Acid‐insoluble ash as a measure of dry matter digestibility in captive African elephants (Loxodonta africana)

There are limited data on the diet dry matter digestibility (DMD) of captive African elephants. Although the total fecal collection method is the standard for determining DMD, it is labor‐intensive, time‐consuming, and expensive. The acid‐insoluble ash (AIA) marker technique has been used successfully to determine DMD in ruminants and monogastrics. The objective of this study was to assess how accurately the AIA marker technique could estimate the DMD of captive African elephants (Loxodonta africana). Three mature male African elephants at Disney's Animal Kingdom in Florida were used in this study. The animals were offered a Bermuda grass hay‐based ration, and the total dry matter intake (DMI) and total fecal output were measured daily over a 7‐day period to determine the total collection DMD. The feed ingredients and fecal samples were also analyzed for AIA. Although there were differences (P<0.05) in total DMI and total fecal outputs, the DMD values did not (P>0.05) differ (35.1±0.72 vs. 37.1±0.72 for total collection and AIA, respectively). There was a linear (y=0.9461x; R²=0.74) relationship between the total collection and AIA marker technique DMD values. These results suggest that AIA can be used to accurately estimate the DMD of captive African elephants. Zoo Biol 0:1–5, 2005. © 2005 Wiley‐Liss, Inc.     

Social factors influence ovarian acyclicity in captive African elephants (Loxodonta africana)

Nearly one-third of reproductive age African elephants in North America that are hormonally monitored fail to exhibit estrous cycle activity, which exacerbates the nonsustainability of the captive population. Three surveys were distributed to facilities housing female African elephants to determine how social and environmental variables contribute to cyclicity problems. Forty-six facilities returned all three surveys providing information on 90% of the SSP population and 106 elephants (64 cycling, 27 noncycling and 15 undetermined). Logistic analyses found that some physiological and social history variables were related to ovarian acyclicity. Females more likely to be acyclic had a larger body mass index and had resided longer at a facility with the same herdmates. Results suggest that controlling the weight of an elephant might be a first step to helping mitigate estrous cycle problems. Data further show that transferring females among facilities has no major impact on ovarian activity. Last, social status appears to impact cyclicity status; at 19 of 21 facilities that housed both cycling and noncycling elephants, the dominant female was acyclic. Further studies on how social and environmental dynamics affect hormone levels in free-living, cycling elephants are needed to determine whether acyclicity is strictly a captivity-related phenomenon. Zoo Biol 28:1–15, 2009. © 2008 Wiley-Liss, Inc.     

The relationship between social behaviour and habitat familiarity in African elephants (Loxodonta africana)

Social associations with conspecifics can expedite animals'' acclimation to novel environments. However, the benefits gained from sociality may change as the habitat becomes familiar. Furthermore, the particular individuals with whom animals associate upon arrival at a new place, familiar conspecifics or knowledgeable unfamiliar residents, may influence the type of information they acquire about their new home. To examine animals'' social dynamics in novel habitats, we studied the social behaviour of African elephants (Loxodonta africana) translocated into a novel environment. We found that the translocated elephants'' association with conspecifics decreased over time supporting our hypothesis that sociality provides added benefits in novel environments. In addition, we found a positive correlation between body condition and social association, suggesting that elephants gain direct benefits from sociality. Furthermore, the translocated elephants associated significantly less than expected with the local residents and more than expected with familiar, but not necessarily genetically related, translocated elephants. The social segregation between the translocated and resident elephants declined over time, suggesting that elephants can integrate into an existing social setting. Knowledge of the relationship between sociality and habitat familiarity is highly important in our constantly changing world to both conservation practice and our understanding of animals'' behaviour in novel environments.     

