reflected reproductive status. Neither nonpregnant nor early-gestational female black rhinoceros' urine samples contained detectable amounts of PDG. Urinary PDG concentrations became measurable in midgestation (9–12 months prior to parturition) and rose steadily throughout the remainder of gestation. PDG levels declined sharply and became nondetectable 1 day postpartum. Though a wide range in PDG levels was observed among individual pregnant animals, each female consistently excreted increasing amounts of PDG through latter pregnancy.     

Characterization of microsatellite loci in the black rhinoceros (Diceros bicornis) and white rhinoceros (Ceratotherium simum): their use for cross-species amplification and differentiation between the two species

As the population sizes of the black and white rhinoceroses continues to decline, more efforts are needed in multiple areas to help with the conservation efforts. One area being explored is the use of genetic diversity information to aid conservation decisions. In this study, we designed 21 microsatellite primers for white and black rhinoceroses, 16 and 17 of which amplified bands in the white and black rhinoceros, respectively. Out of these primers all 16 were polymorphic in the white rhinoceros and 12 of the 17 were polymorphic in the black rhinoceros. The mean number of alleles was 3.31 and 2.12, the expected heterozygosities were 0.420 and 0.372, and the observed heterozygosities were 0.436 and 0.322 for the white and black rhinoceroses, respectively. Seven of the primers produced different allele sizes and variations that distinguished between black and white rhinoceroses. Further genetic analyses with larger wild population sample sizes and markers are recommended to obtain a better understanding of the genetic structure of the black and white rhinoceros populations in order to be useful in the conservation efforts of these critically endangered species. A. Kilbourn—In memoriam.     

Relationships of reproductive performance indicators in black rhinoceros (Diceros bicornis michaeli) with plant available moisture,plant available nutrients and woody cover

Plant available moisture and plant available nutrients in soils influence forage quality and availability and subsequently affect reproductive performance in herbivores. However, the relationship of soil moisture, soil nutrients and woody forage with reproductive performance indicators is not well understood in mega‐browsers yet these three are important in selecting suitable areas for conservation of mega‐browsers. Here, the eastern black rhinoceros (Diceros bicornis michaeli), a mega‐browser, was studied in seven geographically distinct populations in Kenya to understand the relationships between its reproductive performance indicators and plant available moisture, plant available nutrients and woody cover. Reproductive parameters showed a complex relationship with plant available moisture and plant available nutrients. We found an increase in the predicted yearly percentage of females calving as plant available nutrients decreased in areas of high levels of plant available moisture but no relationship with plant available nutrients in areas of low plant available moisture. Age at first calving was earlier, inter‐calving interval was longer and yearly percentage of females calving was higher at higher woody cover. Woody plant cover contributes positively to black rhino reproduction performance indicators, whereas plant available moisture and plant available nutrients add to the selection of conservation areas, in more subtle ways.     

Evaluation of habitat use and ecological carrying capacity for the reintroduced Eastern black rhinoceros (Diceros bicornis michaeli) in Ruma National Park,Kenya

We carried out a postrelease evaluation to determine predictors of habitat use and carrying capacity for the black rhinoceros (Diceros bicornis michaeli), which are critical for monitoring how the Ruma National Park sub-population may contribute to Kenya's meta-population strategy. We determined whether level of elevation, rockiness, shade, distance to fence, roads, and human settlements predict habitat use, differences in habitat and diet preference between female and male black rhinoceros, and the ecological carrying capacity (CC) of black rhinoceros in the park. We used standard ecological methods to collect data on predictors of habitat use, habitat preference and to estimate CC. Results show, first, that none of the environmental and anthropogenic factors evaluated predicted habitat use by black rhinoceros in the park. Second, although there was no significant difference in habitat preference between the sexes (U = 16.50, p = 0.306), there was a 60% difference in Jaccard's dissimilarity in diet selection between the sexes. Third, the park can support 65 black rhinoceros. Altogether, the findings suggest that the park has potential to support other sub-populations in Kenya. We recommend that future similar studies should incorporate population viability analysis and a community-based approach to forecast the species health and extinction risk.     

