Interspecific Information Transfer as a Driver of Community Structure

Lead Research Organisation: University of York
Department Name: Biology


What drives the abundance and distribution of animal species in space and time? This central question in ecology and conservation has so far been approached mainly by investigating the impact of predator-prey relationships and competition for limited resources such as food. However, it is becoming increasingly clear that animals may also have substantial effects on each other by acting as critical sources of information, for example when they communicate the presence of a predator or the location of food. The potential importance of information exchange between species is evident from the widespread occurrence of mixed-species groups in nature, in taxa ranging from spiders and fish to birds and mammals. Yet, currently almost nothing is known about how the advantage of living next to valuable informants affects patterns in social attraction between species.

Our study addresses this question from an integrated theoretical and empirical angle. We will build a general model to predict how information benefits shape the composition of mixed-species groups in nature. Our model will take into account the key costs and benefits from group life: (i) the information benefits from joining many eyes, ears and noses, (ii) the benefit of being close to others that may be eaten instead of you when predators attack, and (iii) the costs from increased competition over limited resources. To test our model in the real world, we focus on the mixed-species groups of herbivores dominating the African savannas in a field study. The many eyes, ears and noses in these herds are known to result in all-important information benefits when alarm signals are emitted. To determine how patterns in social attraction between species in this system depends on their value as informants, we use predator simulations and playback experiments to determine (i) the information contained in the alarm signals from each species in the community and (ii) to what extent this information is transferred between species. We will also obtain measures of the vulnerability of each herbivore species to the various predators in the community as well as costs from food competition when diets overlap. On this basis, we will be able to use social network analysis to test the role of communication (relative to other costs and benefits) as a driver of group formation between species. The savannah herbivore study will thus allow us to address fundamental questions in biology by (i) revealing which species within the community group together and why, and (ii) establishing the nature of coexistence between species: are multispecies groups mutually beneficial or do they rather form when one species parasitizes on the information produced by another?

The project will provide novel insights into the basic links between species in the natural environment by integrating communication benefits into classical food-web-based models of ecosystem structure. By establishing new fundamental principles that shape the social associations between species, the study will bring a deeper understanding of ecosystem dynamics that can be critical for identifying conservation priorities. Our conceptual framework will allow conservationists to assess when declines in key species are likely to have repercussions throughout the ecosystem by affecting the survival of others, and when these indirect effects are likely to become a conservation concern for endangered species.

