Abstract
Patterns of association in animal societies vary through space and time. Understanding such variation is key to predicting inter- and intra-population variation across factors as diverse as gene flow, disease transmission, and resilience to climate change. Here, we use 3.5 years of observational data, coupled with static and dynamic methods in social network analysis, to investigate patterns of social association in wild giraffe in the northern Namib Desert, Namibia (a habitat of extreme aridity for the species). Our static analyses reveal similar robust nested social communities, but less distinct community structures, than those found in giraffe populations inhabiting less arid environments. Furthermore, results of our dynamic social network analyses show increased social connectivity in this population during the morning and hot-dry season. These temporal patterns align with patterns of thermoregulatory and sociosexual behaviour in this population. However, they differ from temporal patterns of connectivity revealed in populations in less arid environments. Combined, results of both our static and temporal analyses suggest that while the major characteristics of giraffe society persist through space and time, there is substantial variation in the strength of patterns of social connectivity both within and between populations at multiple spatiotemporal resolutions.
Significance statement
Animal social networks are inherently dynamic. Patterns of connectivity can be shaped by both social factors and habitat variation across space and time. We reveal that a population of uniquely desert-dwelling giraffe maintains a robust, albeit weaker, community structure in a habitat at the extreme edge of its natural range. We also reveal variation in intra-population temporal patterns of connectivity at both daily and seasonal resolutions. Combined, these results suggest that while the major characteristics of giraffe society persist across habitats and through time, the strength of patterns of social connectivity markedly varies in line with habitat factors both within and between populations.
Similar content being viewed by others
Data availability
All data generated or analysed during this study are included in this published article (and its supplementary information files).
References
Amici F, Call J, Watzek J, Brosnan S, Aureli F (2018) Social inhibition and behavioural flexibility when the context changes: a comparison across six primate species. Sci Rep 8:3067
Ansmann IC, Parra GJ, Chilvers BL, Lanyon JM (2012) Dolphins restructure social system after reduction of commercial fisheries. Anim Behav 84:575–581
Baigger A, Perony N, Reuter M, Leinert V, Melber M, Grünberger S, Fleischmann D, Kerth G (2013) Bechstein’s bats maintain individual social links despite a complete reorganisation of their colony structure. Sci Nat 100:895–898
Bejder L, Fletcher D, Bräger S (1998) A method for testing association patterns of social animals. Anim Behav 56:719–722
Bennie JJ, Duffy JP, Inger R, Gaston KJ (2014) Biogeography of time partitioning in mammals. P Natl Acad Sci USA 111:13727–13732
Bercovitch FB, Bashaw MJ, del Castillo SM (2006) Sociosexual behavior, male mating tactics, and the reproductive cycle of giraffe Giraffa camelopardalis. Horm Behav 50:314–321
Bercovitch FB, Berry PS (2010) Ecological determinants of herd size in the Thornicroft’s giraffe of Zambia. Afr J Ecol 48:962–971
Biernacki C, Celeux G, Govaert G (2000) Assessing a mixture model for clustering with the integrated completed likelihood. IEEE T Pattern Anal 22:719–725
Blonder B, Wey TW, Dornhaus A, James R, Sih A (2012) Temporal dynamics and network analysis. Methods Ecol Evol 3:958–972
Blumstein DT, Fuong H, Palmer E (2017) Social security: Social relationship strength and connectedness influence how marmots respond to alarm calls. Behav Ecol Sociobiol 71:145
Bond ML, König B, Lee DE, Ozgul A, Farine DR (2021a) Proximity to humans affects local social structure in a giraffe metapopulation. J Anim Ecol 90:212–221
Bond ML, König B, Ozgul A, Farine DR, Lee DE (2021b) Socially defined subpopulations reveal demographic variation in a giraffe metapopulation. J Wildlife Manage 85:920–931
Bond ML, Lee DE, Ozgul A, König B (2019) Fission–fusion dynamics of a megaherbivore are driven by ecological, anthropogenic, temporal, and social factors. Oecologia 191:335–347
Brand R (2007) Evolutionary ecology of giraffes (Giraffa camelopardalis) in Etosha National Park, Namibia. PhD thesis, Newcastle University
