Thus, either the systems responsible for sequestration of alkaloids differ in selectivity among different species and/or populations of dendrobatid frogs or the small arthropod fauna presenting itself and used as a diet by different species and/or populations varies even within a small island. The latter appears more likely. It was noted that the dendrobatid frogs raised on leaf-litter in Panama shared more alkaloids with a population of D. auratus from the leaf-litter site than they did with the parental population from a nearby island (34). The second major question concerns what small insects or other arthropods contain such toxic and/or unpalatable alkaloids as the batrachotoxins, the pumiliotoxins, and the histrionicotoxins, the decahydroquinolines, the 5,8-disubstituted indolizidines, the 1,4-disubstituted quinolizidines, and epibatidine. It is remarkable that such small, presumably distasteful arthropods have escaped the attention of researchers. Whether frogs intent on sequestering defensive alkaloids seek out such prey is unknown. With regard to the frogs/toads from the Madagascan family Mantellidae, the Australian family Myobatrachidae and the South American genus Melanophryniscus of the family Bufonidae, which also contain many of the dendrobatid alkaloids, it is unknown whether sequestering systems are present or even whether captive-raised frogs will lack skin alkaloids. If such systems are present, then it is remarkable from an evolutionary standpoint that such unrelated lineages of toads/frogs have independently developed systems for sequestering alkaloids into skin glands from a diet of small, presumably noxious insects for use by the toad/frog in chemical defense.
Poisons are common in nature, where they often serve the organism in chemical defense. Such poisons either are produced de novo or are sequestered from dietary sources or symbiotic organisms. Among vertebrates, amphibians are notable for the wide range of noxious agents that are contained in granular skin glands. These compounds include amines, peptides, proteins, steroids, and both water-soluble and lipid-soluble alkaloids. With the exception of the alkaloids, most seem to be produced de novo by the amphibian. The skin of amphibians contains many structural classes of alkaloids previously unknown in nature. These include the batrachotoxins, which have recently been discovered to also occur in skin and feathers of a bird, the histrionicotoxins, the gephyrotoxins, the decahydroquinolines, the pumiliotoxins and homopumiliotoxins, epibatidine, and the samandarines. Some amphibian skin alkaloids are clearly sequestered from the diet, which consists mainly of small arthropods. These include pyrrolizidine and indolizidine alkaloids from ants, tricyclic coccinellines from beetles, and pyrrolizidine oximes, pre-