Physiological indicators of stress in African forest elephants (Loxodonta africana cyclotis) in relation to petroleum operations in Gabon,Central Africa

Aim Human activities are major determinants of forest elephant (Loxodonta africana cyclotis) distribution in Gabon, but the types and intensity of disturbance that elephants can tolerate are not known. We conducted dung surveys within the Gamba Complex of Protected Areas in SW Gabon to examine (1) the feasibility of noninvasive faecal analyses for monitoring stress physiology, and (2) the influence of petroleum operations on stress levels in forest elephants. Location Gabon, Central Africa. Methods We identified multiple dung piles from the same individual by matching their eight‐locus microsatellite genotypes, and measured faecal concentrations of glucocorticoid metabolites as an indicator of stress in areas subject to different levels of disturbance: (1) Loango National Park (2) an ‘industrial corridor’ dominated by oil fields, and (3) a nearby area of human settlements. Results We obtained unique microsatellite genotypes and faecal glucocorticoid metabolite (FGM) concentrations for 150 forest elephant individuals, which is the largest hormonal data set for wild African forest elephants to date. Adults exhibited higher mean FGM concentrations than juveniles, and in contradiction of our expectations of chronic stress around oil fields, elephants in Loango National Park exhibited significantly higher FGM concentrations than elephants in the industrial corridor. Main conclusions We argue that forest elephants in the industrial corridor of the Gamba Complex have become acclimated to oil fields, resulting in part from oil company regulations that minimize stressful interactions between elephants and petroleum operations. Our findings for a flagship species with substantial ecological requirements bode well for other taxa, but additional studies are needed to determine whether oil operations are compatible over their life span with rain forest ecosystems in Central Africa.     

Behavioral indices of estrus in a group of captive African elephants (Loxodonta africana)

This study investigated behavioral signals of estrus by systematically monitoring the interactions of one male with four female African elephants housed in a naturalistic outdoor enclosure at Disney's Animal Kingdom over a period of 11 months. We measured changes in five spatial behaviors and 22 tactile‐contact behaviors, as well as changes in serum progestagen and LH concentrations, across three ovarian cycles for each female. Two females did not cycle during the study. Three different phases of the ovarian cycle were identified: mid luteal, anovulatory follicular, ovulatory follicular. The male followed more and carried out more genital inspections, flehmen, and trunk‐to‐mouth behaviors toward cycling females during their ovulatory phase. Genital inspections by the male peaked above baseline levels on the day of an LH surge, and up to 9 days before, in both cycling females and, thus, might be a useful behavioral index of estrus. The male also carried out more genital inspections, flehmen, and trunk touches to the back leg toward ovulatory cycling than noncycling females. Overall, our results indicated that: 1) a single subadult African elephant male could discriminate two females in the ovulatory phase of their cycle (i.e., during the 3 weeks preceding ovulation) from the mid luteal phase; 2) the male also discriminated two cycling females in the ovulatory and anovulatory follicular phases from two noncycling females; 3) two females in the ovulatory phase of the cycle displayed a greater variety of tactile‐contact behavior toward the male compared to the other cycle phases. Zoo Biol 0:1–19, 2005. © 2005 Wiley‐Liss, Inc.     

Seed protection through dispersal by African savannah elephants (Loxodonta africana africana) in northern Tanzania.

We examined effectiveness of African savannah elephant dung as a protective barrier for seeds of three tree species, Acacia tortilis Hayne, Tamarindus indica L. and Ximenia aegyptiaca L. Seeds were collected from dung and underneath fruiting trees in Tarangire National Park, Tanzania. Experimental treatments were established to test: (i) the efficacy of dung in protecting seeds of A. tortilis from bruchid beetle infestation and the role of animals larger than insects in removing seeds; (ii) the same tenets as in Experiment 1, using seeds of T. indica; and (iii) the effect of distance on survival of seeds of X. aegyptiaca. Sites were established during two field seasons underneath conspecific trees, where seed predation was likely highest. Repeated‐measures two‐way ANOVA indicated that there was no treatment effect for Experiment 1. For Experiments 2 and 3 in October 2013, seeds in dung experienced less beetle infestation than fresh seeds. Repeated‐measures two‐way ANOVA and Tukey's HSD indicated that treatment effect differed among all treatments. Passed seeds at distances ≥5 m experienced less beetle infestation than fresh seeds underneath conspecifics. African savannah elephants appear to be important seed dispersers of these three tree species.     

