Tropicalization unlocks novel trophic pathways and enhances secondary productivity in temperate reefs

The structure of ecological communities is rapidly changing across the globe due to climate-mediated shifts in species distributions, with novel ecosystem states emerging as new species become dominant. While it is clear that such changes restructure habitat properties and their associated assemblages, how new nutritional resources and consumers may affect the ecosystem trophic structure and energetic dynamics remains poorly resolved. Here, we investigate how the reconfiguration of tropicalized reefs-that is, temperate ecosystems receiving an influx of tropical species-affects the availability of nutritional resources and energy flow through herbivore-based trophic pathways. To do so, we quantified changes in algal and detrital resources along a tropicalization gradient in eastern Australia. We also estimated energy flow towards herbivorous fishes by quantifying their standing biomass (kg/ha), productivity (kg ha(-1) day(-1)), and biomass turnover (% day(-1)), using an individual-level modelling approach that combines estimates of fish growth rates and mortality. Along the gradient, tropicalized reefs had relatively higher amounts of palatable algal turfs and detrital particulates compared to non-tropicalized sites. Feeding intensity by herbivorous fishes was also c. 400 times higher on tropicalized reefs, with tropical turf-cropping surgeon fishes being responsible for >98% of the feeding rates. Turf-driven trophic pathways underpinned virtual all (>99%) of the biomass produced by tropical fishes, which contributed up to 63% of the total herbivorous fish standing biomass and 86% of the productivity despite only representing 35% of the abundance. Turfs also fuelled most of the secondary productivity of tropical fishes on tropicalized reefs across Australia's western tropical-temperate transition zone, although their overall productivity was similar to 5 times lower. This is possibly due to turfs there containing elevated sediment loads, which dilute the nutritional quality of turf-based resources, as well as other differences in the biogeographical context. We propose that algal turfs are central drivers of ecosystem energetic shifts on temperate reefs as they become tropicalized because they support novel algal UWA and detrital trophic pathways that enhance secondary productivity and biomass turnover. Our results also suggest that turf characteristics such as organic content or sediment load may emerge as increasingly important drivers of energy flow in temperate locations where turfs dominate benthic cover.