An introduction presents an overview of chemical signalling and suggests some key definitions, in particular distinguishing cues (chemicals that are used to guide behavior but haven't evolved for that purpose), pheromones (evolved as signals between members of a species, usually innate) and signature mixtures (variable chemical mixes used to identify individuals or colonies, usually learned). A short chapter covers methods, bioassays and so forth. And the longest chapter is on the perception of chemical communications, on receptors, neural structures and so forth.

The penultimate chapter covers the practical applications of pheromones to pest management and the handling of domestic animals. And the last chapter surveys what we know about human odours, the human sense of smell, and the as yet scant evidence for human pheromones ("there seem to be no unique and exclusive marker molecules to distinguish the sexes").

Otherwise, the chapters focus on particular kinds of behavior: mating, mate choice, and sexual selection; aggregation and host-marking; territorial scent marking; recognition and reproduction in social groups; foraging recruitment; alarm communications; orientation; and eavesdropping, mimicry, and disguise. Mammals and insects feature most prominently in most of this, but only because they are the best-studied groups. Mostly these chapters consist of overviews of the research literature, with occasional attempts at arbitration between discordant results or approaches. The material comes from a broad range of disciplines and journals, whose practitioners and readers don't always communicate, so the result is a valuable resource for specialists wanting to know how findings in one taxon fit into the broader picture.

This kind of compendium might have been too dull to be of broader interest, but the saving grace is Wyatt's ability to provide succinct but engaging summaries of individual studies and the intrinsic appeal of the material — natural history vignettes integrating behavior, evolution and biochemistry.

"California spiny lobsters, Panulirus interruptus, form groups of up to 30 animals in dens during the day. The aggregations are caused by attractions to chemical cues in the urine from conspecifics of either sex (Aggio & Derby 2011; Horner et al. 2006). Although vulnerable as individuals, as a group the lobsters wave their robust spiny antennae from the den opening to deter predatory fishes."

"Much less is known about colony recognition in termites but it seems to be based, as in Hymenoptera, on blends of CHCs; a similar role for genetically determined and acquired components in the colony odour is likely (Clément & Bagnères 1998; Dronnet et al. 2006; Kaib et al. 2004). Termites are capable of kind distinctions as subtle as any hymenopteran: DNA fingerprinting suggests that in polygynous and polyandrous termite colonies, workers departing from the nest to their foraging areas tend to form working parties with their kin."

"Orchids are not limited to mimicking sex pheromones. One orchid species mimics an alarm pheromone component, (Z)-11-eicosen-1-ol, of honeybees to attract hornets as pollinators (Brodmann et al. 2009) and another mimics aphid alarm pheromone to attract hoverflies (Stökl et al. 2011)."

These are often illustrated with diagrams and sometimes with small halftones. Each chapter also includes a few paragraphs recommending a handful of books or articles of broad general interest.

Pheromones and Animal Behavior assumes a fair bit of general biology, a decent grasp of evolutionary theory, and some basic biochemistry, but should be accessible to competent undergraduate biology students and anyone else with a similar background. (An appendix contains an introduction to chemical terms for non-chemists, but most of the chemical detail can easily be skipped.) Some of the highlights for me were explanations of how the mammalian and insect olfactory systems work, of the structure of pheromone plumes and how different animals follow them, and of the scent-matching hypothesis for territorial scent marking (that it allows intruders to recognise an owner). Individual chapters could be read by themselves: the chapter on humans is probably the leading candidate here.

Disclaimer: Tristram Wyatt is a colleague in a cycling advocacy organisation.

November 2014

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