pyrrolizidine oxime 236 even higher than levels in wild-caught frogs from the leaf-litter site (34). The second are the tricyclic coccinelline alkaloids that have been found in several frogs/toads. The coccinellines occur as defensive substances in a variety of small beetles (see ref. 6). Thus, beetles represent a possible dietary source for coccinelline-class alkaloids in frog/toad skin. Indeed, the beetle alkaloid precoccinelline is a significant alkaloid in the skin of D. auratus raised in Panama on leaf-litter arthropods (34). The other alkaloids that were found in skin of D. auratus raised on leaf-litter arthropods are three other tricyclic alkaloids, perhaps of the coccinelline-class but of unknown structure, two 1,4-disubstituted quinolizidines, a gephyrotoxin, a decahydroquinoline, and several histrionicotoxins. With the exception of the pyrrolizidine oxime 236, skin levels of the various alkaloids in the captive-raised frogs were low compared to levels of alkaloids in wild-caught frogs from the leaf-litter collection site or from the parental stock of D. auratus on a nearby island (34). Individual variation in wild-caught frogs appears significant, which complicates the comparisons. However, the lack of any pumiliotoxins and the relatively low levels or absence of decahydroquinolines and histrionicotoxins in the captive-raised frogs suggests that dietary sources for these alkaloids have been missed in the paradigm using large funnels to collect the arthropods from the leaf litter.

In summary, poisons used in chemical defense are widespread in nature. In amphibians, the defensive substances seem to be elaborated by the amphibian in the case of amines, peptides, proteins, bufadienolides, and the salamander alkaloids of the samandarine class. For the tetrodotoxin class of water-soluble alkaloids, the origin is unclear, but symbiotic bacteria have been suggested for marine organisms (4). For the so-called dendrobatid alkaloids, a dietary source now appears a likely explanation for the lack of skin alkaloids in dendrobatid frogs raised in captivity. Certainly, dendrobatid frogs of the dendrobatid genera Phyllobates, Dendrobates, and Epipedobates, which in the wild contain skin alkaloids, have highly efficient systems for accumulating selectively into skin a variety of dietary alkaloids (25, 34). A biological system for sequestration of alkaloids for chemical defense finds precedence in the transfer of pyrrolizidine alkaloids from plants via aphids to ladybug beetles (35). Accumulation of cantharidins in muscle of ranid frogs after feeding on beetles has been documented (36). Frogs of the dendrobatid genus Colostethus, which in the wild do not contain skin alkaloids, do not accumulate dietary alkaloids (25).

The proposal that all alkaloids found in skin glands of dendrobatid frogs and used in chemical defense against predators have a dietary origin leads to many questions. First, the profile of alkaloids has been found in many instances to be characteristic of a species or a population.



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