Planned Impact

We expect the work to benefit the following key end-users, nationally and internationally:
ACADEMIC COMMUNITY: A new conceptual framework that explicitly incorporates both social and trophic connections in models of community structure and resilience will advance the knowledge base for a broad range of academic disciplines, from Animal Behaviour and Evolutionary Biology to Ecology to Biomodeling. Specifically, the project will also provide high quality postdoctoral training for an accomplished Dphil-student by furthering his expertise in social network analysis, mathematical modelling, acoustic analysis and ecological fieldwork.
CONSERVATION MANAGERS: The development of a multi-layered network approach to account for interspecific interactions will provide a conservation tool to support the effectiveness of ecosystem management by allowing the consequences of species declines for the population viability of coexisting species to be assessed. In particular the impact of communication on population performance is relevant for managers of social wildlife species, notably mammals and birds, which include some of the most highly threatened taxa and therefore can be susceptible to even modest changes in ecological processes. Specifically, in collaboration with our project partner Kenya Wildlife Service we are organizing a conservation management workshop for stakeholders in conservation of the Mara ecosystem. This provides an urgently needed forum for addressing the extremely serious management issues that faces the region in these years. Drastic declines in wildlife populations have been documented due to intensifying illegal livestock grazing and other activities. The theme of the meeting will be the importance of interspecific interactions for ecosystem functioning and conservation, including wildlife-livestock conflicts. The main aim is to inform and influence the regional management practices of one of the world's most unique wildlife areas, a UNESCO World Heritage Site, at a critical time. Technologically, the project will specifically benefit managers of rangelands by groundtruthing satellite-derived vegetation indexes to assess their usefulness for quantifying sward characteristics in grassland systems.
ECOTOURISM INDUSTRY: The tourism sector is the 2nd largest source of foreign exchange revenue in Kenya, UK residents being the most frequent visitors. Wildlife is the main attraction and Masai Mara, the proposed study site, is by far the most visited park, receiving around 300,000 foreign visitors anually. By informing the management of savanna ecosystems in general and Masai Mara in particular, the project will support a wide range of stakeholders in the tourist industry. These include poor local households whose income relies on tourism directly or indirectly, companies in the tourist trade, both in Africa and worldwide, and the tourists themselves. The project will also enrich the tourist product by uncovering new facts about the behavioural ecology of African wildlife.
ANIMAL WELFARE: The project will benefit management of captive mammals by showing when anxiety is likely to transmit between species and welfare may be improved by separation (eg during veterinary interventions).
GENERAL PUBLIC: The large mammals of the African savanna hold a special fascination for the general public, and we will take advantage of this to improve the public understanding of environmental issues and science in general. The public will benefit from a deeper understanding of the interconnectedness of species in natural systems and increased societal awareness of the importance of sustainable use of natural resources.
SCHOOLS: Again, the study system provides an exceptional opportunity to engage school children in environmental science. We seek to stimulate pupils and students who are at critical points in their education with the aim of increasing their environmental awareness and potentially inspiring them to pursue a science career.


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Description This project has expanded our knowledge on how environmental, ecological and social factors interact to drive the formation of mixed-species groups in nature, thereby providing a deeper understanding of the structure and function of ecological communities which is relevant for natural resource management, in particular for biodiversity conservation. The insights were obtained by combining the construction of a general theoretical model with a large-scale field study on the African savannah herbivore community. We have already published findings showing that the responsiveness of African savannah herbivores to each other's alarm calls depends on both adaptive and non-adaptive processes. Specifically, responses are stronger to the species who are most consistent in denoting the presence of predators relevant to the receiver and in risky environments. Stronger responses to alarm calls that resemble the conspecific alarm call suggest sensory constraints on the ability to decode alarm calls. That opportunities of learning alarm calls matter as well is suggested by stronger responses to more abundant callers. Several of our main findings relating to community level processes are currently being submitted for publication and are expected to be publicly available soon. New areas for research identified include: - The need to understand how the increasing impact of human activities on natural environments affects the performance of wildlife populations and their evolutionary dynamics because of alterations in social relations between species, and - the need to understand the cognitive mechanisms underlying the responses to alarm calls of other species as these will determine the ability of species to adjust to human-induced changes in the species composition of communities. In addition to school outreach, we organized forums that promoted the role of ecological research in conservation, both globally and focused on the wider study area, the Mara region of Kenya, which is a top conservation priority globally. Thus, in 2017, we organized and hosted the workshop "Future research priorities for the conservation management of the Mara" in Kenya. The workshop brought together researchers and conservation managers and followed a structured format to identify and rank research priorities for the Mara ecosystem, both in terms of importance and urgency. Top priorities included research into habitat fragmentation by fencing and settlement, planning of wildlife corridors, livestock management and disease transmission. We disseminated the findings in a popular article in SWARA, the leading conservation magazine in East Africa. In 2018, we hosted a two-day NERC/ZSL symposium "Linking behaviour to populations and communities: how can behavioural ecology inform conservation?" at the Zoological Society of London with talks by 22 experts from around the globe. As an outcome of the meeting, we are currently editing a special issue of the Philosophical Transactions of the Royal Society B (planned publication 2019). The project also allowed JBJ to secure follow-on funding from the International Opportunities Fund (NERC) to continue research on the same system with the project "Parasite transmission at the wildlife-livestock interface and its implications for management - a pilot study of an African savannah ecosystem" in collaboration with Kenya Wildlife Service, Moredun Research Institute and Queens University Belfast.
Exploitation Route The results can be used to inform conservation efforts and used by academics studying mixed-species groups and community structure.
Sectors Environment,Other