Burger AL, Fennessy J, Fennessy S, Dierkes PW (2020) Nightly selection of resting sites and group behavior reveal antipredator strategies in giraffe. Ecol Evol 10:2917–2927
Cairns SJ, Schwager SJ (1987) A comparison of association indices. Anim Behav 35:1454–1469
Calenge C (2006) A tool for the analysis of space and habitat use by animals. Ecol Modell 197:516–519
Cameron AC, Windmeijer AG (1997) An R-squared measure of goodness of fit for some common nonlinear regression models. J Econom 77:329–342
Cameron EZ, du Toit JT (2005) Social influences on vigilance behaviour in giraffes, Giraffa camelopardalis. Anim Behav 69:1337–1344
Cameron EZ, Setsaas TH, Linklater WL (2009) Social bonds between unrelated females increase reproductive success in feral horses. P Natl Acad Sci USA 106:13850–13853
Carter KD, Brand R, Carter JK, Shorrocks B, Goldizen AW (2013a) Social networks, long-term associations and age-related sociability of wild giraffes. Anim Behav 86:901–910
Carter KD, Seddon JM, Frère CH, Carter JK, Goldizen AW (2013b) Fission–fusion dynamics in wild giraffes may be driven by kinship, spatial overlap and individual social preferences. Anim Behav 85:385–394
Castles M (2018) Drivers of sociability and fission-fusion dynamics in a population of wild giraffe (Giraffa Camelopardalis). PhD thesis, The University of Queensland, Australia
Chen C, Jin J, Vandeleest J, Beisner B, McCowan B, Fushing H (2019) DCG: data cloud geometry (DCG): using random walks to find community structure in social network analysis. R package version 0.9.3, https://CRAN.R-project.org/package=DCG
Clutton-Brock TH, O’Riain MJ, Brotherton PN, Gaynor D, Kansky R, Griffin AS, Manser M (1999) Selfish sentinels in cooperative mammals. Science 284:1640–1644
Crall JP, Stewart CV, Berger-Wolf TY, Rubenstein DI, Sundaresan SR (2013) Hotspotter—patterned species instance recognition. In: 2013 IEEE workshop on applications of computer vision (WACV), Clearwater Beach, pp 230–237
Csardi G, Nepusz T (2006) The igraph software package for complex network research. InterJ Complex Syst 1695:1–9
Dekker D, Krackhardt D, Snijders TA (2007) Sensitivity of MRQAP tests to collinearity and autocorrelation conditions. Psychometrika 72:563–581
Ellis S, Snyder-Mackler N, Ruiz-Lambides A, Platt ML, Brent LJ (2019) Deconstructing sociality: the types of social connections that predict longevity in a group-living primate. Proc R Soc B 286:20191991
Farine DR (2013) Animal social network inference and permutations for ecologists in R using asnipe. Methods Ecol Evol 12:1187–1194
Farine DR (2014) Measuring phenotypic assortment in animal social networks: weighted associations are more robust than binary edges. Anim Behav 89:141–153
Farine DR (2017) A guide to null models for animal social network analysis. Methods Ecol Evol 8:1309–1320
Farine DR (2018) When to choose dynamic vs. static social network analysis. J Anim Ecol 87:128–138
Farine DR, Whitehead H (2015) Constructing, conducting and interpreting animal social network analysis. J Anim Ecol 84:1144–1163
Fennessy J, Bidon T, Reuss F, Kumar V, Elkan P, Nilsson MA, Vamberger M, Fritz U, Janke A (2016) Multi-locus analyses reveal four giraffe species instead of one. Curr Biol 26:2543–2549
Fennessy J (2004) Ecology of desert-dwelling giraffe Giraffa camelopardalis angolensis in northwestern Namibia. PhD thesis: University of Sydney
Funke T, Becker T (2019) Stochastic block models: a comparison of variants and inference methods. PLoS ONE 14:0215296
Fushing H, McAssey MP (2010) Time, temperature, and data cloud geometry. Phys Rev 82:061110
Fushing H, Wang H, VanderWaal K, McCowan B, Koehl P (2013) Multi-scale clustering by building a robust and self correcting ultrametric topology on data points. PLoS ONE 8:e56259
Hart EE, Fennessy J, Chari S, Ciuti S (2020a) Habitat heterogeneity and social factors drive behavioral plasticity in giraffe herd-size dynamics. J Mammal 101:248–258
Hart EE, Fennessy J, Hauenstein S, Ciuti S (2020) Intensity of giraffe locomotor activity is shaped by solar and lunar zeitgebers. Behav Process 178:104178
Hart EE, Fennessy J, Rasmussen HB, Butler-Brown M, Muneza AB, Ciuti S (2020c) Precision and performance of an 180g solar-powered GPS device for tracking medium to large-bodied terrestrial mammals. Wildlife Biol 2020:2
Hart EE, Fennessy J, Wells E, Ciuti S (2021) Seasonal shifts in sociosexual behaviour and reproductive phenology in giraffe. Behav Ecol Sociobiol 75:15
