African Savannah Elephants Research Articles

Development of a new Human-Elephant Coexistence Toolbox for communities living with African savannah elephants (Loxodonta Africana)

This field note is to invite our colleagues to peer review and test a new illustrated Human-Elephant Coexistence (HEC) Toolbox that is being developed in Kenya by Save the Elephants (STE) under the organizations’ mission to secure a future for elephants and to develop a tolerant relationship between humans and elephants. Through presenting the first edition here (Fig. 1), we are inviting our elephant colleagues and community leaders from across the African savannah elephant range States to provide feedback, or any corrections, on the tools, as well as sharing content for additional methods not yet represented. By publishing our process and methods for how we are compiling this encyclopaedia of HEC tools and this novel approach to the peer review process, we hope to provide a transparent process to gauging the validity of the methods presented. This is particularly important because some of the technical advice around the conflict reduction tools presented are not published formally in the scientific literature.

Social integration of translocated wildlife: a case study of rehabilitated and released elephant calves in northern Kenya

Conservation translocations have the potential to strengthen populations of threatened and endangered species, but facilitating integration of translocated individuals with resident populations remains a substantial challenge. Developing functional social relationships like cooperative partnerships or establishing clear dominance hierarchies may be critical to integration of released individuals. Developing such relationships has not received much attention in translocation research, especially for long-lived, socially complex animals for which establishment and navigation of social environments is often a lengthy process that requires sustained monitoring to understand. Here, we present a case study of the social associations of African savannah elephant (Loxodonta africana) calves that have been rehabilitated and released into a fenced wildlife sanctuary in northern Kenya with a resident population of elephants. We use focal follows of interactions pre-release and GPS tracking post-release to quantify social associations of calves with each other and with resident elephants at the release site. We demonstrate how this approach supports translocation monitoring by capturing temporal trends in social patterns within and between release cohorts and among released elephants and wild elephants already resident at the site during a transitional soft release period. Our results show that initial post-release social behavior of rehabilitated calves is related to histories of interaction with familiar individuals and cohort membership and that released calves increased their associations with residents over time. This information provides new behavioral insights for guiding elephant release projects, like the strength of relationships within and among release cohorts, the time to integration with the resident population, and the occurrence and increased incidence of societal fission–fusion. Further, this study provides an example of the utility of animal behavior research to achieve and assess progress towards conservation objectives, and to develop monitoring tools for conservation managers.

Personality and behaviour of three captive African savannah elephants (Loxodonta africana)

This study explores the diversity in individual response for three captive African Savannah Ele-phants (Loxodonta africana) to variations in the daily number of zoo visitors. The nocturnal behav-iour of the elephants was observed in two periods with a factor 5.1 difference in numbers of zoo visitors. Ten behavioural expressions for each elephant were compared for the two periods. Data was collected with surveillance cameras for ten hours (20:00 to 06:00) throughout six nights. Fur-thermore, the data was compared to a similar study on the same elephants performed two years earlier. Both studies revealed a significant difference in behaviour between low activity and high visitor activity periods for all three elephants in the behaviour ’Walk’. No overall significant differ-ence was found in the other behavioural expressions. Clear difference in reaction norms was found between the matriarch and the two subordinates for the behaviours: ’Feed from ground’, ’Inactive’, ’Walk’ and ’Other’. This study shows that elephants in captivity are influenced by an increase in the number of zoo visitors, and that the personality or social status of the elephant may provide them with different resilience to disturbance.

Movement behaviour after birth demonstrates precocial abilities of African savannah elephant, Loxodonta africana, calves

