Table 1.

Winged flight as an innovation ecosystem. This table shows multiple (non-exhaustive) biological levels of organization, how they are associated with invention, novelty and innovation, and how these associations manifest in the example of winged flight in birds, which originated in dinosaurs and persists to this day in their avian descendants with numerous adaptive and novel modifications [68,69], and loss of flight in certain lineages. ‘Winged flight’ is a convenient label for an innovation, but the underlying adaptations and novelties form a trait complex, and together with environmental interactions, compose an innovation ecosystem. All components of this innovation ecosystem (e.g. developmental pathways, tissue architectures, and behaviours) contribute to winged flight, and generic examples given (e.g. in the ‘molecules, cells, tissues’ level) are non-exhaustive. Components may themselves be novelties or innovations, may be singular to wings and flight, and may have been co-opted or modified from other trait complexes (see e.g. [70] for the origins of feathers). Symbol x (?) indicates that the entity does (may) affect the associated biological level.

biological level	genes	molecules, cells, tissues	organs, systems, appendages	individual	population	community, ecosystem
role in innovation ecosystem	encoding	expression and development	physiology and behaviour	ecological function and performance	selection and evolution	ecological impact
invention	x	x	?	?
novelty	x	x	x	x	?
innovation	x	x	x	x	x	x
example of winged flight in birds	genes encoding the development and expression of bird appendages and wings in particular	development and expression (tissue structure and metabolism) produce and maintain wings and their substructures (muscles, feathers, cartilage)	wings enable propelled flight, nervous systems permit diverse flight behaviours	flight enables escape from antagonists, local foraging, long-distance displacement, thereby increasing survival and resource acquisition	enhanced performance increases fitness and representation of heritable novel trait in future generations	ecological success of innovated species and subsequent adaptive radiation may have impacts
references	[71]	[71–73]	[67,70]	[74–76]	[75,77,78]	[79,80]