Characterization of tetranucleotide microsatellite loci in the African Savannah Elephant (Loxodonta africana africana)

Most African elephant (Loxodonta africana africana) populations are isolated and thus threatened by a loss of genetic diversity. As a consequence, genetic analysis of African elephant populations will play an increasing role in their conservation, and microsatellite loci will be an important tool in these analyses. Previously published sets of polymorphic microsatellites developed for African elephants are all dinucleotide repeats, which are prone to typing error. Here, we characterize 11 tetranucleotide microsatellite loci in the African elephant. All loci were polymorphic in 32 faecal samples and two tissue samples from 33 individual African savannah elephants.     

Assessing the effect of roads on impala (Aepyceros melampus) stress levels using faecal glucocorticoid metabolites

In Tanzania, a proposed road through the Serengeti National Park has caused international debate as to whether it will cause irreversible damage to the ecosystem. We assessed the effect of current roads and traffic in the Serengeti ecosystem on a model ungulate species, the impala antelope (Aepyceros melampus). We measured how faecal glucocorticoid metabolites (FGM), an indicator of physiological stress levels, were affected by different levels of traffic while accounting for other stress‐related variables such as distance to roads, vegetation type and group size. A total of 196 faecal samples were collected. FGM were measured using an enzyme immunoassay (EIA) validated for several ruminants. FGM levels were significantly elevated near major roads with higher traffic levels compared to roads with less traffic. Predators and illegal hunting might also affect FGM levels in the impala, but the potential effect of these variables was inconsistent with the recorded FGM levels and cannot explain the observed variation. These results show for the first time that an African mammal exhibits significant physiological stress in relation to roads and traffic.     

Differences in woody vegetation are unrelated to use by African elephants (Loxodonta africana) in Mkhuze Game Reserve, South Africa

The reintroduction of African elephants into fenced game reserves throughout South Africa has presented managers with several challenges. Although elephants are a natural part of southern African ecosystems, their confinement to fenced protected areas in South Africa has exacerbated their potential to impact their habitats negatively. However, many studies investigating the impact of elephants have failed to control for the effects of other browsers on the vegetative community. In this study, we used location data on an elephant herd to delineate high-use and low-use areas. This paired design allowed us to minimize confounding factors that could explain differences in the structure, diversity and utilization of woody species. We found little evidence to suggest elephant-mediated change in, or selection for, the structure or diversity of woody species; however, our results suggest that elephants may be altering the composition of species by preferentially using areas with higher canopy diversity and by enhancing sapling recruitment. Although stripping of bark was higher in high-use areas, there was no evidence of differential mortality of tree species. Therefore, in our study area, and over the current time scale, elephants are having a negligible impact on the vegetative community.     

Seasonal habitat use by Elephants (Loxodonta africana) in the Mole National Park of Ghana

To avoid unnecessary waste of limited resources and to help prioritize areas for conservation efforts, this study aimed to provide information on habitat use by elephants between the wet and dry seasons in the Mole National Park (MNP) of Ghana. We compiled coordinates of 516 locations of elephants’ encounters, 256 for dry season and 260 for wet season. Using nine predictor variables, we modeled the probability of elephant's distribution in MNP. We threshold the models to “suitable” and “nonsuitable” regions of habitat use using the equal training sensitivity and specificity values of 0.177 and 0.181 for the dry and wet seasons, respectively. Accuracy assessment of our models revealed a sensitivity score of 0.909 and 0.974, and a specificity of 0.579 and 0.753 for the dry and wet seasons, respectively. A TSS of 0.488 was also recorded for the dry season and 0.727 for the wet season indicating a good model agreement. Our model predicts habitat use to be confined to the southern portion of MNP due to elevation difference and a relatively steep slope that separates the northern regions of the park from the south. Regions of habitat use for the wet season were 856 km² and reduced significantly to 547.68 km² in the dry season. We observed significant overlap (327.24 km²) in habitat use regions between the wet and dry seasons (Schoener's D = 0.922 and Hellinger's‐based I = 0.991). DEM, proximity to waterholes, and saltlicks were identified as the key variables that contributed to the prediction. We recommend construction of temporal camps in regions of habitat use that are far from the headquarters area for effective management of elephants. Also, an increase in water point's density around the headquarters areas and selected dry areas of the park will further decrease elephant's range and hence a relatively less resource use in monitoring and patrols.     