Living in groups has many benefits including predator protection and information sharing that can outweigh the associated costs. African savannah elephants live in a matriarchal fission–fusion social structure where groups merge or split depending on the relative costs of group living. However, elephants are also nonsynchronous breeders which may cause optimal movement patterns to differ between group members, particularly around parturition. Both the mother and the group must assess whether the benefits outweigh the costs of group cohesion. Understanding the impact of gestation, parturition and the presence of a newborn calf on the mother's movement patterns could provide key insights into the drivers of fission–fusion social dynamics. Here, we examined the relationship between reproductive state and movement using GPS tracking data collected during 23 parturitions from multiparous elephants (≥ second calving) and three from primiparous females. We analysed daily mean speed before, during and after parturition, and used a three-state hidden Markov model to determine whether parturition results in a distinctive change in speed. The results show that, although speed decreased on the estimated parturition date, parturition could only be detected in four of 23 multiparous parturition events when placed in the context of speed over a 2-month period. Furthermore, the speed on the day before and the day after parturition (i.e. first complete day of the calf's life) was not significantly different from the yearly mean. By contrast, a speed decrease was detected in the limited sample of primiparous females but was of short duration. Our results indicate that the precocial abilities of calves facilitate the maintenance of group cohesion within the matriarchal fission–fusion social structure of elephants. We speculate that, alongside cognitive development, elephants may have evolved a long gestation period to facilitate an advanced stage of fetal physical development. • We analysed the movement behaviour of parturient African savannah elephants. • Gestation, parturition and neonatal calves had little impact on mothers' movements. • We demonstrate the precocial abilities of elephant calves. • Movement after birth could be a significant advantage for a social species. • Extended gestation of elephants may have evolved for advanced fetal development.

The relationship between distal trunk morphology and object grasping in the African savannah elephant (Loxodonta africana).

BackgroundDuring reach-to-grasp movements, the human hand is preshaped depending on the properties of the object. Preshaping may result from learning, morphology, or motor control variability and can confer a selective advantage on that individual or species. This preshaping ability is known in several mammals (i.e., primates, carnivores and rodents). However, apart from the tongue preshaping of lizards and chameleons, little is known about preshaping of other grasping appendages. In particular, the elephant trunk, a muscular hydrostat, has impressive grasping skills and thus is commonly called a hand. Data on elephant trunk grasping strategies are scarce, and nothing is known about whether elephants preshape their trunk tip according to the properties of their food.MethodsTo determine the influence of food sizes and shapes on the form of the trunk tip, we investigated the morphology of the distal part of the trunk during grasping movements. The influence of food item form on trunk tip shape was quantified in six female African savannah elephants (Loxodonta africana). Three food item types were presented to the elephants (elongated, flat, and cubic), as well as three different sizes of cubic items. A total of 107 ± 10 grips per individual were video recorded, and the related trunk tip shapes were recorded with a 2D geometric morphometric approach.ResultsHalf of the individuals adjusted the shape of the distal part of their trunk according to the object type. Of the three elephants that did not preshape their trunk tip, one was blind and another was subadult.Discussion and perspectivesWe found that elephants preshaped their trunk tip, similar to the preshaping of other species’ hands or paws during reach-to-grasp movements. This preshaping may be influenced by visual feedback and individual learning. To confirm these results, this study could be replicated with a larger sample of elephants.

Impacts of African savannah elephants (Loxodonta africana) on tall trees and their recovery within a small, fenced reserve in South Africa

AbstractAfrican savannah elephants (Loxodonta africana) can have detrimental impacts on trees; studies exploring elephant impact within small, fenced reserves, have lacked focus in determining elephant high‐use areas, the direct effect of fence line and tree recovery. The aim of this study is to assess whether elephants cause significant impact on trees in the small‐fenced Karongwe Private Game Reserve (KPGR) and to determine the levels of tree recovery. Trees ≥5 m in height were surveyed (n = 634 trees). Elephant location data were used to identify high‐ and low‐use areas. Five species accounted for 80% of the records; these were used for further analysis. Trees in high‐use areas were significantly less likely to show signs of debarking and push over. Tall trees were significantly more likely to be impacted by elephants, being associated with heightened risks of debarking and branches being broken but lower risks of being pushed over. Trees close to the fence line were not more impacted than trees near the centre of the reserve. The level of use, distance to the fence and tree height were not significant predictors of tree recovery indicators. Future mitigation efforts should focus on trees with high levels of impact and low levels of recovery.

Comparison of African savanna elephant (Loxodonta africana) fatty acid profiles in whole blood, whole blood dried on blood spot cards, serum, and plasma.

