Abstract

Animal cooperation in the wild often involves multiple individuals that must tolerate each other in close proximity. However, most cooperation experiments in the lab are done with two animals, that are often also physically separated. Such experiments are useful for answering some pertinent questions, for example about the understanding of the role of the partner and strategies of partner control, but say little about factors determining successful cooperation with multiple partners in group settings. We explored the influence of dominance, rank distance, tolerance, affiliation, and coordination by testing kea parrots with a box requiring two, three, or four chains to be pulled simultaneously to access food rewards. The reward could be divided unevenly, but not monopolized completely. Eventually dyadic, triadic, and tetradic cooperation tasks were solved, showing that non-human animals are capable of tetradic cooperation in an experimental setup. Starting with two chains, we found that in a dyad monopolization of the box by the highest-ranking bird was the largest obstacle preventing successful cooperation. High-ranking birds learned to restrain themselves from monopolizing the box during a single session in which monopolization was hindered by the presence of a large number of birds. Thereafter, restraint by dominants remained the strongest factor determining success in the first trial in dyadic, triadic, and tetradic setups. The probability of success increased with the degree of restraint shown by all dominant subjects present. Previous experience with the task contributed to success in subsequent sessions, while increasing rank distance reduced success notably in the four-chain setup.

Highlights

  • IntroductionOver the last two decades, experiments testing cooperation in animals have been conducted successfully with many different species, including corvids (Corvus corax – AsakawaHaas, Schiestl, Bugnyar, & Massen, 2016; Massen, Ritter, & Bugnyar, 2015; Corvus frugilegus – Seed, Clayton, & Emery, Comparative Cognition Unit, Messerli Research Institute, University of Veterinary Medicine Vienna, University of Vienna, Medical University Vienna, Veterinärplatz 1, 1210 Vienna, AustriaHaidlhof Research Station, University of Veterinary Medicine, University of Vienna, Haidlhof 204, 2540 Bad Vöslau, AustriaPsychologie, Université de Strasbourg, 12 Rue Goethe, 67000 Strasbourg, FranceCollege of Forestry, Guangxi University, No 100 Daxue Road, Nanning, Guangxi 530005, People’s Republic of China2008; Corvus moneduloides – Jelbert, Singh, Gray, Taylor, & Marshall, 2015), parrots (Psittacus Erithacus – Péron, RatFischer, Lalot, Nagle, & Bovet, 2011; Nestor notabilis – Schwing, Jocteur, Wein, Massen, & Noë, 2016; Schwing, Reuillon, Conrad, Noë, & Huber, 2020), primates (Pongo pygmeus – Chalmeau, Lardeux, Brandibas, & Gallo, 1997; Cebus apella – Mendres & de Waal, 2000; Visalberghi, Quarantotti, & Tranchida, 2000; Saguines oedipus – Cronin, Kurian, & Snowdon, 2005; Callithrix jacchus – Werdenich & Huber, 2002; Pan troglodytes – Hare, Melis, Woods, Hastings, & Wrangham, 2007; Melis, Hare, & Tomasello, 2006), canines (Canis lupus – Marshall-Pescini, Schwarz, Kostelnik, Virányi, & Range, 2017; Canis familiaris), as well as other mammals (Crocuta crocuta – Drea & Carter, 2009; Elephas maximus – Plotnik, Lair, Suphachoksahakun, & De Waal, 2011; Tursiops truncatus – Jaakkola, Guarino, Donegan, & King, 2018)

  • A subject that waits for its partner in the loose-string paradigm has shown the ability for coordination, or at least synchrony, as the spatial aspect is often artificially restricted by the laboratory setting ( see, e.g., Marshall-Pescini, Schwarz, Kostelnik, Virányi, & Range, 2017, or Schwing et al, 2020, for setups with a spatial aspect by presenting subject(s) with two apparatuses simultaneously)

  • We analyzed four in more detail because we considered them important based on both results of cooperation experiments with other species and our previous experience with kea: (1) monopolization of the apparatus, or essential parts thereof, by dominant animals, (2) reward division, (3) the power difference of subjects, as measured by rank distance, and (4) the affiliation qualities of their relationships

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Summary

Introduction

Over the last two decades, experiments testing cooperation in animals have been conducted successfully with many different species, including corvids (Corvus corax – AsakawaHaas, Schiestl, Bugnyar, & Massen, 2016; Massen, Ritter, & Bugnyar, 2015; Corvus frugilegus – Seed, Clayton, & Emery, Comparative Cognition Unit, Messerli Research Institute, University of Veterinary Medicine Vienna, University of Vienna, Medical University Vienna, Veterinärplatz 1, 1210 Vienna, AustriaHaidlhof Research Station, University of Veterinary Medicine, University of Vienna, Haidlhof 204, 2540 Bad Vöslau, AustriaPsychologie, Université de Strasbourg, 12 Rue Goethe, 67000 Strasbourg, FranceCollege of Forestry, Guangxi University, No 100 Daxue Road, Nanning, Guangxi 530005, People’s Republic of China2008; Corvus moneduloides – Jelbert, Singh, Gray, Taylor, & Marshall, 2015), parrots (Psittacus Erithacus – Péron, RatFischer, Lalot, Nagle, & Bovet, 2011; Nestor notabilis – Schwing, Jocteur, Wein, Massen, & Noë, 2016; Schwing, Reuillon, Conrad, Noë, & Huber, 2020), primates (Pongo pygmeus – Chalmeau, Lardeux, Brandibas, & Gallo, 1997; Cebus apella – Mendres & de Waal, 2000; Visalberghi, Quarantotti, & Tranchida, 2000; Saguines oedipus – Cronin, Kurian, & Snowdon, 2005; Callithrix jacchus – Werdenich & Huber, 2002; Pan troglodytes – Hare, Melis, Woods, Hastings, & Wrangham, 2007; Melis, Hare, & Tomasello, 2006), canines (Canis lupus – Marshall-Pescini, Schwarz, Kostelnik, Virányi, & Range, 2017; Canis familiaris), as well as other mammals (Crocuta crocuta – Drea & Carter, 2009; Elephas maximus – Plotnik, Lair, Suphachoksahakun, & De Waal, 2011; Tursiops truncatus – Jaakkola, Guarino, Donegan, & King, 2018). Boesch and Boesch (1989) in describing behavior in chimpanzees, suggested four levels of growing complexity with regard to hunting, all of which were considered cooperative as they were all directed at the same prey item: (1) similarity – similar actions but without relation in time and space; (2) synchrony – similar actions with relation in time; (3) coordination – similar actions with relation in time and space; and (4) collaboration – different actions that are complementary in nature in working to achieve success Based on these levels, a subject that waits for its partner in the loose-string paradigm has shown the ability for coordination, or at least synchrony, as the spatial aspect is often artificially restricted by the laboratory setting ( see, e.g., Marshall-Pescini, Schwarz, Kostelnik, Virányi, & Range, 2017, or Schwing et al, 2020, for setups with a spatial aspect by presenting subject(s) with two apparatuses simultaneously). Tolerance in a lab setting is generally used to describe the occurrence of co-feeding in artificial and natural shareable food patches – notably in the primate literature (Hare, Melis, Woods, Hastings, & Wrangham, 2007; Kasper, Voelkl, & Huber, 2008; Melis et al, 2006; Mendres & de Waal, Mendres & de Waal, 2000; Petit et al, 1992; Suchak, Eppley, Campbell, & de Waal, 2014), and in cooperation studies with corvids (Massen et al, 2015; Seed, Clayton, & Emery, 2008)

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