Multi-level fission-fusion societies, characteristic of a number of large brained mammal species including some primates, cetaceans and elephants, are among the most complex and cognitively demanding animal social systems. Many free-ranging populations of these highly social mammals already face severe human disturbance, which is set to accelerate with projected anthropogenic environmental change. Despite this, our understanding of how such disruption affects core aspects of social functioning is still very limited. We now use novel playback experiments to assess decision-making abilities integral to operating successfully within complex societies, and provide the first systematic evidence that fundamental social skills may be significantly impaired by anthropogenic disruption. African elephants (Loxodonta africana) that had experienced separation from family members and translocation during culling operations decades previously performed poorly on systematic tests of their social knowledge, failing to distinguish between callers on the basis of social familiarity. Moreover, elephants from the disrupted population showed no evidence of discriminating between callers when age-related cues simulated individuals on an increasing scale of social dominance, in sharp contrast to the undisturbed population where this core social ability was well developed. Key decision-making abilities that are fundamental to living in complex societies could be significantly altered in the long-term through exposure to severely disruptive events (e.g. culling and translocation). There is an assumption that wildlife responds to increasing pressure from human societies only in terms of demography, however our study demonstrates that the effects may be considerably more pervasive. These findings highlight the potential long-term negative consequences of acute social disruption in cognitively advanced species that live in close-knit kin-based societies, and alter our perspective on the health and functioning of populations that have been subjected to anthropogenic disturbance.

Lose the tusks: Harvest and poaching of wildlife have increased as the human population and our technology have grown. These pressures now occur on such a scale that they can be considered selective drivers. Campbell-Staton et al . show that this phenomenon has occurred in African elephants, which are poached for their ivory, during the 20-year Mozambican civil war (see the Perspective by Darimont and Pelletier). In response to heavy poaching by armed forces, African elephant populations in Gorongosa National Park declined by 90%. As the population recovered after the war, a relatively large proportion of females were born tuskless. Further exploration revealed this trait to be sex linked and related to specific genes that generated a tuskless phenotype more likely to survive in the face of poaching. —SNV