BackgroundAfrican elephants in managed care have presented differences in the balance between omega-3 and omega-6 fatty acids, a situation primarily thought to be due to dietary differences between the managed animals and their free-ranging counterparts. Because of this, circulating fatty acid status is included in routine monitoring of elephant health. A method of blood collection that requires only a few drops of whole blood, dried on filter paper (DBS) and can be used for analyzing full fatty acid profiles offers advantages in clinical application.MethodsThis study compared the use of whole blood, and whole blood DBS, serum or plasma for use in evaluating circulating fatty acid composition in African savannah elephants. Samples from six African elephants (two males and four females) were collected during the same week at the NC Zoo, Asheboro, NC.ResultsResults found only 2 of 36 individual fatty acids and none of the 10 fatty acid groupings were different when comparing the four blood fraction sample types to each other with Mann-Whitney U-Test pairwise comparisons. Myristic acid (14:0) was lower in the DBS samples than in whole blood, serum, and plasma and pentadecaenoic acid (15:1) was slightly more concentrated in DBS and whole blood.DiscussionResults indicate that fatty acid profile of serum, plasma, whole blood, and DBS are comparable in African elephants. The DBS method offers advantages in acquisition and handling and may be preferable to other methods in both routine health assessment of captive animals and field research on free ranging animals.

Social and Seasonal Factors Contribute to Shifts in Male African Elephant (Loxodonta africana) Foraging and Activity Patterns in Kruger National Park, South Africa.

Simple SummaryAfrican savannah elephants are able to greatly modify the vegetation around them through their foraging activities. Accordingly, studying the factors that affect elephant foraging behaviour during different seasons are important to understand their impact on the environment and will also aid in predicting how elephants might react to potential threats such as climate change and land transformation. This article aimed to reinforce current knowledge regarding elephant foraging behaviour by examining how the behaviour is affected across (a) season (wet versus dry); (b) time of day (before or after noon); (c) presence or absence of other elephants; and (d) reproductive state; for six adult elephant bulls monitored in Kruger National Park. Results indicated that elephant foraging behaviour is indeed affected by seasonal and social factors. This highlights how these animals are able to adjust their foraging behaviour during the day to aid in thermoregulation, or during different seasons to fulfil their nutritional requirements. Furthermore, this study opens the door for further research regarding how reproductive activity affects the foraging behaviour of male elephants.African savannah elephants (Loxodonta africana) are well-known as ecosystem engineers with the ability to modify vegetation structure. The present study aimed to examine how male elephant foraging behaviour is affected across (a) season (wet versus dry); (b) time of day (before or after noon); (c) presence or absence of other elephants; and (d) reproductive state (musth versus no musth). Six radio-collared adult elephant bulls were observed twice per week from June 2007–June 2008 in Kruger National Park (KNP), South Africa. Using generalized linear mixed effect modeling, results indicate that elephant bulls graze more during the wet season and browse more during the dry season. To potentially offset the costs associated with thermoregulation during the heat of the day, KNP elephants spent more time foraging during the morning, and more time resting during the afternoon. Male elephants also foraged significantly less when they were associated with females compared to when they were alone or with other males. This is likely due to male–female associations formed mainly for reproductive purposes, thus impeding on male foraging behaviours. In contrast, the condition of musth, defined by the presence of related physical signs, had no significant effect on foraging behaviour.

Movement of Rehabilitated African Elephant Calves Following Soft Release Into a Wildlife Sanctuary

The ability to locate essential resources is a critical step for wildlife translocated into novel environments. Understanding this process of exploration is highly desirable for management that seeks to resettle wildlife, particularly as translocation projects tend to be expensive and have a high potential for failure. African savannah elephants (Loxodonta africana) are very mobile and rely on large areas especially in arid environments, and are translocated for differing management and conservation objectives. Thus, research into how translocated elephants use the landscape when released may both guide elephant managers and be useful for translocations of other species that adjust their movement to social and ecological conditions. In this study, we investigated the movement of eight GPS tracked calves (translocated in three cohorts) following their soft release into a 107 km2 fenced wildlife sanctuary in northern Kenya and compared their movement with that of five tracked wild elephants in the sanctuary. We describe their exploration of the sanctuary, discovery of water points, and activity budgets during the first seven, 14, and 20 months after release. We explored how patterns are affected by time since release, ecological conditions, and social factors. We found that calves visited new areas of the sanctuary and water points during greener periods and earlier post-release. Social context was associated with exploration, with later release and association with wild elephants predictive of visits to new areas. Wild elephants tended to use a greater number of sites per 14-day period than the released calves. Activity budgets determined from hidden Markov models (including the states directed walk, encamped, and meandering) suggested that released calves differed from wild elephants. The first two cohorts of calves spent a significantly greater proportion of time in the directed walk state and a significantly lower proportion of time in the encamped state relative to the wild elephants. Our results represent a step forward in describing the movements of elephant orphan calves released to the wild following a period of profound social disruption when they lost their natal family and were rehabilitated with other orphan calves under human care. We discuss the implications of the elephant behavior we observed for improving release procedures and for defining success benchmarks for translocation projects.