Milk intake and development in a newborn captive African elephant (Loxodonta africana)

In August 2003 the San Diego Zoo's Wild Animal Park (WAP) and the Lowry Park Zoo, under the auspices of the AZA's Elephant SSP and a USFWS permit, imported 3.8 African elephants (Loxodonta africana) from the Kingdom of Swaziland. When they were captured, transrectal ultrasound examinations revealed that one nulliparous cow was approximately 10 months pregnant. At the time of their arrival (August 2003), all of the animals were estimated to be approximately 13 years old and were thought to be nulliparous. Based on the ultrasound examination results and the average African elephant gestation period, parturition was predicted to be 20 February 2004. In this report, we provide the first detailed data about nursing activity around the clock and newborn calf development, describe maternal and neonatal nighttime activity budgets, and explore maternal weight changes during suckling and lactation. The newborn calf suckled significantly more at night than during the day, but suckled for only about 2 hr per 24‐hr period. Regression analysis revealed that through the first 3 months of life the calf gained 0.385 kg/day while it suckled on a regular basis. We compare our findings with published information on wild elephants, and conclude that although the growth rate is reduced compared to hand‐reared elephant calves, the suckling patterns are almost identical to those reported for wild calves. Zoo Biol 0:1–7, 2005. © 2005 Wiley‐Liss, Inc.     

The use of low-frequency vocalizations in African elephant(Loxodonta africana) reproductive strategies

Fertility-advertisement calls in females are predicted to occur in nonmonogamous species where males and females are widely separated in space. In African elephants, low-frequency vocalizations have thus been suggested as a reproductive strategy used by fertile females to attract mates. This study examined the use of low-frequency vocalizations with respect to different phases of the estrous cycle in African elephants by simultaneously monitoring vocalizations, behavior, and hormonal profiles. Subjects were one male and six female African elephants housed at Disney's Animal Kingdom. No acoustically distinct vocalizations were restricted to the ovulatory follicular phase. However, overall rate of low-frequency vocalization as well as the rate of one acoustically distinct vocalization changed over the estrous cycle, with highest rates of calling related to the first period of follicular growth, or anovulatory follicular phase. Elevated rates of vocalization thus were not restricted to behavioral estrus and occurred much earlier in the estrous cycle than in most species that produce fertility-advertisement calls. Both herd composition and elephant identity also affected rates of vocalization. Vocalizations therefore may not be reliable signals of actual fertility. However, the increase in vocalizations in advance of estrus may attract males to the herd prior to ovulation, facilitating both male-male competition and female choice. Once present in the herd, males may then switch strategies to use more reliable chemical and visual cues to detect ovulating females.     

Relationship between group dynamics and spatial distribution of African elephants in a semi‐arid environment

Group dynamics related to distribution of African elephants in the semi‐arid environment of Tsavo East National Park, Kenya was studied between 2007 and 2010. I examined the seasonal distribution of lone bulls, bull groups, family units and mixed groups and group size. Lone bulls were widely distributed in the dry season and localized in the wet season, whereas bull groups were localized in both seasons. On average bull group size did not differ with seasons or areas, but larger groups were observed in preferred areas in the dry season. Family units were widely distributed in the dry season and localized in the wet season, whereas mixed groups were localized in both seasons. Although family units and mixed groups tended to be large in all areas in the wet season, large groups were also observed in the dry season. Large family units were widely distributed, whereas large mixed groups were localized. The relationship between group dynamics and distribution was attributed to the combination of social, ecological and poaching factors. This study provided insight into distribution patterns that can be applied to the security of the Tsavo elephant population.     