This short paper is intended to alert our colleagues to the existence of The Elephant Ethogram: A Library of African Elephant Behaviour. It describes its purpose, form and scope, and appeals for contributions of undocumented, rare, novel or cultural Loxodonta africana behaviour. We do not present descriptions of behaviours, methodologies, results or discussion; these may be found online within The Elephant Ethogram. The Elephant Ethogram is an ElephantVoices initiative to document the complex, diverse and nuanced repertoire of behaviour and communication of African savannah elephants (L. africana). In a unique, user-friendly, fully searchable and publicly accessible database, The Elephant Ethogram chronicles the rich postural, gestural, tactile, chemical and acoustic communication and behaviour of Africa's savannah elephants. It includes commonly displayed, unusual, novel and culturally learned behaviours, as well as those expressed in response to people. The Elephant Ethogram is based on published descriptions of behaviour and the decades of behavioural studies and photographic, acoustic and video-graphic collections from Amboseli National Park (NP), the Maasai Mara ecosystem (Mara), Kenya, and Gorongosa National Park, Mozambique conducted by ElephantVoices. It is also built from behaviour captured for documentaries filmed in the Mara and Gorongosa and video clips of unusual behaviour collected by colleagues and members of the general public. The Elephant Ethogram replaces ElephantVoices' online Elephant Gestures and Elephant Calls Databases originally developed in 2003 and revamped in 2011 (Poole 2011; Poole and Granli 2011), that were based on our elephant studies in Amboseli National Park, Kenya between 1975 and 2009 (Poole 1987; Poole et al. 1988; Poole 1989a; Poole 1989b; Poole et al. 2005) and the work of other scientists (Douglas-Hamilton 1972; Berg 1983; Moss 1983; Kahl and Armstrong 2000; Kahl and Armstrong 2002). Between 2011 and 2019 we carried out elephant behaviour studies and conservation projects in the Mara, and Gorongosa NP, during which we completed elephant, field notes, images and videos of the behaviour of known individuals. Furthermore, in a ground-breaking collaboration with copyright owners Off the Fence, Gorongosa Media Project and Bob Poole Films, hundreds of hours of raw footage of elephants, originally shot for documentaries in Gorongosa and the Mara, were granted to ElephantVoices for use in science and education. Since we collaborated with the filmmakers on site, the footage primarily depicts known individuals. In 2020 we collected additional footage of behaviour in Amboseli National Park. The Elephant Ethogram combines and significantly improves the structure and functionality of the original databases, includes hundreds of additional behaviours, 2,400 annotated video clip examples from three populations, higher-resolution images, additional audio files. Elephant behaviour has been documented by hunters, naturalists and scientists for hundreds of years starting with the earliest scholarly notes of Aristotle (1862 translation) to those of Darwin (1872) to Kühme’s (1962; 1963) research on captive African elephants. The study of free-ranging elephants by Douglas-Hamilton (1972) stimulated the work of many who followed, including the five decades-long work of Moss and her colleagues, and our own. From hundreds of published studies, we know that savannah elephants show great richness, variation and flexibility in their behaviour. Some of these publications have described elephant behaviours relevant to the aims of their particular study (Moss 1983 (oestrus); Poole 1987 (musth); O’Connell-Rodwell et al. 2011 (male-male relationships); Goldenberg and Wittemyer 2020 (death)), but only Kahl and Armstrong’s work in Hwange National Park, Zimbabwe (2000; 2002) and our own in Amboseli National Park (Poole 2011; Poole and Granli 2011) aimed to document the full repertoire of behaviour of the species. We worked closely with Kahl to share data and to agree on terminology and definitions. Until his untimely death in 2012 Kahl’s plan was to publish a detailed elephant ethogram. The construction of “exclusive ethograms” to describe a species' behaviour or activity patterns is commonly used in behavioural studies, where the ethogram focuses on the behaviours of interest. It is more unusual to find catalogues that attempt to produce an “exhaustive” ethogram of all known behaviours of a given species. One example is the work of Nishida et al. (1999) on chimpanzees, Pan troglodytes, which aims to provide an exhaustive list of behaviours. This body of work defines 515 behaviours, recording whether they were idiosyncratic, limited to a small group, to one population or were, to a greater or lesser extent, cross-cultural. Another example is the work of Bolgan et al. (2014) on the Arctic charr, Salvelinus alpinus, a fish species. A draw-back of these studies is that they are non-searchable, written documents without video-graphic documentation. Nishida et al. (2010) solved this problem for their study of chimpanzees by publishing a book with an accompanying CD. Very few studies; however, have attempted to produce an exhaustive, searchable web-based ethogram of a species with video examples of behaviour. One example is Mouse Ethogram: An Ethogram for the Laboratory Mouse developed in the Stanford Medical School in the Laboratory of Joseph Garner. Since this study was carried out in a captive environment it is unlikely to be exhaustive for the species. African elephants (Savannah and Forest) are among the most socially complex non-human species (Moss and Poole 1983; McComb et al. 2000; Archie et al. 2005; Wittemyer et al. 2005), as well as one of the more heavily exploited (Meredith 2001; Wittemyer et al. 2014). As scientists continue to document their extraordinary behaviour, elephants are increasingly impacted by humans to...

The social complexity hypothesis (SCH) for communication states that the range and frequency of social interactions drive the evolution of complex communication systems. Surprisingly, few studies have empirically tested the SHC for vocal communication systems. Filling this gap is important because a co-evolutionary runaway process between social and vocal complexity may have shaped the most intricate communication system, human language. We here propose the African elephant Loxodonta spec. as an excellent study system to investigate the relationships between social and vocal complexity. We review how the distinct differences in social complexity between the two species of African elephants, the forest elephant L. cyclotis and the savanna elephant L. africana, relate to repertoire size and structure, as well as complex communication skills in the two species, such as call combination or intentional formant modulation including the trunk. Our findings suggest that Loxodonta may contradict the SCH, as well as other factors put forth to explain patterns of vocal complexity across species. We propose that life history traits, a factor that has gained little attention as a driver of vocal complexity, and the extensive parental care associated with a uniquely low and slow reproductive rate, may have led to the emergence of pronounced vocal complexity in the forest elephant despite their less complex social system compared to the savanna elephant. Conclusions must be drawn cautiously, however. A better understanding of vocal complexity in the genus Loxodonta will depend on continuing advancements in remote data collection technologies to overcome the challenges of observing forest elephants in their dense rainforest habitat, as well as the availability of directly comparable data and methods, quantifying both structural and contextual variability in the production of rumbles and other vocalizations in both species of African elephants.

