An understanding of biological fitness is central to theory and practice in ecology and evolution, yet fitness remains an elusive concept to define and challenging to measure accurately. Fitness reflects an individual's ability to pass its alleles on to subsequent generations. Researchers often quantify proxies for fitness, such as survival, growth or reproductive success. However, it can be difficult to determine lifetime fitness, especially for species with long lifespans. The abiotic and biotic environment strongly affects the expression of fitness, which means that fitness components can vary through both space and time. This spatial and temporal heterogeneity results in the impressive range of adaptations that we see in nature. Here, we review definitions of fitness and approaches to measuring fitness at the level of genes, individuals, genotypes, and populations and highlight that fitness is a key concept linking ecological and evolutionary thought.