Feed intake and dietary composition of iron (Fe), copper (Cu), vitamin E,and tannic acid of five captive black rhinoceros (Diceros bicornis) in a UK collection

The black rhinoceros (Diceros bicornis) is a critically endangered species facing multiple anthropogenic pressures in its natural home range across Africa. Black rhinoceros are difficult to maintain ex situ and subject to diseases that are linked with captive dietary factors. Hemochromatosis is of particular concern, as it is a common finding at necropsy of captive adults, and has been linked to excessive dietary iron intake. This intake study investigates the select nutrient composition of the diets offered to and consumed by five captive black rhinoceros in a UK zoo to evaluate, ensure adequacy, and/or make adjustments if necessary. Alfalfa hay, pellets and six browse species offered were analyzed for iron (Fe), copper (Cu), vitamin E, and tannic acid content. Intakes were quantified and evaluated against levels found in wild diets and the currently available feeding guidelines for black rhinoceros. Diets eaten by five individual rhinoceros (1.4%–2.3% of bodyweight dry matter [DM] intake), comprising 68%–82% hay, 6%–13% pellets, and 13%–27% browse, contained 76–98 mg/kg Fe (on a DM basis), fell within the ranges of plants eaten by free‐ranging rhinoceros (45–140 mg/kg DM), as well as values recommended for captive‐fed browsing rhinoceros (50–100 mg/kg DM). Consumed diets were found to be marginal to adequate in Cu (9–11 mg/kg DM) compared with the recommended 10 mg/kg DM; dietary vitamin E ranged from 54 to 79 IU/kg DM, and tannic acid measured 13–14 g/kg DM. Commercial pellets were the primary contributor of dietary Fe, Cu, and vitamin E, containing up to 10 times more of each of those nutrients than the forages. Native browses were important sources of lower Fe ingredients, as well as appropriate levels of dietary Cu and vitamin E (dependent on species). Interestingly, pellets (23 g/kg) and alfalfa hay (14 g/kg) contained higher concentrations of tannic acid compared with any of the browses fed (4–13 g/kg). All nutritional parameters evaluated were close to recommended dietary levels, diets resembled values consumed in the wild, and the animals remained clinically healthy throughout the study. Overall, diets were considered nutritionally adequate for captive feeding of black rhinoceros; evaluating the nutrient composition of all ingredients, including browse plants in diets, provides important information for achieving optimal nutrient balance.     

Filters and templates: stonefly (Plecoptera) richness in Ouachita Mountains streams, U.S.A.

1.  We collected adult stoneflies periodically over a 1-year period at 38 sites in two headwater catchments in the Ouachita Mountains, Arkansas, U.S.A. The 43 species collected were a subset of the Ozark-Ouachita fauna and the much larger fauna of the eastern U.S.A. We estimated 78–91% species coverage in the two catchments using jackknife extrapolation of species richness from our survey.
2.  Many streams, especially small ones, lacked surface water for months, but others, both small and large, flowed permanently.
3.  Using published regional presence–absence and coarse ecological data in a discriminant function analysis (DFA), we identified stream size (negative) and regional frequency of occurrence (positive) as predictors of presence in these headwater catchments. For the combined catchments, the extrapolated richness (51 spp.) was similar to an estimate (48 spp.) based on predicted absences from DFA and the Ouachita provincial total of known stonefly species (57 spp.).
4.  Local species richness (1–27 spp. per site) was correlated strongly with stream size (catchment area) but was independent of stream drying. Generic richness was correlated negatively with stream drying and positively, but less strongly, with stream size.
5.  Regionally endemic stoneflies dominated in drying streams, and widely distributed species dominated in more permanent streams. The composition of stonefly assemblages was associated with regional factors (species pools, regional abundance, evolution of tolerant endemic species, regional climate) and local factors (drying, stream size).     

Plant size,breeding system,and limits to reproductive success in two sister species of Ferocactus (Cactaceae)

Plant reproductive output can be limited by a variety of factors, bothintrinsic and extrinsic. I investigated the reproductive biologies of twospecies of unbranched short-columnar cacti, Ferocactuscylindraceus and F. wislizeni. I recordedfemalereproductive output (flowers produced, fruit set, seeds per fruit and seedmass), plant size and growth, and used hand-pollination experiments todeterminebreeding systems and pollen limitation. In both species, the ability to selfvaried among individuals, but self-pollination resulted in very few seeds,suggesting strong inbreeding depression. Neither species was pollen-limited.Numbers of flowers produced increases with plant size for both species, andseeds per fruit may also be related to plant size, although the relationship isunclear. Seed mass is not correlated with plant size. Flower production wassimilar in both species, but F. cylindraceus producedfewerseeds per fruit than F. wislizeni, and its seeds weighedless. Fruit set by F. cylindraceus was heavily impacted bya florivorous lepidopteran. Fruit set was very high (94 to 96%) inF. wislizeni, suggesting that architectural constraints(e.g., meristem limitation) are more limiting than resource levels or the levelof pollinator services. In F. cylindraceus, numbers ofseeds per fruit was positively correlated with seed mass, whereas inF. wislizeni, the relationship was negative (tradeoff).Thegrowth rates of F. wislizeni are affected by rainfall theprevious season, and growth rates increase as the plant ages.Ferocactus cylindraceus and F.wislizeni are thought to be sister species, meaning that observeddifferences between them are more likely to be the result of recentevolutionaryprocesses in their lineages rather than differing phylogenetic histories.     

