How does a frugivore handle its meal? It is clear that the appropriate technique will vary with fruit type. For instance, we deal with strawberries (by swallowing) rather differently than avocados. There is a growing realization that costs are important to frugivores (e.g. Sorensen 1981, Martin 1985, Foster 1987, Levey 1987, Hegde et al. 1991, Levey and Grajal 1991, Levey and Duke 1992). It is the purpose of this note to point out that there are several implicit assumptions in the formulation of These assumptions appear unrealistic for many fruit-frugivore interactions (and perhaps many other relationships as well). A richer description of the ways frugivores deal with their food may be more realistic and predictive. Handling time is a common concept in studies of interactions between consumers and their food. Holling (1959) introduced the term, and distinguished between the events leading up to contact between a predator and its prey (search) and the subsequent events, as the predator pursues, subdues, and consumes its prey (handling). The concept has proven useful in a number of subdisciplines within ecology, most notably the study of foraging behaviour. Handling time has gradually come to mean the time between periods of search, time spent in activities such as pursuit, capture, killing and consumption of prey, but also all activities necessary to resume search. Begon et al. (1986: 322-325) give a good summary of the relevant ideas and data. Under the classical definition, as used in the influential models of MacArthur and Pianka (1966) and many followers, time can be spent either on search or on handling, but not simultaneously on both. In the case of many large predators, such a model may be appropriate; for some others it is not. In particular, some components of may occur synchronously with search or other activities. For instance, some animals appear to be gut-process limited; ingestion is limited by the rate at which gut volume becomes available for more food (e.g. frugivores: Walsberg 1975, Sorensen 1984, Tedman and Hall 1985, Levey and Grajal 1991, Levey and Duke 1992; and nectarivores: Diamond et al. 1986, Karasov et al. 1986, Karasov and Diamond 1987). Interestingly, some frugivorous birds can offset the problem of gut process constraints by storing ingested fruits in a distensible esophagus (Levey and Duke 1992). In this way, such frugivores can consume more fruits per feeding bout than their gizzard and intestine can process at any one time. Digestion is therefore clearly part of handling; but during periods of digestion, individuals can engage in other activities, such as territorial defence, sampling behaviour, or search. Similarly, studies of optimal oviposition behaviour in insects have begun to recognize the importance of egg maturation processes (Griffiths 1969, Jones 1977, Courtney et al. 1989). A female butterfly can be simultaneously maturing an egg, and searching for a place to deposit it (Courtney 1986). To return to the original definition of handling, it is clear that it has been used to cover several behaviours (pursuit, capture, consumption, digestion, etc.). We now argue that it is productive to distinguish among these different components, precisely because some may occur synchronously with search or other activities, while others are exclusive of alternative behaviours. This distinction is not fussy: it becomes interesting when applied to frugivores. We also emphasize the need to recognize that digestion is part of handling. Studies which do not incorporate digestion into time (e.g. Kaspari 1990, Woodrey 1990, Hegde et al. 1991) may underestimate the influence gut-processing constraints have on prey choice. For example, Hegde et al. (1991) measured time for red-vented bulbuls (Pycnonotus cafer L.) eating fruits of Solanum pubescens Willd. (Solanaceae) as the time from picking to swallowing. Hegde et al. concluded that fruit preference was based upon the benefits and costs of handling; we do not disagree with their results, but emphasize that handling of fruits in the gut may have also influenced preference. Recently, a number of studies have recognized the importance of digestive physiology (Martinez del Rio et al. 1988, 1989, Karasov and Levey 1990,