Enzyme‐immunoassay for the measurement of luteinizing hormone in the serum of African elephants (Loxodonta africana)

Circulating patterns of progesterone and luteinizing hormone (LH) in the elephant have been well characterized, and routine monitoring of these hormones is now viewed as a valuable tool for making informed decisions about the reproductive management of elephants in captivity. Currently, LH monitoring in elephants is done with radio‐immunoassays (RIAs); unfortunately, the use of radioactive materials in RIAs limits their application to institutions with laboratory facilities equipped for the storage and disposal of radioactive waste. Enzyme‐immunoassays (EIAs) offer an inexpensive and more zoo‐friendly alternative to RIA. This work reports on an EIA capable of quantifying circulating LH in African elephants. The EIA employs a biotin label and microtiter plates coated with goat anti‐mouse gamma globulin. LH surges in African elephants (n=3) increased fivefold over baseline concentrations (1.00±0.1 ng/ml vs. 0.2±0.1 ng/ml) and occurred 19.3±0.2 days apart. Ovulatory LH surges were associated with an increase in serum progestogens from 4.8±0.4 ng/ml to 11.7±0.4 ng/ml. The ability to quantify reproductive hormones in elephants via EIA is an important step in the process of making endocrine monitoring more accessible to zoos housing these species. Zoo Biol 21:403–408, 2002. © 2002 Wiley‐Liss, Inc.     

Circulating levels of α‐tocopherol and retinol in free‐ranging African elephants (Loxodonta africana)

To date, there are no detailed reports of circulating levels of plasma α‐tocopherol and retinol for large samples of free‐ranging African elephants (Loxodonta africana). This survey study measured natural circulating levels of α‐tocopherol as a measure of vitamin E activity and retinol as an indicator of vitamin A activity, in 70 free‐ranging African elephants captured at Kruger National Park as part of a translocation program. Mean levels of α‐tocopherol and retinol were found to be 0.613 ± 0.271 μg/mL and 0.039± 0.007 μg/mL, respectively, and did not vary significantly across sex or age class. Elephants appear to normally have low circulating levels of both these nutrients compared with domestic herbivore species; values from healthy, free‐ranging elephant populations may provide useful data for assessing nutrient status of captive animals. Zoo Biol 18:319–323, 1999.© 1999 Wiley‐Liss, Inc.     

Estrous synchrony in a group of African elephants (Loxodonta africana) under human care

Synchrony of estrous, and consequently of conception and birth of young, may be of adaptive significance for certain mammals. Among the species in which estrous synchrony has been suspected several times are elephants, but clear evidence is still missing. We determined estrous cycles of African elephants (Loxodonta africana) (n=4) at the Vienna Zoo, Austria, between June 2003 and January 2006 by measuring serum progesterone levels from weekly blood samples. Except for the dominant female when she was intensively lactating, all animals showed clear cycles or progesterone release with a mean period of 105.3+/-15.37 days. For most of the study period, estrous cycles were asynchronous between females. However, after re-occurrence of the progesterone cycle in the dominant female following the first period of lactation, all four females showed high synchrony of progesterone release over the two subsequent cycles. Large changes in individual period lengths indicated that synchronization was due to the adjustment of cycle length in subdominants to that of the dominant female. We used a bootstrap procedure, based on resampling measured times of progesterone peaks, to determine if this apparent synchrony could have been caused by chance alone. This statistical analysis indicated that between-individual variances of the timing of progesterone peaks were much smaller that to be expected by chance (P=0.009). This finding represents the first evidence for estrous synchrony between elephants. We discuss various hypotheses to explain the biological function of cycle synchrony in elephants.     

A survey of elephants (Loxodonta africana) in the Kahuzi-Biega National Park lowland sector and adjacent forest in eastern Zaire

There has been considerable uncertainty about the abundance estimation of Loxodonta africana within tropical lowland moist forests in Zaire. We surveyed a 15,570 km² area within the forests of eastern Zaire using transect sampling methods and estimated the elephant population to be 3720 (range 2300–5000) individuals. Dung pile densities were significantly different between adjacent settlement, deep forest, and deep forest core strata, with the most remote area harbouring the highest density. Evidence of elephant poaching was encountered throughout the survey area suggesting that elephant populations continue to be at severe risk.     

