Integrative developmental ecology: a review of density-dependent effects on life-history traits and host-microbe interac

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Integrative developmental ecology: a review of density‑dependent effects on life‑history traits and host‑microbe interactions in non‑social holometabolous insects Anh The Than1,2 · Fleur Ponton1 · Juliano Morimoto3  Received: 18 February 2020 / Accepted: 24 August 2020 / Published online: 2 September 2020 © The Author(s) 2020

Abstract Population density modulates a wide range of eco-evolutionary processes including interand intra-specific competition, fitness and population dynamics. In holometabolous insects, the larval stage is particularly susceptible to density-dependent effects because the larva is the resource-acquiring stage. Larval density-dependent effects can modulate the expression of life-history traits not only in the larval and adult stages but also downstream for population dynamics and evolution. Better understanding the scope and generality of densitydependent effects on life-history traits of current and future generations can provide useful knowledge for both theory and experiments in developmental ecology. Here, we review the literature on larval density-dependent effects on fitness of non-social holometabolous insects. First, we provide a functional definition of density to navigate the terminology in the literature. We then classify the biological levels upon which larval density-dependent effects can be observed followed by a review of the literature produced over the past decades across major non-social holometabolous groups. Next, we argue that host-microbe interactions are yet an overlooked biological level susceptible to density-dependent effects and propose a conceptual model to explain how density-dependent effects on host-microbe interactions can modulate density-dependent fitness curves. In summary, this review provides an integrative framework of density-dependent effects across biological levels which can be used to guide future research in the field of ecology and evolution. Keywords  Niche theory · Ecological specialisation · Ecological theory · Population density · Microbiome · Host-microbiome interaction

* Juliano Morimoto [email protected] 1

Department of Biological Sciences, Macquarie University, Sydney, NSW 2109, Australia

2

Department of Entomology, Vietnam National University of Agriculture, Trau Quy, Gia Lam, Hanoi, Vietnam

3

School of Biological Sciences, University of Aberdeen, Zoology Building, Tillydrone Ave, Aberdeen AB24 2TZ, UK



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Evolutionary Ecology (2020) 34:659–680

Introduction Population density is a key factor affecting life-history trait expression and trade-offs (Mueller et  al. 1991). Individuals from low-density populations often display higher expression of fitness-related traits (e.g., higher fecundity, larger size) compared to individuals from high-density populations (Dey and Joshi 2018; Prasad and Joshi 2003). This is because high-density increases intraspecific competition which limits per capita resource acquisition (Klepsatel et al. 2018). On the other hand, social interactions may benefit individuals t