Operant control and call usage learning in African elephants.

Elephants exhibit remarkable vocal plasticity, and case studies reveal that individuals of African savannah (Loxodonta africana) and Asian (Elephas maximus) elephants are capable of vocal production learning. Surprisingly, however, little is known about contextual learning (usage and comprehension learning) in elephant communication. Usage learning can be demonstrated by training animals to vocalize in an arbitrary (cue-triggered) context. Here we show that adult African savannah elephants (n = 13) can vocalize in response to verbal cues, reliably producing social call types such as the low-frequency rumble, trumpets and snorts as well as atypical sounds using various mechanisms, thus displaying compound vocal control. We further show that rumbles emitted upon trainer cues differ significantly in structure from rumbles triggered by social contexts of the same individuals (n = 6). Every form of social learning increases the complexity of a communication system. In elephants, we only poorly understand their vocal learning abilities and the underlying cognitive mechanisms. Among other research, this calls for controlled learning experiments in which the prerequisite is operant/volitional control of vocalizations.This article is part of the theme issue ‘Vocal learning in animals and humans’.

Effects of Male Age and Female Presence on Male Associations in a Large, Polygynous Mammal in Southern India: The Asian Elephant

We present a detailed study of male associations in the Asian elephant, using 6 years of data on identified, non-musth males. Adult males spent greater proportions of their time solitarily than in mixed-sex or in all-male groups. Old (over 30 years) males were sighted more frequently with their age-peers and less frequently with young (15–30 years) males than expected at random in all-male groups. Young males were not sighted more frequently with old males than with young males, and did not disproportionately initiate associations with old males. These results suggest that male associations, in the absence of females, may allow for old non-musth males to test strengths against age-peers. Social learning from older individuals did not seem to be important in male associations, unlike that observed in the African savannah elephant. We also found a constraint on the sizes of all-male groups, similar to that seen in female groups in our study population, and all-male groups were rarer and smaller than those in African savannah elephant. Although male associations were weak, most males had a significant top associate, with whom their association was the strongest, in female absence. In mixed-sex groups, male associations occurred at random, suggesting that males were tracking female groups independently. Differences in male social organization from that of the related African savannah elephant that occupies a similar niche possibly arise from differences in ecology.

Field evidence supporting monitoring of chemical information on pathways by male African elephants

When animals move along well-established pathways, sensory cues along the path may provide valuable information concerning other individuals that have used the same route. Yet the extent to which animals use pathways as sources of public social information is poorly understood. Here we quantified the responses of wild African savannah elephants, Loxodonta africana , to olfactory information along natural elephant pathways, habitual routes that link predictable critical resources in the environment. By monitoring the behaviour of elephants travelling on pathways in a predominantly male study population, we found that elephants were highly olfactorily responsive to pathway substrate. Lone travellers were more responsive than elephants travelling in groups, suggesting elephants without social companions may be more dependent on olfactory cues on pathways during navigation. Furthermore, by experimentally presenting olfactory cues on pathways we found that male African elephants exhibited focused olfactory responses to urine cues of same-sex conspecifics for at least 48 h from time of deposition, and that urine from adult elephants was more likely to elicit vomeronasal system responses than subadult urine. African elephants may therefore potentially be able to discern the age and maturity of individuals they can expect to encounter in the environment from remote urine cues on pathways. We suggest elephant pathways act as a public information resource, assisting navigating elephants via the depositing of urine and dung by previous travellers on the route. These results could help inform elephant management, which may manipulate olfactory information on pathways in high human–wildlife conflict areas, or could use olfactory urine cues to improve the efficiency of corridors that link protected areas for elephants. • Male elephants use the olfactory sense extensively when travelling on pathways. • Those travelling alone are more responsive to pathways than group travellers. • Olfactory responses are made to urine for at least 48 h after it is deposited. • Adult urine elicits vomeronasal system responses more than subadult urine.

