Ecosystem engineers have been widely studied for terrestrial systems, but global trends in research encompassing the range of taxa and functions have not previously been synthesised. We reviewed contemporary understanding of engineer fauna in terrestrial habitats and assessed the methods used to document patterns and processes, asking: (a) which species act as ecosystem engineers and with whom do they interact? (b) What are the impacts of ecosystem engineers in terrestrial habitats and how are they distributed? (c) What are the primary methods used to examine engineer effects and how have these developed over time? We considered the strengths, weaknesses and gaps in knowledge related to each of these questions and suggested a conceptual framework to delineate "significant impacts" of engineering interactions for all terrestrial animals. We collected peer-reviewed publications examining ecosystem engineer impacts and created a database of engineer species to assess experimental approaches and any additional covariates that influenced the magnitude of engineer impacts. One hundred and twenty-two species from 28 orders were identified as ecosystem engineers, performing five ecological functions. Burrowing mammals were the most researched group (27%). Half of all studies occurred in dry/arid habitats. Mensurative studies comparing sites with and without engineers (80%) were more common than manipulative studies (20%). These provided a broad framework for predicting engineer impacts upon abundance and species diversity. However, the roles of confounding factors, processes driving these patterns and the consequences of experimentally adjusting variables, such as engineer density, have been neglected. True spatial and temporal replication has also been limited, particularly for emerging studies of engineer reintroductions. Climate change and habitat modification will challenge the roles that engineers play in regulating ecosystems, and these will become important avenues for future research. We recommend future studies include simulation of engineer effects and experimental manipulation of engineer densities to determine the potential for ecological cascades through trophic and engineering pathways due to functional decline. We also recommend improving knowledge of long-term engineering effects and replication of engineer reintroductions across landscapes to better understand how large-scale ecological gradients alter the magnitude of engineering impacts.

SUMMARYThe interaction between temperature and diet quality can affect the life history of ectotherms. The rate and ratio at which protein and carbohydrate are obtained from food are important aspects of diet quality, and insects have a well-developed capacity to adjust their feeding behaviour and post-ingestive physiology to regulate intake and allocation of these nutrients. If the supply of protein and carbohydrate varies with temperature (e.g. via effects on intake, digestion or metabolism), then herbivorous insects can use thermoregulatory behaviour to help achieve nutritional homeostasis. When fed the host grass Triticum aestivum, Locusta migratoria nymphs absorbed and allocated protein and carbohydrate to growth with the same efficiency at 38°C as at 32°C; however, at the higher temperature, they ingested more food. In contrast, when feeding on Themeda triandra, the nymphs absorbed carbohydrate with higher efficiency at 32°C, and protein at 38°C. Using synthetic diets, we induced either a protein or a carbohydrate deficiency in experimental insects and showed that locusts placed in a thermal gradient following a meal of T. triandra selected 32°C when deprived of carbohydrate, and 38°C when deprived of protein. This capacity to use thermoregulatory behaviour to redress an imposed nutritional imbalance improved with experience of feeding on T. triandra. As predicted, locusts fed T. aestivum always chose higher temperatures, irrespective of nutritional state. Our results have consequences for understanding host-plant choice by herbivores and interpreting the effects of changed environmental temperatures and microclimate on animal-plant interactions. Supplementary material available online at

Because key nutritional processes differ in their thermal optima, ectotherms may use temperature selection to optimize performance in changing nutritional environments. Such behaviour would be especially advantageous to small terrestrial animals, which have low thermal inertia and often have access to a wide range of environmental temperatures over small distances. Using the locust, Locusta migratoria, we have demonstrated a direct link between nutritional state and thermoregulatory behaviour. When faced with chronic restrictions to the supply of nutrients, locusts selected increasingly lower temperatures within a gradient, thereby maximizing nutrient use efficiency at the cost of slower growth. Over the shorter term, when locusts were unable to find a meal in the normal course of ad libitum feeding, they immediately adjusted their thermoregulatory behaviour, selecting a lower temperature at which assimilation efficiency was maximal. Thus, locusts use fine scale patterns of movement and temperature selection to adjust for reduced nutrient supply and thereby ameliorate associated life-history consequences.

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