Title EIDC database for NE/L007185/1 
Description Data collected during the grant NE/L007185/1. This dataset contains information about various aspects of the alarm communication network of African savannah herbivores. Data were collected in April 2015 and between September 2015 and October 2016 in the Masai Mara National Reserve, in southern Kenya (1°30'S, 35°10'E). Research focused on the 12 most common herbivore species in the ecosystem. For each of these species, the dataset provides information on vigilance rates, the probability to alarm call in response to different predators, the responsiveness to heterospecific alarm calls, as well as the relative abundance and grouping behaviour of species. Details on study species, study design and data analysis are provided in the supporting documentation. 
Type Of Material Database/Collection of data 
Year Produced 2019 
Provided To Others? No  
Impact The publications published, submitted and in preparation from this study draw on this database. 
Title Multispecies grouping model (in progress) 
Description This stochastic model examines the costs and benefits of two species in grouping together and with their own species (and a mix). The model examines the effect of differences between the species in predation risk and information value (vigilance and contribution to dilution) and resource overlap competition and predicts the optimal mix of any two species. 
Type Of Material Computer model/algorithm 
Provided To Others? No  
Impact Modelling in progress 
Title Savanna herbivore database (in progress) 
Description Data in this database is accommulating as fieldwork progresses. It consists of outcomes of playback experiments, predator simulation experiments, species counts, vegetation sampling, vigilance recording and leader-follower interactions as described in the proposal. 
Type Of Material Database/Collection of data 
Provided To Others? No  
Impact Database is under construction. 
Description Kenya Wildlife Service 
Organisation Kenya Wildlife Service
Country Kenya 
Sector Private 
PI Contribution Kenya Wildlife Service (KWS) is responsible for management of all wildlife in Kenya, and we are currently collecting data on interspecific interactions between savannah herbivores in order to establish the mutual dependency of population performance between species. The data will constitute the basis for derivation of a general model.
Collaborator Contribution We have had several meetings with representatives from KWS both at the HQ in Nairobi and on site in Masai Mara National Reserve, during which we have had constructive discusssions about the design of the project and the format of a future collaborative workshop. KWS have been supportive in providing approvals for research clearance.
Impact N/A
Start Year 2014
Description Conference talk by Kristine Meise: Multiple processes determine responsiveness to heterospecific alarm calls in African savanna herbivores 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact Conference of the Ethological Society in Potsdam
Year(s) Of Engagement Activity 2018
Description Guide training (Kenya) 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact Capacity-building by project staff during the fieldwork phases: delivery of biweekly training sessions for safari guides in Kenya's tourist industry.
Year(s) Of Engagement Activity 2014,2015,2016,2017,2018
Description Hosting workshop "Future research priorities for the conservation management of the Mara" for research directors, senior conservation managers and others in Masai Mara National Reserve, Narok Kenya, September 2017. 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact The workshop was hosted successfully as planned on Friday 22 September 2017 at Fig Tree Camp in the Maasai Mara National Reserve. The around 20 participants included research directors and managers of the protected area in the region, including the chief park warden of the Maasai Mara National Reserve itself. The exercise resulted in a ranked priority list of 30 research questions for the conservation management of the Mara region. The list has been made available to protected area management and researchers working in the area. A more detailed description of the workshop is available in the attached manuscript which has been submitted for publication the January 2018 issue of SWARA, the leading conservation magazine in East Africa, published by the East African Wildlife Society. Participants generally expressed a deep wish for the workshop forum for communication between managers and researchers to become an annual event, emphasising that the consensus approach would strengthen their voice in trying reach local and national government.
The ranked priority list of research questions are as follows:
How does fencing and human settlement impact habitat fragmentation and thereby wildlife connectivity, and which species are most affected?