Camera trap survey of mammals in the Fazao‐Malfakassa National Park, Togo, West Africa

AbstractFor the first time in Togo, we used camera trapping to investigate the mammal community in Fazao‐Malfakassa National Park (FMNP), a forest and savannah mosaic landscape subject to poaching and other detrimental anthropogenic activity. We compiled a species inventory, that is species occurrence, habitat use and activity patterns during dry and wet seasons. Based on images from >80 locations during 9,007 camera days, we identified 32 mammal species, which, when combined with other published studies, increases the total number of mammals (excluding bats) historically reported to 57 species. Our results confirm the presence of five mammal species evaluated as threatened according to the IUCN Red List, highlighting the significant conservation value of the FMNP. Specifically, it appears to be the only protected area in Togo where the African savannah elephant and the African forest elephant occur sympatrically, and only the second site currently known with direct observations of Walter's duiker. We also report the presence of poachers and other anthropogenic activity within the FMNP. We recommend that continued survey efforts should be combined with detailed ecological data collection, effective law enforcement, community outreach and eco‐tourism development, to safeguard the remaining mammal species communities necessary for a functioning ecosystem in the park.

African forest and savannah elephants treated as separate species

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Importance of old bulls: leaders and followers in collective movements of all-male groups in African savannah elephants (Loxodonta africana)

In long-lived social species, older individuals can provide fitness benefits to their groupmates through the imparting of ecological knowledge. Research in this area has largely focused on females in matrilineal societies where, for example, older female African savannah elephants (Loxodonta africana) are most effective at making decisions crucial to herd survival, and old post-reproductive female resident killer whales (Orcinus orca) lead collective movements in hunting grounds. In contrast, little is known about the role of older males as leaders in long-lived social species. By analysing leadership patterns of all-male African savannah elephant traveling groups along elephant pathways in Makgadikgadi Pans National Park, Botswana, we found that the oldest males were more likely to lead collective movements. Our results challenge the assumption that older male elephants are redundant in the population and raise concerns over the biased removal of old bulls that currently occurs in both legal trophy hunting and illegal poaching. Selective harvesting of older males could have detrimental effects on the wider elephant society through loss of leaders crucial to younger male navigation in unknown, risky environments.

Understanding the drivers of mortality in African savannah elephants.

Populations of African savannah elephants (Loxodonta africana) have been declining due to poaching, human-elephant conflict, and habitat loss. Understanding the causes of these declines could aid in stabilizing elephant populations. We used data from the Great Elephant Census, a 19-country aerial survey of savannah elephants conducted in 2014 and 2015, to examine effects of a suite of variables on elephant mortality. Independent variables included spatially explicit measures of natural processes and human presence as well as country-level socioeconomic measures. Our dependent variable was the carcass ratio, the ratio of dead elephants to live plus dead elephants, which is an index of recent elephant mortality. Carcass ratios are inversely proportional to population growth rates of elephants over the 4 yr prior to a survey. At the scale of survey strata (n=275, median area=1,222 km2 ), we found strong negative associations for carcass ratios with vegetation greenness at the time of the survey, overseas development aid to the country, and distance to the nearest international border. At the scale of ecosystems (n=42, median area=12,085 km2 ), carcass ratios increased with drought frequency and decreased with human density and overseas development aid to the country. Both stratum- and ecosystem-scale models explained well under one-half of the variance in carcass ratios. The differences in results between scales suggest that the drivers of mortality may be scale-specific and that the corresponding solutions may vary by scale as well.

