The nitrogen content of a plant is only one of the many plant characteristics that are vitally important to herbivores. However, because of its central role in all metabolic processes as well as in cellular structure and genetic coding, nitrogen is a critical element in the growth of all organisms. Supplementary N often elicits enhanced health, growth, reproduction, and survival in many organisms. This suggests that N is a limiting factor. Since N makes up a large portion of the earth's atmosphere (about 78%), the problem is not an absolute but a relative shortage-that is, a scarcity of usable or metaboliza­ ble N during critical growth periods (159, 328). Plants encounter shortages of inorganic nitrogen (nitrate and/or ammonium ions); animals experience shortages of organic nitrogen (specific proteins and/or amino acids). This article reviews and examines the evidence (a) that N is scarce and perhaps a limiting nutrient for many herbivores, and (b) that in response to this selection pressure, many herbivores have evolved specific behavioral, morphological, physiological, and other adaptations to cope with and uti­ lize the ambient N levels of their normal haunts. McNeill & Southwood (201) and White (328) have also reviewed these general questions. There­ fore, this review explores additional evidence and further develops the fundamental arguments. The review is organized into three major divisions. The first focuses on important sources of variation in plant N (seasonal and ontogenetic trends, different tissues and species, etc) because such variation may be the basis

Herbivory in relation to plant nitrogen content

▪ Abstract Floral evolution has often been associated with differences in pollination syndromes. Recently, this conceptual structure has been criticized on the grounds that flowers attract a broader spectrum of visitors than one might expect based on their syndromes and that flowers often diverge without excluding one type of pollinator in favor of another. Despite these criticisms, we show that pollination syndromes provide great utility in understanding the mechanisms of floral diversification. Our conclusions are based on the importance of organizing pollinators into functional groups according to presumed similarities in the selection pressures they exert. Furthermore, functional groups vary widely in their effectiveness as pollinators for particular plant species. Thus, although a plant may be visited by several functional groups, the relative selective pressures they exert will likely be very different. We discuss various methods of documenting selection on floral traits. Our review of the literatur...

Pollination Syndromes and Floral Specialization

Proponents of optimal foraging theory attempt to predict the behavior of animals while they are foraging; this theory is based on a number of assump­ tions ( 133 , 155 , 2 10, 23 1 ) . First, an individual's contribution to the next generation (i.e. its "fitness") depends on its behavior while foraging. This contribution may be measured genetically or culturally as the proportion of an individual's genes or "ideas", respectively, in the next generation. In the former case, the theory is simply an extension of Darwin's theory of evolution. Second, it is assumed that there should be a heritable component of foraging behavior, i.e. an animal that forages in a particular manner should be likely to have offspring that tend to forage in the same manner. This heritable compo­ nent can be either the actual foraging responses made by an animal or the rules by which an animal learns to make such responses. In other words, optimal foraging theory may apply regardless of whether the foraging behavior is learned or innate. Given these first two assumptions, it follows that the proportion of individuals in a population foraging in ways that enhance their fitness will tend to increase over time. Unless countervailed by sufficiently strong group selection (see 287, 242), foraging behavior will therefore evolve, and the average foraging behavior will increasingly come to be characterized by those characteristics that enhance individual fitness. The third assumption is that the relationship between foraging behavior and fitness is known. This relationship is usually referred to as the currency of fitness (23 1 ) . In general, any such currency will include a time scale, although in some cases it may be assumed that fitness is a function of some rate.

Optimal Foraging Theory: A Critical Review

In his introduction to this detailed survey of knowledge of insect societies, the author points out that research on insect sociology has proceeded in three phases, the natural history phase, the physiological phase and the population-biology phase. Advances in the first two phases have permitted embarkation in the third phase on a more rigorous theory of social evolution based on population genetics and writing this book, the author wished to relate the three phases of research on insects and to express insect sociology as population biology. A glossary of terms, a considerable bibliography and a general index are included. Other CABI sites 

The Insect Societies

Invasions by non-native ants are an ecologically destructive phenomenon affecting both continental and island ecosystems throughout the world. Invasive ants often become highly abundant in their introduced range and can outnumber native ants. These numerical disparities underlie the competitive asymmetry between invasive ants and native ants and result from a complex interplay of behavioral, ecological, and genetic factors. Reductions in the diversity and abundance of native ants resulting from ant invasions give rise to a variety of direct and indirect effects on non-ant taxa. Invasive ants compete with and prey upon a diversity of other organisms, including some vertebrates, and may enter into or disrupt mutualistic interactions with numerous plants and other insects. Experimental studies and research focused on the native range ecology of invasive ants will be especially valuable contributions to this field of study.

