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Influence of Intraspecific Relationships on the Allocation of Temporal Resources in the Lekking Ruff, Calidris Pugnax

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Influence of Intraspecific Relationships on the Allocation of Temporal Resources in the Lekking Ruff, Calidris Pugnax Influence of Intraspecific Relationships on the Allocation of Temporal Resources in the Lekking Ruff, Calidris pugnax Jaani Mustonen MSc Thesis University of Oulu Faculty of Science Department of Biology January 2020 Abstract Alternative Reproductive Tactics (ARTs) are utilized by animals of both sex to optimize their reproductive success in an intrasexually competitive environment where they are unable to prevail through use of the predominant tactic for obtaining mates. Often manifested as subtle differences in physical size, species where distinct genetically fixed morphological differences are associated with specific tactics are rare. The ruff (Calidris pugnax), a lekking palearctic wader, exhibits three fixed morphs (territorial and combative “independents”, cooperative and non-aggressive “satellites” and the female mimicking “faeders”) with distinct morphologies and reproductive behaviours which have their reproductive success interwoven to one-another on the leks. This diversity creates intricate connections between the morphs and sexes, giving rise to a web of influential relationships that have not been investigated thoroughly. These relationships were explored through utilization of stand-alone cameras to obtain precise information on the time use and reproductive success of male morphs and females and the evolution of their time investment throughout the lekking season. The degree of influence morphs and sexes exerted on the allocation of temporal resources by others through their own corresponding time investment was particularly investigated. The gradual progress of the female receptivity eventually resulting in a rapid copulatory peak was described, as well as and the central role of the time investment by dominant males on the attractiveness of a lek, both to females and other males alike. Indications of a more complex role of the satellite males in the overall success of a lek were identified, suggesting of even greater intricacy in the connections between the morphs. Table of Contents 1. Introduction ............................................................................................................................................. 1 1.1 Alternative Reproductive Tactics.......................................................................................................... 1 2. Ruff: A Life history ......................................................................................................................................................... 5 2.1 Description ..................................................................................................................................................... 5 2.2 Range & Habitat ........................................................................................................................................... 6 2.3 Male morph variation and its genetic background ....................................................................... 8 2.3.1 The Independent Male ........................................................................................................................... 9 2.3.2 The Satellite Male ................................................................................................................................. 10 2.3.3 The Faeder Male .................................................................................................................................... 11 2.3.4 The Female .............................................................................................................................................. 12 3. Reproduction ......................................................................................................................................... 14 3.1 Lekking paradox ....................................................................................................................................... 14 3.2 Lekking in Scolopacidae ........................................................................................................................ 14 3.4 Mating ........................................................................................................................................................... 16 3.5 Nesting .......................................................................................................................................................... 18 4.0 Key Questions ..................................................................................................................................... 18 5.0 Materials and Methods..................................................................................................................... 20 5.1 Study sites .................................................................................................................................................... 20 5.2 Lek recording and field observations............................................................................................... 20 5.3 Nest location and bird handling ......................................................................................................... 21 5.4 Statistical analysis ................................................................................................................................... 22 6.0 Results ................................................................................................................................................... 23 6.1 Weather ........................................................................................................................................................ 23 6.2 Lek and field observations .................................................................................................................... 24 6.3 Receptiveness and time use .................................................................................................................. 28 6.4 Correlations between morphs, sexes and temporal variables ............................................... 30 6.5 Lek attendance and copulations frequency ................................................................................... 32 7.0 Discussion ............................................................................................................................................ 34 7.1 Progress of lekking .................................................................................................................................. 34 7.2 Influential relationships ........................................................................................................................ 35 7.3 Decisions influencing time and resource allocation .................................................................. 37 8.0 Conclusions: ........................................................................................................................................ 41 8.1 Answering the key questions ............................................................................................................... 41 8.2 Review of methodology .......................................................................................................................... 42 8.3 Guidelines for future studies................................................................................................................ 43 References ................................................................................................................................................. 44 Supplementary materials ..................................................................................................................... 50 1. Introduction 1.1 Alternative Reproductive Tactics Alternative Reproductive Tactics, referred to as ARTs, are consistent intraspecific and intrasexual variations in reproductive behaviours. They serve the purpose of maximizing the number of fertilizations obtained by an individual animal, both male and female. A strategy is defined as a set of decision rules with a genetic basis that give rise to tactics, which are behavioural phenotypes resulting from a strategy. Multiple different tactics can be included within a strategy (Gross 1996; Taborsky et al. 2008). ARTs are generally characterized by traits that have evolved towards the same functional end and display discontinuous distribution. This can be observed in many forms, such as size dimorphism, colour polymorphism, dimorphic morphological structures and behavioural alternatives. There are multiple pathways for ARTs to manifest in an individual (Figure 1). A genetically fixed tactic occurs inevitably but if fixation is developmental and alternatives exist, environmental conditions may influence which tactic is manifested. Variation in tactics utilized by monomorphic individuals can rise from multiple factors, such as resource availability, level of competition or passing a threshold such as size or age (Taborsky et al. 2008). Figure 1. A simplified overview of possible developmental Alternative Reproductive Tactic pathways. Red and blue colours represent different tactics whereas black colour represents an immature state where no tactic has manifested. Edited from Shcradin (2019). 1 In a classical paper on Alternative Reproductive Tactics, Gross (1996) introduced three categories of strategies (Figure 2): Figure 2. Categories of reproductive strategies by Gross (1996), edited from Schradin (2019). Alternative strategies are genetically fixed, polymorphic and based on frequency-dependent selection where
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