He P, Maldonado-Chaparro AA, Farine DR (2019) The role of habitat configuration in shaping social structure: a gap in studies of animal social complexity. Behav Ecol Sociobiol 73:9
Heathcote RJ, Darden SK, Franks DW, Ramnarine IW, Croft DP (2017) Fear of predation drives stable and differentiated social relationships in guppies. Sci Rep 7:41679
Hemson G, Johnson P, South A, Kenward R, Macdonald RR, D, (2005) Are kernels the mustard? Data from global positioning system (GPS) collars suggests problems for kernel home-range analyses with least-squares cross-validation. J Anim Ecol 74:455–463
Henzi SP, Lusseau D, Weingrill T, van Schaik CP, Barrett L (2009) Cyclicity in the structure of female baboon social networks. Behav Ecol Sociobiol 63:1015–1021
Hobson EA, Avery ML, Wright TF (2013) An analytical framework for quantifying and testing patterns of temporal dynamics in social networks. Anim Behav 85:83–96
Hoppitt WJ, Farine DR (2018) Association indices for quantifying social relationships: how to deal with missing observations of individuals or groups. Anim Behav 136:227–238
Hubert L, Arabie P (1985) Comparing Partitions J Classif 2:193–218
Jacobson PJ, Jacobson KN, Seely MK (1995) Ephemeral rivers and their catchments: sustaining people and development in western Namibia. Desert Research Foundation of Namibia, Windhoek
Knüsel MA, Lee DE, König B, Bond ML (2019) Correlates of home range sizes of giraffes, Giraffa camelopardalis. Anim Behav 149:143–151
Krause J, Ruxton GD (2002) Living in groups. Oxford University Press, Oxford
Kummer H (1971) Primate societies: group techniques of ecological adaptation. Aldine, Chicago
Laver PN, Kelly MJ (2008) A critical review of home range studies. J Wildlife Manage 72:290–298
Lavista Ferres JM, Lee DE, Nasir M, Chen YC, Bijral AS, Bercovitch FB, Bond ML (2021) Social connectedness and movements among communities of giraffes vary by sex and age class. Anim Behav 180:315–328
Lee C, Wilkinson DJ (2019) A review of stochastic block models and extensions for graph clustering. Appl Netw Sci 4:122
Lee DE, Bond ML, Bolger DT (2017) Season of birth affects juvenile survival of giraffe. Popul Ecol 59:45–54
Matias C, Miele V (2017) Statistical clustering of temporal networks through a dynamic stochastic block model. J R Stat Soc B 79:1119–1141
McDonald DB (2007) Predicting fate from early connectivity in a social network. P Natl Acad Sci USA 104:10910–10914
Meise K, Franks DW, Bro-Jørgensen J (2019) Using social network analysis of mixed-species groups in African savannah herbivores to assess how community structure responds to environmental change. Phil Trans R Soc B 374:20190009
Muller Z, Bercovitch F, Brand R et al (2018a) Giraffa camelopardalis (amended version of 2016 assessment). The IUCN Red List of Threatened Species 2018a: e.T9194A136266699, https://doi.org/10.2305/IUCN.UK.2016-3.RLTS.T9194A136266699.en
Muller Z, Cantor M, Cuthill IC, Harris S (2018b) Giraffe social preferences are context dependent. Anim Behav 146:37–49
Muller Z, Cuthill IC, Harris S (2019) Giraffe (Giraffa camelopardalis) social networks in areas of contrasting human activity and lion density. Ethology 125:702–715
Nandini S, Keerthipriya P, Vidya TNC (2017) Seasonal variation in female Asian elephant social structure in Nagarahole-Bandipur, southern India. Anim Behav 134:135–145
Newman M (2003) Modularity, community structure, and spectral properties of networks. Phys Rev E 67:026126
Noonan MJ, Tucker MA, Fleming CH et al (2019) A comprehensive analysis of autocorrelation and bias in home range estimation. Ecol Monogr 89:e01344
Packer C, Swanson A, Ikanda D, Kushnir H (2011) Fear of darkness, the full moon and the nocturnal ecology of African lions. PLoS ONE 6:e22285
Papageorgiou D, Farine DR (2021) Multilevel societies in birds. Trends Ecol Evol 36:15–17
Peignier M, Webber QM, Koen EL, Laforge MP, Robitaille AL, Vander Wal E (2019) Space use and social association in a gregarious ungulate: testing the conspecific attraction and resource dispersion hypotheses. Ecol Evol 9:5133–5145
Pilot M, Dahlheim ME, Hoelzel AR (2010) Social cohesion among kin, gene flow without dispersal and the evolution of population genetic structure in the killer whale (Orcinus orca). J Evol Biol 23:20–31
Prehn SG, Laesser BE, Clausen CG, Jønck K, Dabelsteen T, Brask JB (2019) Seasonal variation and stability across years in a social network of wild giraffe. Anim Behav 157:95–104