The African Elephant (<i>Loxodonta africana</i>): Mini-Review of an Endangered Species

Humankind is responsible for conditioning the distribution of species, including the African elephant. From its great physical form to its fascinating social behaviour, Loxodonta africana can trigger an admiration in some and be undervalued by others, often ending up being a victim of human activities. Increasingly, the apparent impossibility of the coexistence of these two species represents a problem for the conservation of the large African mammal. Here it was reviewed the published bibliography about the African savannah elephant (L. africana), between July 2015 and December 2019. The Web of Science (WoS) database enabled the search for the words “Loxodonta africana”. From this research, we obtained 310 articles, 149 of them were analysed. Elephants have characteristics that define its uniqueness that are constantly reported, such as: its cognition and extraordinary memory, factors that influence its movements and its distribution, its specific vocalization and the strong dependence on social connections. Threats to the conservation of the species and the urgency of interventions are also an important subject of discussion. Poaching for ivory represents a major obstacle on conservation and has been drastically reducing the number of elephant populations and causing future consequences for those who thrive. Law enforcement in reducing/prohibit poaching has proven to be insufficient, so there is an urgent need to combat the demand for ivory. Also, it is necessary to bypass the conflict, promote tolerance and enable cohabitation. We believe that educating human populations to better understand the species may be the difference to guarantee a future for the species. Conservationists may be the enablers of these educational actions, however, to make the difference, it is necessary to know the species in detail. Mastering knowledge in these behavioural areas will contribute to improve mitigation strategies, which could make the difference to guarantee a future for the species.

Factors affecting detection probability of elephant (Loxodonta africana) carcasses in an African conservation area

Poaching for ivory has led to massive population declines of African savannah elephants (Loxodonta africana). To prioritize anti-poaching efforts, elephant poaching hotspot maps were created. However, these might be biased because they are not corrected for the detection probability of elephant carcasses. Carcass decomposition state was defined as a proxy for detection probability. Detection probability was influenced by the Normalized Difference Vegetation Index (NDVI) influencing live elephant densities across the area. Our results show the importance of accounting for detection probability to achieve true estimates when predicting poaching risk to successfully direct anti-poaching efforts in African conservation areas.

Los objetos de adorno en marfil de proboscídeos del hipogeo de Padru Jossu (Sanluri, Cerdeña, Italia) y el Campaniforme mediterráneo

In the present work, we examine the personal adornment in proboscidean ivory from the Bell Beaker period at the hypogeum of Padru Jossu, Sanluri (Sardinia, Italy) currently preserved in the Museo Civico Archeologico Villa Abbas of Sardara. For the first time, a complete study –morphological, use wear and archaeometric– of this material has been conducted. The typological study established two categories: buttons and pins. Those categories were also subdivided into three groups respectively. Technological and functional analyses were made difficult by the strong degradation of the items and the presence of glue and varnish. The archaeometric study highlighted the diverse provenances of the proboscidean ivories, suggesting a chronological difference in the geographical sources, as well as in the mobility patterns implicit in the movements of the raw material. The ivory from the older Stratum iii is predominantly from the Asian elephant, and in the later Stratum ii the exclusive supplier species is the African Savannah elephant. It is also important to mention that in the ensemble from Stratum iii, one of the items seems related to the Eastern types of ossi a globuli, linking this Asian ivory with an Aegean and Oriental axis of mobility.

Incipient signs of genetic differentiation among African elephant populations in fragmenting miombo ecosystems in south‐western Tanzania

AbstractHabitat fragmentation can play a major role in the reduction of genetic diversity among wildlife populations. The Ruaha‐Rungwa and Katavi‐Rukwa ecosystems in south‐western Tanzania comprise one of the world's largest remaining African savannah elephant metapopulations but are increasingly threatened by loss of connectivity and poaching for ivory. To investigate the genetic structure of populations, we compared the genotypes for nine microsatellite loci in the western, central and eastern populations. We found evidence of genetic differentiation among the three populations, but the levels were low and mostly concerned the younger cohort, suggesting recent divergence probably resulting from habitat loss between the two ecosystems. We identified weak isolation by distance, suggesting higher gene flow among individuals located less than 50 km apart. In a long‐lived species with overlapping generations, it takes a long time to develop genetic substructure even when there are substantial obstacles to migration. Thus, in these recently fragmented populations, inbreeding (and the loss of heterozygosity) may be less of an immediate concern than the loss of adults due to illegal hunting.