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

Bateman’s principle states that male fitness is usually limited by the number of matings achieved, while female fitness is usually limited by the resources available for reproduction. When applied to flowering plants this principle leads to the expectation that pollen limitation of fruit and seed set will be uncommon. However, if male searching for mates (including pollen dissemination via external agents) is not sufficiently successful, then the reproductive success of both sexes (or both sex functions in hermaphroditic plants) will be limited by number of matings rather than by resources, and Bateman’s principle cannot be expected to apply. Limitation of female success due to inadequate pollen receipt appears to be a common phenomenon in plants. Using published data on 258 species in which fecundity was reported for natural pollination and hand pollination with outcross pollen, I found significant pollen limitation at some times or in some sites in 159 of the 258 species (62%). When experiments were performed multiple times within a growing season, or in multiple sites or years, the statistical significance of pollen limitation commonly varied among times, sites or years, indicating that the pollination environment is not constant. There is some indication that, across species, supplemental pollen leads to increased fruit set more often than increased seed set within fruits, pointing to the importance of gamete packaging strategies in plant reproduction. Species that are highly self-incompatible obtain a greater benefit relative to natural pollination from artificial application of excess outcross pollen than do self-compatible species. This suggests that inadequate pollen receipt is a primary cause of low fecundity rates in perennial plants, which are often self-incompatible. Because flowering plants often allocate considerable resources to pollinator attraction, both export and receipt of pollen could be limited primarily by resource investment in floral advertisement and rewards. But whatever investment is made is attraction, pollinator behavioral stochasticity usually produces wide variation among flowers in reproductive success through both male and female functions. In such circumstances the optimal deployment of resources among megaspores, microspores, and pollinator attraction may often require more flowers or more ovules per flower than will usually be fertilized, in order to benefit from chance fluctuations that bring in large number of pollen grains. Maximizing seed set for the entire plant in a stochastic pollination environment might thus entail a packaging strategy for flower number or ovule number per flower that makes pollen limitation of fruit or seed set likely. Pollen availability may limit female success in individual flowers, entire plants (in a season or over a lifetime), or populations. The appropriate level must be distinguished depending on the nature of the question being addressed.

Bateman's principle and plant reproduction: The role of pollen limitation in fruit and seed set

By analyzing 296 published and unpublished data sets describing annual variation in seed output by 144 species of woody plants, this article addresses the following questions. Do plant species naturally fall into distinct groups corresponding to masting and nonmasting habits? Do plant populations generally exhibit significant bimodality in annual seed output? Are there significant relationships between annual variability in seed production and pollination and seed dispersal modes, as predicted from economy of scale considerations? We failed to identify distinct groups of species with contrasting levels of annual variability in seed output but did find evidence that most polycarpic woody plants seem to adhere to alternating supra-annual schedules consisting of either high or low reproduction years. Seed production was weakly more variable among wind-pollinated taxa than animal-pollinated ones. Plants dispersed by mutualistic frugivores were less variable than those dispersed by either inanimate means or animals that predominantly behave as seed predators. We conclude that there are no objective reasons to perpetuate the concept of mast fruiting in the ecological literature as a shorthand to designate a distinct biological phenomenon. Associations between supra-annual variability in seed output and pollination and seed dispersal methods suggest the existence of important reproductive correlates that demand further investigation.

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Annual variability in seed production by woody plants and the masting concept: reassessment of principles and relationship to pollination and seed dispersal.

All individuals of all known populations of Banksia goodii were assessed for seed production. Small populations produced no or only a few seeds per unit canopy area. Effects of population size on seed production per unit area and seed production per plant were present over the whole range of population sizes, indicating that even in large populations seed production may still not be at its maximum. Resource differences could not explain this disproportionate decrease in seed production with decline in population size, because there were no differences in soil properties and understorey or overstorey cover between the small and large populations. Although plants in small and large populations were similar in size, seed production per plant was much lower in small populations. This was not because plants in small populations produced fewer cones but because the fraction of these cones that was fertile was much lower. Five of the nine smallest populations (<200 m2) produced no fertile cones over the last 10 years. The number of seeds per fertile cone did not depend on population size. The results are discussed in relation to pollination biology.

Population fragmentation may reduce fertility to zero in Banksia goodii — a demonstration of the Allee effect

Several taxa of the Australian Proteaceae have invaded South African fynbos and require costly management programmes to bring under control. Banksia spp. have been introduced only recently to fynbos regions. The invasive potential of Banksia ericifolia (Proteaceae) was investigated by comparing its recruitment potential with that of an indigenous proteaceous shrub, Leucadendron laureolum. Both species are overstorey shrubs that are killed by fire and rely on canopy-stored seeds (serotiny) for recruitment. Eight year old B. ericifolia shrubs produced an average of 16 500 seeds per plant, which is thirty times more than the average of 570 seeds produced by 10 year old L. laureolum shrubs. The seed bank of B. ericifolia was not only larger than that of L. laureolum (1098 vs 525 viable seeds m−2 projected canopy cover), but also considerably larger than that described for the species in its native environment (200–330 seeds m−2 in a 9 year old stand north of Sydney). Leucadendron laureoleum released most of its seed a few days after the cones were burnt, whereas seed release in B. ericifolia was spread over 12 weeks. The seeds of B. ericifolia had lower wingloading and fall rates than L. laureolum and were dispersed over greater distances. The relative seedling growth rates of the two species were very similar (0.03 g per day), but below-ground biomass was greater and proteoid roots were more developed in B. ericifolia seedlings than in L. laureolum after 100 days. Four year old B. ericifolia plants growing in the field had attained over twice the height of indigenous pro-teoids and accumulated up to 10 times the fresh biomass of L. xanthoconus, a species which is ecologically similar to L. laureolum. The Bioclimatic Prediction System (BIOCLIM) was used to create a bioclimatic profile of B. ericifolia and identify climatically suitable areas in the Cape Province. Results show that its potential distribution covers most fynbos areas in the southwestern Cape. It is concluded that B. ericifolia has the potential to be highly invasive in fynbos.

The invasive potential of Australian banksias in South African fynbos: A comparison of the reproductive potential of Banksia ericifolia and Leucadendron laureolum

Patterns of flowering, levels of fruit set and pollination limitation of fruit set were examined over four years in a group of four co-occurring Banksia species (Proteaceae) in south-eastern Australia

B. J. Copland

Papers

Seasonal variation in flowering intensity and pollination limitation of fruit set in four co-occurring Banksia species