The evolution of "niche breadth," or "niche width," was a more popular topic in the evolutionary ecological literature of the 1960s and 1970s than it has been recently (109, 118, 120, 134, 155, 156). This review summarizes current hypotheses on the evolution of specialization and generalization and suggests areas in which future research might be rewarding. The topic is so broad that every area of biology bears on it. We cannot hope to offer an exhaustive review of evidence and in particular have slighted much of the ecological literature to emphasize genetic and evolutionary perspectives. We limit our discussion almost entirely to animals. We adopt Hutchinson's (86) representation of a population's ecological niche as an n-dimensional hypervolume, the axes of which are environmental variables or resources. Along each of these, the population displays a wide or narrow tolerance or pattern of utilization, relative to other populations or species. Specialization and generalization must be defined with reference to particular axes (e.g. temperature, range of food particle sizes). Brown (9) suggests that niche breadth along different axes is positively correlated and that this explains positive correlations across species between local abundance and breadth of geographic range. Multidimensional specialization might be expected if species arise in localized regions that differ in several ecological respects from those occupied by parent species. Cody (20), however, suggested that the breadth of habitat is negatively correlated with diet breadth among certain bird species. In practice, quantitative measurement of niche breadth can be difficult (22,

The Evolution of Ecological Specialization

Single cell responses from olfactory receptors of small ermine moths to sex-attractants

Selective factors in the evolution of host choice by phytophagous insects

Sex pheromone communication in the nine European species of small ermine moths (Yponomeuta) is reviewed in regard to the potential role of pheromones in the speciation process. Six of the nine species studied (viz.,Y. evonymellus, Y. cagnagellus, Y. padellus, Y. irrorellus, Y. plumbellus, andY. vigintipunctatus) use a mixture of (E)-11-and (Z)-11-tetradecenyl acetate in different ratios as primary pheromone components, with combinations of tetradecyl acetate, (Z)-9-tetradecenyl acetate, (Z)-11-hexadecenyl acetate and the corresponding alcohols of the acetates as additional pheromone components. Analysis of (Z)- to (E)-11-tetradecenyl acetate ratios produced by individual females of these species demonstrated significant variation among females of all species. However, the ranges of ratios produced byY. cagnagellus, Y. irrorellus, andY. plumbellus, sharing the same host-plant species, spindle tree, did not overlap. Niche separation of all six species mentioned required consideration of at least one additional pheromone component or of temporal aspects. The remaining three species,i.e. Y. malinellus, Y. mahalebellus andY. rorellus, have pheromones that differ qualitatively. Biosynthetic routes to the pheromone components identified are proposed on the basis of fatty acid pheromone precursors found in the pheromone glands. A phylogenetic tree for the genus is constructed based on allozyme frequency data and changes in pheromone composition are superimposed on this tree. We suggest that the ancestral ermine moth pheromone is a mixture of (Z)-11- and (E)-11-tetradecenyl acetate and the corresponding alcohols, and a scenario of how present-day patterns evolved is outlined. The pheromone differences among the three species using spindle tree as their host-plant might have evolved throughreproductive character displacement upon secondary contact between populations that had already diverged genetically in allopatry. Pheromone differences within the so-calledpadellus-complex (includingY. cagnagellus, Y. mahalebellus, Y. malinellus, Y. padellus, andY. rorellus) in which species might have originated sympatrically, may have evolved byreinforcing selection as these species still hybridise and produce viable offspring when confined in cages. The role of pheromones in reproductive isolation amongYponomeuta species is emphasised by (1) the function of pheromone components of some of the species as behavioural antagonists to other species, (2) the cross-attraction under experimental conditions between allochronic species with similar pheromones, and (3) the formation of hybrids in the laboratory between species that are isolated in nature by pheromone differences.

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Sex pheromones and their potential role in the evolution of reproductive isolation in small ermine moths (Yponomeutidae)

8 – Selective Factors in the Evolution of Host Choice by Phytophagous Insects*

Insect—host-plant relationships are often studied in cultivated species of economic importance. While the thorough knowledge acquired in modern plant-protection research is appreciable, the study of insects and plants under more natural conditions may add new data relevant to an understanding of the evolution of their relationship.

W. M. Herrebout

Papers

Small Ermine Moths of the Genus Yponomeuta and their Host Relationships (Lepidoptera, Yponomeutidae)