Anthropogenic risk increases night-time activities and associations in African elephants (Loxodonta africana) in the Ruaha-Rungwa ecosystem,Tanzania

Elephants face diverse threats from human activities and use temporal and social strategies to reduce human-induced mortality risk. We used data from camera trap surveys in 2018–2019 (n = 1625 independent detection events from 11,751 sampling days) to investigate elephant responses to anthropogenic risk in the Ruaha-Rungwa ecosystem, Tanzania. The study was conducted in one low-risk and three high-risk sites using 26–40 paired camera trap stations per site. Risk influenced the active periods, use of roads and water sources, social associations and behaviour of elephants. Elephants demonstrated significantly more night-time and reduced daytime activity in the high-risk sites relative to the low-risk site. This higher night-time activity in the high-risk sites was observed for both males and females, though it was more pronounced for cow–calf groups than lone males. Foraging events and use of water sources were more frequent at night in the high-risk sites. Elephants used roads as movement routes in the low-risk site but avoided roads in the high-risk sites. Males were significantly more likely to associate with other males and cow–calf groups in the high-risk sites. Fewer occurrences of relaxed behaviours were observed in the high-risk sites compared to the low-risk site. We discuss the potential implications of our findings for elephant survival and reproduction.     

Landscape-scale habitat response of African elephants shows strong selection for foraging opportunities in a human dominated ecosystem

Balancing trade-offs between foraging and risk factors is a fundamental behavior that structures the spatial distribution of species. For African elephants Loxodonta africana, human pressures from poaching and conflict are primary drivers of species decline, but little is known about how elephants structure their spatial behavior in the face of human occupancy and predation. We seek to understand how elephants balance trade-offs between resource access, human presence and human predatory risk factors (poaching and conflict killing) in an unfenced, dynamic ecosystem where elephants persist primarily outside protected areas in community rangelands. We used tracking data from 101 elephants collected between 2001 and 2016. We investigated elephant behavior in response to landcover, topography, productivity, water, human features and human predation risk using third-order resource selection functions. We extended this analysis by employing a mixed-effects multinomial regression to identify temporal shifts in habitat use, and evaluated temporal shifts in movement patterns by estimating mean squared displacement across different productivity periods. Across periods, elephants displayed strong selection for productive areas and areas near water. Temporal shifts in habitat use showed that, during the dry period, elephants were clustered around permanent water sources where humans also congregated. At the onset of the wet period, a shift occurred where elephants moved away from permanent water and from permanent settlements towards seasonal water sources and seasonal settlements. Our findings indicate that foraging and water access are important limiting factors affecting elephants that potentially restrain their spatial responses to humans at the scale of our analysis. Given that pastoralists and elephants rely on the same resources, increasing human and livestock populations enhance pressure on shared resources and space in Africa's drylands. The long-term conservation of elephants will require approaches that reduce poaching as well as landscape level planning to prevent negative impacts from increasing competition for preferred resources.     

Diurnal variation of salivary cortisol in captive African elephants (Loxodonta africana) under routine management conditions and in relation to a translocation event

The present study assessed the diurnal variation in salivary cortisol in captive African elephants during routine management (baseline) and in relation to a potential stressor (translocation) to evaluate to what extent acute stress may affect diurnal cortisol patterns. Under baseline conditions, we collected morning and afternoon saliva samples of 10 animals (three zoos) on different days in two study periods (n = 3–10 per animal, daytime and period). Under stress conditions, we sampled the transported cow (newcomer) and the two cows of the destination zoo before and after the transport in the morning and afternoon (n = 3–9 per animal, daytime and transport phase), as well as after the first introduction of the newcomer to the bull (n = 1 per animal). Cortisol was measured in unextracted samples by enzyme immunoassay. Under baseline conditions, we observed the expected diurnal variation with higher cortisol levels in the morning than in the afternoon. Under stress conditions, neither a significant difference between pre- and posttransport, nor between morning and afternoon levels was found. The percentage difference between morning and afternoon cortisol after the transport, however, was remarkably lower than before the transport in the newcomer potentially indicating a stress response to familiarization. Saliva samples taken immediately after the introduction of the newcomer to the bull revealed a marked cortisol increase. Our findings indicate that stressors may disturb the diurnal cortisol rhythm. Furthermore, provided that samples can be collected promptly, salivary cortisol is a useful minimally invasive measure of physiological stress in the African elephant.