title: The relationship between poison frog chemical defenses and age, body size, and sex.
abstract: BACKGROUND
  Amphibians secrete a wide diversity of chemicals from skin glands as defense against predators, parasites, and pathogens. Most defensive chemicals are produced endogenously through biosynthesis, but poison frogs sequester lipophilic alkaloids from dietary arthropods. Alkaloid composition varies greatly, even among conspecific individuals collected at the same time and place, with some individuals having only a few micrograms of one or a few alkaloids and others possessing >1 mg of >30 alkaloids. The paucity of alkaloids in juveniles and their abundance in adults suggests that alkaloids accumulate over time; however, alkaloid diversity is highly variable among adult poison frogs and has never been studied in relation to individual age. Using skeletochronology to infer individual ages and gas chromatography-mass spectrometry and vapor phase Fourier-transform infrared spectral analysis to identify the defensive chemicals of 63 individuals, we tested the relationship between defensive chemicals and age, size, and sex in the Brazilian red-belly toad, Melanophryniscus moreirae, a poison frog that possesses both sequestered alkaloids and the biosynthesized indolealkylamine bufotenine.
  
  
  RESULTS
  Adult females were, on average, older and larger than adult males. Juveniles were smaller but not necessarily younger than adults and possessed bufotenine and 18 of the 37 alkaloids found in adults. Alkaloid richness was positively related to age, but not size, whereas the quantities of sequestered alkaloids and bufotenine were positively related to size, but not age. Defensive chemicals were unrelated to sex, independent of size.
  
  
  CONCLUSIONS
  The relationship between alkaloid richness and age appears to result from the gradual accumulation of alkaloids over a frog's lifetime, whereas the relationship between the quantity of defensive chemicals and size appears to be due to the greater storage capacity of larger individuals. The decoupling of age and size effects increases the amount of individual variation that can occur within a population, thereby possibly enhancing anti-predator efficacy. Further, given that both richness and quantity contribute to the overall chemical defense of individual frogs, our results suggest that older, larger individuals are better defended than younger, smaller ones. These considerations underscore the importance of including age in studies of the causes and consequences of variation in poison frog chemical defenses.
authors.affiliation: Department of Zoology, Institute of Biosciences, University of São Paulo, 05508-090 São Paulo, São Paulo Brazil.
authors.forename: Adriana M
authors.initials: AM
authors.lastname: Jeckel
authors.affiliation: Department of Biology, John Carroll University, University Heights, OH 44118 USA.
authors.forename: Ralph A
authors.initials: RA
authors.lastname: Saporito
authors.affiliation: Department of Zoology, Institute of Biosciences, University of São Paulo, 05508-090 São Paulo, São Paulo Brazil.
authors.forename: Taran
authors.initials: T
authors.lastname: Grant
keywords: Alkaloids; Amphibia; Anura; Bufonidae; Bufotenine; Melanophryniscus; Sequestration; Skeletochronology