Where should corridors and new conservancies be prioritized?
What is the extent and number of livestock encroachment into the park?
What is the effect of livestock management of forage quality?
What is the impact of tourist camps and associated structures and activities in key habitats, notably by rivers, on different aspects of ecosystem health?
Which road sections would have the biggest impacts on animal mortality?
Does an increase in conservancies cause an increase in livestock grazing within the park?
Which would be more beneficial, reducing numbers and improving quality of livestock or managing grazing?
What is the relationship between livestock disease and transmission to wildlife?
What is the prevalence and drivers of wildlife poisonings?
What is the spatial pattern in decline of woody vegetation caused by human activities?
How does off-road driving relate to sward degradation?
What are the short/long-term costs and benefits to people of building fences?
What is the impact of climate change on biodiversity and species distributions in plants, and how does this impact resource competition in herbivores?
How are future landuse changes in terms of deforestation and agriculture within the ecosystem going to affect water flow in rivers and thereby ecosystem health?
What is the impact of the various aspects of ballooning on different wildlife species?
What is the effect of climate change on herbivore migration patterns and their secondary effect on human-wildlife conflict?
How do different intensities and practices of game-viewing affect the vital rates of key species such as threatened predators, the black rhino and crossing wildebeest?
What is the economic value from tourism of different aspects of biodiversity?
What are the consequences of snaring for population dynamics of affected species?
What is the effectiveness of community consolation schemes in reducing human-wildlife conflict?
How do socioeconomic changes affect the intensity of bushmeat hunting?
How does use of water and firewood by tourist facilities affect ecosystem health?
How do the presence of shoats and cattle differ in their effect on wildlife?
How does human-caused changes in water quality affect aquatic life?
What is the role of a declining natural prey population in driving human-wildlife conflict?
How does predator competition change with changes in resources?
How can responsible dog ownership reduce the spread of canine distemper and rabies to wildlife?
What are the drivers of the presence invasive species, e.g. can the occurrence of invasive species be linked to tourist camps / human settlements / climate change?
What is the impact of fires and different fire management regimes on biodiversity?
What is the effect of the presence of invasive species (excluding livestock) on biodiversity?
Year(s) Of Engagement Activity 2017
Description Manazine article in SWARA 
Form Of Engagement Activity A magazine, newsletter or online publication
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact Magazine article linking the kind of behavioural ecology that we are doing to conservation
Year(s) Of Engagement Activity 2018
Description Organzing NERC-ZSL 2-day Symposium "Linking behaviour to populations and communities: how can behavioural ecology inform conservation?" at the Zoological Society of London 22&23 Novermber 
Form Of Engagement Activity Participation in an activity, workshop or similar
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact Following peer-review, my proposal was accepted for organizing a two-day symposium aiming at bridging the gap between scientists working in behavioural ecology and conservation practitioners was accepted by the Zoological Society of London (GBP2,500 co-funding from NERC). The symposium was nearly sold out and included presentations and discussions by 22 leading experts from across the globe. The meeting took place in November 2018 and received very positive feedback from attendees.
Year(s) Of Engagement Activity 2018
Description Project website 
Form Of Engagement Activity Engagement focused website, blog or social media channel
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Public/other audiences
Results and Impact The website presents our project and allow interested parties to follow its progress (typical 93 visits per week by 54 individual visitors)
Year(s) Of Engagement Activity 2015,2016,2017,2018
Description Symposium presentation by Kristine Meise "Using social network analysis to predict the community-wide repercussions of severing the Great Migration in the Mara Ecosystem" (NERC/ZSL Symposium in London) 
Form Of Engagement Activity A talk or presentation
Part Of Official Scheme? No
Geographic Reach International
Primary Audience Professional Practitioners
Results and Impact Symposium talk of project findings
Year(s) Of Engagement Activity 2018