Vocalizations often vary in structure within a species, from the individual to population level. Vocal differences among social groups and populations can provide insight into biological processes such as vocal learning and evolutionary divergence, with important conservation implications. As vocal learners of conservation concern, intraspecific vocal variation is of particular interest in elephants. We recorded calls from individuals in multiple, wild elephant social groups in two distinct Kenyan populations. We used machine learning to investigate vocal differentiation among individual callers, core groups, bond groups (collections of core groups) and populations. We found clear evidence for vocal distinctiveness at the individual and population level, and evidence for much subtler vocal differences among social groups. Social group membership was a better predictor of call similarity than genetic relatedness, suggesting that subtle vocal differences among social groups may be learned. Vocal divergence among populations and social groups has conservation implications for the effects of social disruption and translocation of elephants.

Monitoring populations of endangered species is critical to understanding the threats they face and to managing interventions to ensure their long-term survival. Individual recognition further allows for essential studies of life history, home range, population dynamics, social behaviour, and photographic capture-recapture, all of which can make conservation interventions more effective. African elephants (Loxodonta Africana, L. cyclotis), endangered keystone species, have been subjects of several long-term studies involving individual recognition, yielding critical knowledge for their conservation. Ongoing concern for elephant survival has also led to increased interest in engaging non-scientists in monitoring populations. Tools are, therefore, needed to easily reidentify individual elephants and to provide user-friendly collection, upload, analysis and sharing of data. We describe the development of the Elephants Who’s Who & Whereabouts, a relational web-based database (DB) system, for registering, reidentifying and monitoring elephants. We tailored the DB for studying elephants in two populations: Maasai Mara ecosystem, Kenya, and Gorongosa National Park, Mozambique. The main components of the system are: a searchable database (Who’s Who) of registered adult elephants with admin and user interfaces; a searchable database of observations (Whereabouts) including elephant sightings, signs, sick and wounded individuals, and mortalities, with admin and user interfaces; a user interface (My Observations) for entering and edit of data; a Smartphone application (EleApp) permitting collection and upload of data; a searchable Google-Earth-based Mapping interface with export functionality; a Features Guide (photographs and text describing how to identify elephants); a User Info. We describe the system’s structure, functionality, and ease of use. La surveillance des populations d’espèces menacées est essentielle à la compréhension des dangers qu’elles rencontrent et pour la gestion des interventions mises en place afin d’assurer leur survie à long terme. En outre, la reconnaissance individuelle ouvre la voie à une analyse nécessaire du cycle biologique, du domaine vital, de la dynamique des populations, du comportement social et de la capture-recapture photographique, autant d’éléments qui favorisent une meilleure efficacité des interventions. Les éléphants d’Afrique (Loxodonta Africana, L. cyclotis), espèce clé en voie de disparition, ont fait l’objet de plusieurs études de long-terme intégrant la technique de reconnaissance individuelle, qui ont constitué une base de connaissance indispensable à leur protection. Le sujet de la survie des éléphants suscite également un intérêt accru pour l’implication de personnes non-scientifiques dans la surveillance des populations. Des outils sont donc nécessaires pour une nouvelle identification des éléphants ainsi que pour fournir un environnement facile d’utilisation pour la collecte, le chargement, l’analyse et le partage des données. Nous décrivons ici le développement d’un système de base de données relationnelle en ligne (DB), « Who’s Who & Whereabouts » (« Qui est qui et où vont-ils ? ») pour la surveillance, la reconnaissance et le recensement des éléphants. Nous avons configuré la base de données en deux catégories de population : l’écosystème Maasai Mara au Kenya et le parc national de Gorongosa au Mozambique. Les principaux éléments de cet outil sont les suivants : deux bases de données consultables avec interface administrateur et utilisateur, l’une répertoriant les éléphants adultes (Who’s who), l’autre axée sur les observations d’individus, de leurs traces, des sujets malades, blessés, ou morts (Whereabouts). On trouve également une interface utilisateur, « My Observations » (« Mes observations »), permettant la saisie et la modification de données, une application sur smartphone (« EleApp ») pour la collecte et le chargement de données, une interface cartographique (« Mapping ») via Google Earth avec possibilité d’export, un guide des caractéristiques (« Feature Guide ») avec photos et textes expliquant comment identifier les éléphants et une information utilisateur (« User Info »). Nous détaillons ici la structure du système, ses fonctionnalités et sa simplicité d’utilisation.