Body size is manifestly one of the most important attributes of an organism from an ecological and evolutionary point of view. Size has a predominant influence on an animal's energetic requirements, its potential for resource exploitation, and its susceptibility to natural enemies. A large literature now exists on how physiological, life history, and population parameters scale with body dimensions (24, 131). The ecological literature on species interactions and the structure of animal communities also stresses the importance of body size. Differences in body size are a major means by which species avoid direct overlap in resource use (153), and size-selective predation can be a primary organizing force in some communities (20, 70). Size thus imposes important constraints on the manner in which an organism interacts with its environment and influences the strength, type, and symmetry of interactions with other species (152, 207). Paradoxically, ecologists have virtually ignored the implications of these observations for interactions among species that exhibit size-distributed populations. For instance, it has been often suggested that competing species

https://www.jstor.org/stable/pdf/2096954.pdf

The ontogenetic niche and species interactions in size-structured populations

Beginning with Emlen (1966) and MacArthur and Pianka (1966) and extending through the last ten years, several authors have sought to predict the foraging behavior of animals by means of mathematical models. These models are very similar,in that they all assume that the fitness of a foraging animal is a function of the efficiency of foraging measured in terms of some "currency" (Schoener, 1971) -usually energy- and that natural selection has resulted in animals that forage so as to maximize this fitness. As a result of these similarities, the models have become known as "optimal foraging models"; and the theory that embodies them, "optimal foraging theory." The situations to which optimal foraging theory has been applied, with the exception of a few recent studies, can be divided into the following four categories: (1) choice by an animal of which food types to eat (i.e., optimal diet); (2) choice of which patch type to feed in (i.e., optimal patch choice); (3) optimal allocation of time to different patches; and (4) optimal patterns and speed of movements. In this review we discuss each of these categories separately, dealing with both the theoretical developments and the data that permit tests of the predictions. The review is selective in the sense that we emphasize studies that either develop testable predictions or that attempt to test predictions in a precise quantitative manner. We also discuss what we see to be some of the future developments in the area of optimal foraging theory and how this theory can be related to other areas of biology. Our general conclusion is that the simple models so far formulated are supported are supported reasonably well by available data and that we are optimistic about the value both now and in the future of optimal foraging theory. We argue, however, that these simple models will requre much modification, espicially to deal with situations that either cannot easily be put into one or another of the above four categories or entail currencies more complicated that just energy.

Citation classic - optimal foraging - a selective review of theory and tests

"The question of the return of birds to their homes is one of perennial interest. How faithfully do adult birds-males and females return to their territories? How far from their birth place do young birds settle? Over how much ground does one family scatter?" (91). Over 40 years ago, ornithologists studying the movement of birds in relation to their birth and breeding sites were preoccupied with estimating the extent of mixing of individuals within a species's range. There were major disagreements about how far young birds dispersed. Some authors felt that young birds did not tend to return to their birthplaces (101) but selected a nesting site anywhere within the species's natural range (27, 81). Others concluded that birds attempting to breed for the first time did, on the whole, return to their birthplaces though the extent of this fidelty was less than that of adults to their previous breeding sites (36, 69, 70, 91). It was generally agreed that adult birds did return to a previous breeding place. Gradually the position of the supporters of the random dispersal theory was undermined as more data accumulated and methodological problems of measuring dispersal came to be better appreciated. A number of longterm studies were reported between the late 1930s and the early 1950s (4,

The natal and breeding dispersal of birds

Structural complexity of the habitat often reduces predatory efficiency by reducing prey capture rates. Prey density is often positively correlated with habitat structure because it pro- vides food and substrate to the prey as well as a relative refuge from predators. Dense structure inhibits foraging, allowing abundant, highly profitable prey to coexist with predators. Sparse structure allows efficient foraging and generally contains few highly profitable prey. This suggests that feeding rates of predators may be maximized at intermediate structure. If this is true, we might also expect predator growth rates to be higher in intermediate structure habitats. Since diet breadth is thought to be related to rates of encounter with profitable prey, we also expect diets of predators to be narrower at intermediate structure than in either sparsely or densely structured habitats. Bluegill sunfish (Lepomis macrochirus) restricted to experimental ponds varying in vegetation density grew better and consumed more prey at intermediate macrophyte density than fish held at either low or high macrophyte densities. Fish at low macrophyte density had narrower diets than expected due to high initial prey availability relative to prey available at intermediate and high macro- phyte density. Fish at high macrophyte density ate fewer, but larger, prey and thus had a narrower diet than expected. Fish predation reduced total prey biomass as well as mean prey size and altered the prey community structure by removing large active invertebrate predators and herbivores with subsequent release of smaller invertebrate predators and herbivores. These changes in prey com- munity structure were also mediated by habitat structure. Habitat structure-food density interactions may be added to temperature and presence of predators as variables that influence the use of resources by fishes.

