Multimodal communication and audience directedness in the greeting behaviour of semi-captive African savannah elephants.

Many species communicate by combining signals into multimodal combinations. Elephants live in multi-level societies where individuals regularly separate and reunite. Upon reunion, elephants often engage in elaborate greeting rituals, where they use vocalisations and body acts produced with different body parts and of various sensory modalities (e.g., audible, tactile). However, whether these body acts represent communicative gestures and whether elephants combine vocalisations and gestures during greeting is still unknown. Here we use separation-reunion events to explore the greeting behaviour of semi-captive elephants (Loxodonta africana). We investigate whether elephants use silent-visual, audible, and tactile gestures directing them at their audience based on their state of visual attention and how they combine these gestures with vocalisations during greeting. We show that elephants select gesture modality appropriately according to their audience's visual attention, suggesting evidence of first-order intentional communicative use. We further show that elephants integrate vocalisations and gestures into different combinations and orders. The most frequent combination consists of rumble vocalisations with ear-flapping gestures, used most often between females. By showing that a species evolutionarily distant to our own primate lineage shows sensitivity to their audience's visual attention in their gesturing and combines gestures with vocalisations, our study advances our understanding of the emergence of first-order intentionality and multimodal communication across taxa.

Differences in Mother-Infant Bond and Social Behavior of African Elephant Calves Living In Situ and Ex Situ.

African zoo elephants live in safe environments with sufficient resources, are protected from threats, and have their health and body conditions cared for. Calves ex situ undergo the same developmental stages as in situ and are raised by the whole family unit. However, due to environmental differences, there might be behavioral modifications between calves in situ and ex situ. We hypothesize that these differences increase with ongoing generations. This ethological study compares social and general behavior and the distance calves kept to their mothers' between calves of the first (F1) and second (F2) zoo generation and the wild. Using ethological methods, data were collected for ~90 in situ calves and 16 ex situ (8 F1, 8 F2) between the ages of 0.5 to 4 years (120 observation hours per group). Results showed that in situ calves spent significantly more time close to mothers than the F1 and the F2 zoo generations (F1/in situ: p = <0.001; F2/in situ: p = 0.007). The behaviors of eating, drinking, trunk movement, washing, and affiliative behaviors showed significant differences between in situ and ex situ calves. The amount and distribution of affiliative and agonistic behavior initiated and received by calves was displayed with a greater variety ex situ. Ex situ calves not only performed affiliative but, in contrast to the in situ, also agonistic behavior (F1/in situ: initiated p = 0.002, received p = 0.010; F2/in situ: initiated p = 0.050, received p = 0.037). The comparison of zoo generations suggests that differences did not increase with the generation. The more casual binding between mothers and offspring in zoos and the age-dependent improvement of social behavior of zoo-born calves are seen as a result of elephants' adaptation to secure zoo conditions. The results of this study agree with the faster development of ex situ African elephants, like earlier puberty and more frequent breeding patterns, as known from the literature.

Behaviour and Welfare Impacts of Releasing Elephants from Overnight Tethers: A Zimbabwean Case Study.

Within the southern African elephant tourism industry, chaining or tethering elephants is still a relatively routine practice, despite the known negative impacts. Cited reasons for chaining include fear of aggressive interactions between elephants when handlers are absent, or a general increase in expression of aggressive behaviours (both to other elephants and to their human handlers). In Zimbabwe, concerns expressed include the danger of elephants escaping and entering human-inhabited areas. Four male semi-captive elephants at a Zimbabwe tourist facility were taken off overnight (~12 h) tethers and were placed in small pens ('bomas'), approximate sizes from 110 m2 to 310 m2), as part of a strategy to improve elephant welfare. Behavioural data were collected from overnight videos from December 2019 to March 2020, between 18:00 to 06:00, using focal, instantaneous sampling (5-min interval). Data were collected for three nights at three time periods: (i) Tethered; (ii) approximately four weeks post-release; (iii) approximately eight weeks post-release. Behavioural change over these time points was analysed using general linear models with quasibinomial error structures. Behavioural changes indicative of improved welfare were observed following these management changes, and no significant increases in aggression were observed either between elephants, or towards their human handlers. Proportion of time engaging in lying rest was higher in the first month after release from tethering (mean ± SD, 50 ± 14%) than when elephants were tethered (20 ± 18%) (p < 0.05). Additionally, although not statistically significant, stereotypies were reduced when elephants were no longer tethered (4 ± 6% observations tethered compared to 2 ± 2% off tethers), and positive social behaviour also increased (1 ± 1% on tethers, 2 ± 2% off tethers), with the greatest improvements seen in the pair-housed elephants. To improve elephant welfare in southern African tourism facilities we strongly advocate that less restrictive management practices which enable greater choice and freedom of movement overnight are implemented.