Current and historical mortality from cancer and cardiovascular disease in relation to age: U.S. and world values

Data from the World Health Organization, U.S. Vital Statistics, the Historical Abstracts of the United States and the census of the United States were used to compare current world and states data on cancer death rate and age with historical data that shows the changing percentage of elderly. Current annual death rates from malignant neoplastic disease among the 73 countries for which data exist are highly correlated with the percent of the population ≥ 60 yrs of age and a virtually identical relation exists presently among the 50 states of the U.S. The current world and states data provide a remarkably accurate snapshot of U.S. cancer death rates over the past century during which time the U.S. elderly population increased substantially. Data on U.S. lung cancer death rates since 1940 were similarly examined and these historical numbers also show good agreement with current values among countries that today span the same proportion of elderly that the U.S. had over this period. The agreement of current and historical CA death rate and age contrasts sharply with the picture that emerges when cardiovascular death rates and death rate from all causes are examined. The geographic and time invariance seen in the cancer data seems incosistent with identical exposure to carcinogens and suggests that biological aging is an important intrinsic factor in cancer.     

Male search behavior of the stonefly,Pteronarcella badia (hagen) (Plecoptera: Pteronarcyidae), in relation to drumming

The mating system of many northern hemisphere stoneflies involves vibrational duetting, yet searching behavior in relation to the percussive communication has remained unreported. The communication-search system of a Colorado population of the ubiquitous western stonefly Pteronarcella badiain an experimental arena entails (1) a ranging search by calling virgin or polygynous males until duet establishment with virgin females, (2) continued intermittent duetting while the male engages in a local search and the female remains stationary, and (3) a tactile-based find of the female, followed by immediate mounting and mating. The average 234-s find time for pairs engaging in strong, continuous duets was significantly shorter than those for nonduetting or anomalously duetting pairs. Males of strongly duetting pairs made significantly more turns toward the female during search than in anomalously duetting pairs, and their local search pattern appears to be a triangulation aided by resource-specific (female) vibrational cues. Males displayed a wide range of fitness, based on searching time, and an increased number of duets significantly reduced finding time. Potential female selection and possible extrapolation of these experimental results to natural field conditions are discussed.     

Relationship between androgens, environmental factors and reproductive behavior in male white rhinoceros (Ceratotherium simum simum)

We conducted a longitudinal study of the endocrine activity of free-range male white rhinos. An enzyme immunoassay to measure androgens in the feces was developed and validated to show that it can be used to study testicular activity. We identified two fecal metabolites similar to testosterone and dihydrotestosterone. Several lines of evidence suggest that these metabolites clearly reflect testicular activity. Firstly, the stimulation of testicular activity with synthetic GnRH caused a 156% increase in androgen metabolite concentrations in the feces 1 day after treatment. Secondly, androgen metabolite concentrations increased with sexual maturity in rhinos, and finally there was a correlation between testosterone concentrations in plasma and androgen metabolite concentrations in feces. Using the method that we developed, it was possible to establish whether a relationship exists between androgen metabolite concentrations, the behavior and environmental factors. Adult territorial males (n = 5) had elevated androgen metabolite concentrations during months of high rainfall (September-February) compared to months of little or no rainfall (March-August). The increase in concentrations coincided with the beginning of the rainy season, suggesting a seasonal trend in reproduction. This trend was confirmed by behavior observations showing both a higher frequency of conceptions within the first 4 months of increased androgen metabolite concentrations, and a higher number of inter-sexual conflicts, reflecting the initial aggression between the sexes during the consort period. It was also evident that males accompanying a receptive female had higher fecal androgen metabolite concentrations compared to being alone. The elevated levels were likely induced by female presence.