Habitat structural complexity and the interaction between bluegills and their prey

Effects of submersed macrophytes on ecosystem processes

Analysis of the fish faunas of Lake Opinicon and other small, cold temperate Ontario water bodies shows that the component species differ in body size, morphology, abundances, habitats, diurnal and seasonal habitat utilization patterns, diets, dietary changes with age, reproductive strategies, and population turnover rates. These differences are detailed. The number of species occurring in a lake is partly due to historic factors, the number of habitats available, and morphological, behavioral and ecological adaptations that, by channelling their owners towards alternative resources, permit species to co-occur. Diet overlap values between most species are low except for the congeneric bluegill and pump-kinseed sunfishes, where values are moderate. These are the two commonest species in the lake and other parameters must increase the ecological differences between these two species. Lake Opinicon is a highly variable ecosystem. Part of this variability stems from the seasonal nature of the environment and the fact that different resources reach their peak abundance at different times of the year. Ecological overlap levels between fish species fluctuate greatly in the course of the season as species switch from, or move on to, different resources. Population levels in different habitats also vary seasonally. Species adaptations and interaction patterns were presumably evolved over a long period; most of these adaptations undoubtedly developed before the component species colonized the lake.

Trophic and spatial interrelationships in the fish species of an Ontario temperate lake

The exotic macrophyte Myriophyllum spicatum spread widely in Lake Opinicon, Ontario, after 1974, dominating the 2.0- to 3.5-m depth zone which had previously been largely open water. In the shallow...

The introduced aquatic macrophyte, Myriophyllum spicatum, as habitat for fish and their invertebrate prey

Handling time for prey of different sizes and morphological types was studied in the largemouth bass (Micropterus salmoides). Prey ranged in size from about 1/10 to 2/3 the length of the bass. Hand...

The effect of prey morphology and size on handling time in a piscivore, the largemouth bass (Micropterus salmoides)

Analysis of the piscivore guild in fish species-rich lake and stream systems in eastern Ontario showed the co-occurrence of three types: (1) specialists that became piscivorous at the age of a few weeks (Esox, Micropterus); (2) ‘secondary’ piscivores that are fish-eaters only later in life (Perca, Ambloplites); and (3) species in which fish consumption is limited to taking some larvae (Lepomis macrochirus). In the first group the basic series of dietary shifts that characterize many long-lived fish (i.e. zooplankton followed by small invertebrates then large invertebrates and finally fish); is greatly accelerated. Prey size increases with growth. Overall prey selection was on the basis of body size and abundance. Most piscivores took a range of fish prey. There was little evidence of specialization at the species level. Esox and Micropterus spawn some weeks ahead of their major prey species. This is seen as adaptive. Their young harvest the larvae of the latter. The ensuing predator/prey association with growth is highly advantageous to the piscivore as prey of optimum body size are thus continually available.

The piscivore feeding guild of fishes in small freshwater ecosystems

We tested the hypothesis that fish species diversity would be positively related to resource diversity in a series of northern and southeastern Ontario lakes. At a macrospatial scale (among lakes),...

Allen Keast

Papers

Trophic and spatial interrelationships in the fish species of an Ontario temperate lake

The introduced aquatic macrophyte, Myriophyllum spicatum, as habitat for fish and their invertebrate prey

The effect of prey morphology and size on handling time in a piscivore, the largemouth bass (Micropterus salmoides)

The piscivore feeding guild of fishes in small freshwater ecosystems

Resource heterogeneity and fish species diversity in lakes