Qi XG, Garber PA, Ji W et al (2014) Satellite telemetry and social modeling offer new insights into the origin of primate multilevel societies. Nat Commun 5:5296
R Core Team (2019) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria, https://www.R-project.org/
Sah P, Mann J, Bansal S (2018) Disease implications of animal social network structure: a synthesis across social systems. J Anim Ecol 87:546–558
Shizuka D, Farine DR (2016) Measuring the robustness of network community structure using assortativity. Anim Behav 112:237–246
Shizuka D, Johnson AE (2020) How demographic processes shape animal social networks. Behav Ecol 31:1–11
Snijders L, Blumstein DT, Stanley CR, Franks DW (2017) Animal social network theory can help wildlife conservation. Trends Ecol Evol 32:567–577
Snijders TA, Borgatti SP (1999) Non-parametric standard errors and tests for network statistics. Connections 22:161–170
Snyder-Mackler N, Burger JR, Gaydosh L et al (2020) Social determinants of health and survival in humans and other animals. Science 368:6493
Stander P, Steenkamp W, Steenkamp L (2018) Vanishing kings: lions of the Namib Desert. HPH Publishing, Johannesburg
Suraud JP (2011) Identifier les contraintes pour la conservation des dernières girafes de l’Afrique de l’Ouest: Déterminants de la dynamique de la population et patron d’ occupation spatiale. PhD thesis, L’Université Claude Bernard, Lyon, France
Sutton N (2019) Effects of food limitation on social grouping and foraging in a fission-fusion species. MSc Thesis, University of Canterbury
VanderWaal KL, Wang H, McCowan B, Fushing H, Isbell LA (2014) Multilevel social organization and space use in reticulated giraffe (Giraffa camelopardalis). Behav Ecol 25:17–26
Vazquez C, Rowcliffe JM, Spoelstra K, Jansen PA (2019) Comparing diel activity patterns of wildlife across latitudes and seasons: time transformations using day length. Methods Ecol Evol 10:2057–2066
Warton DI, Hui FKC (2011) The arcsine is asinine: the analysis of proportions in ecology. Ecology 92:3–10
Whitehead H (2008) Precision and power in the analysis of social structure using associations. Anim Behav 75:1093–1099
Whitehead H, Dufault S (1999) Techniques for analyzing vertebrate social structure using identified individuals. Adv Stud Behav 28:33–74
Wittemyer G, Douglas-Hamilton I, Getz WM (2005) The socioecology of elephants: analysis of the processes creating multitiered social structures. Anim Behav 69:1357–1371
Wolf TE, Ngonga Ngomo AC, Bennett NC, Burroughs R, Ganswindt A (2018) Seasonal changes in social networks of giraffes. J Zool 305:82–87
Worton BJ (1989) Kernel methods for estimating the utilization distribution in home-range studies. Ecology 70:164–168
Acknowledgements
We thank the reviewers for helpful comments that improved the manuscript.
Funding
This project was funded by the Giraffe Conservation Foundation, the School of Biology and Environmental Science (SBES) at University College Dublin (UCD). Wilderness Wildlife Trust, Columbus Zoo and Aquarium, Idea Wild, and Woodland Park Zoo also provided financial support and TOSCO, ATI Holidays, Karibu Safari, Ultimate Safaris, Matiti Safaris, and Charlottenberg Arabians contributed towards the purchase of a research vehicle.
Author information
Authors and Affiliations
Corresponding author
Ethics declarations
Ethical approval
Data were collected under the Giraffe Conservation Foundation, Namibian Ministry of Environment and Tourism (MET), Namibian Chamber of Environment and National Commission on Research Science and Technology research permit numbers 2162/2016, 2277/2017 and RCIV00042018, and under the University College Dublin (UCD) ethics approval number AREC-E-16–50. All applicable international, national, and/or institutional guidelines for the use of animals were followed.
Conflict of interest
The authors declare no competing interests.
Additional information
Communicated by D. Paul Croft.
Publisher’s note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Supplementary Information
Below is the link to the electronic supplementary material.
Rights and permissions
About this article
Cite this article
Hart, E.E., Ciuti, S., Herrmann, L. et al. Static and dynamic methods in social network analysis reveal the association patterns of desert-dwelling giraffe. Behav Ecol Sociobiol 76, 62 (2022). https://doi.org/10.1007/s00265-022-03167-9
Received:
Revised:
Accepted:
Published:
DOI: https://doi.org/10.1007/s00265-022-03167-9