The Causes and Consequences of Ant Invasions

Floral nectar sugar constituents in relation to pollinator type

IT has been known since ancient times that nectar, usually secreted from specialized glands within flowers, is utilized by anthophilous (flower-visiting) animals for the energy-providing sugars that it contains. It is usually assumed that protein-making materials will be obtained elsewhere, from larval feeding (Lepidoptera), from pollen (Hymenoptera—Apioideae) or (by vertebrates) from insects. However, some butterflies are known to be attracted to decaying flesh, faeces, urine and stagnant water1 and well as sweat2, while many take phloem-sap from a wound or honey-dew excreted by aphids1. Moths are known to drink fruit juices, sweat, secretions from the eyes of animals, and even mammalian blood3. Unusual feeding-behaviour is shown by butterflies of the genus Heliconius which collect pollen, steep it in nectar, and subsequently ingest the amino-acids that diffuse from the grains4.

Amino-acids in Nectar and their Evolutionary Significance

There are several sources of potential error in calculating the concentration or energy value of floral nectar. Errors resulting from confusing data become substantial with increasing concentration. The different methods of expressing sugar concentration are here clarified, and the correct methods of converting from one to the other are provided. Refractometers in use in field studies usually read on a weight per total weight basis; this is recommended as the mode of statement. The perils of oversimplifying conversions from this mode, as is often done, are pointed out.

On the calculation of sugar concentration in flower nectar.

Pollen grains of 990 species were examined and measured to test four predictions: (1) "Primitive" angiosperms will have starch-containing pollen; more advanced families will have starchless pollen. (2) Where Hymenoptera and Diptera use pollen nutritionally, there will be selection of starchless (oilcontaining) pollen, particularly where pollen is the only reward for visitors. (3) Conversely, in autogamous species, anemophilous species, and those pollinated by Lepidoptera or birds, the energetically more economical accumulation of starch will be seen. (4) Small pollen grains will be more likely to be starchless (oil-rich); starch-containing grains will tend to be larger. Prediction 1 is only partially supported by the results; contemporary representatives of primitive families often show starchless grains, and so do most "advanced" families. Strong intra-familial resemblances are found (and some between groups of families), but other families show mixtures of "starchy" and "starchless" species. The latter are more likely to show a fit between food reserve and pollination system. Prediction 2 is well supported. Prediction 3 is supported in that autogamous and anemophilous species show relatively high proportions of starchy grains, but the picture is complicated for them by apparent selection for starchlessness (oil-richness) in small pollen grains (prediction 4), and for the lepidopteran- and bird-pollinated species by selection for large starchless (or very large starchy grains) in species

Starch in Angiosperm Pollen Grains and Its Evolutionary Significance

Plants pollinated and dispersed by different groups of birds offer different kinds of sugars in nectar and fruit pulp. The preferences and physiological traits of avian pollinators and seed dispersers are broadly correlated with the sugar composition of the nectar and fruit that they feed on and appear to have influenced the evolution of the sugar composition of the rewards that plants offer. Hummingbirds prefer sucrose whereas many nectar- and fruit-eating passerines prefer glucose and fructose. Preference for hexoses in passerines seems to be associated with poor sucrose assimilation resulting from two physiological mechanisms: lack of intestinal sucrase activity and fast passage rates. Sucrase activity absence appears to be restricted to a single phylogenetic group (the sturnid-muscicapid lineage). Fast passage rates seem to be characteristic of many small frugivores and to hinder the assimilation of complex nutrients that require hydrolysis before absorption. Hummingbirds have extremely specialized digestive traits that allow them to assimilate sucrose at high rates and with extremely high efficiency. These specialized digestive traits appear not to be present in many nectar-feeding passerines.

Ecological and evolutionary implications of digestive processes: Bird preferences and the sugar constituents of floral nectar and fruit pulp

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