might better be categorized as ''noxious" rather than "poisonous," although at high enough dosages all of these compounds would be poisons.

Toads and salamanders have been considered noxious creatures for centuries and indeed the majority of amphibians have now been found to contain noxious and sometimes poisonous substances in their skin secretions (8). The type of biologically active substance found in amphibians appears to have phylogenetic significance. Thus, indole alkylamines are typically present in high levels in bufonid toads of the genus Bufo, phenolic amines in leptodactylid frogs, vasoactive peptides in a great variety in hylid frogs, particularly of genus Phylomedusa (10), and bufadienolides in parotoid glands and skins of bufonid toads of the genus Bufo as well as in skin of related bufonid genera Atelopus and probably Dendrophryniscus and Melanophryniscus (11). The water-soluble alkaloid tetrodotoxin occurs in newts of the family Salamandridae, toads of the brachycephalid genus Brachycephalus and the bufonid genus Atelopus, and now in one frog species of the dendrobatid genus Colostethus (12). Lipophilic alkaloids have been found only in salamanders of the salamandrid genus Salamandra; in frogs of the dendrobatid genera Phyllobates, Dendrobates, Epipedobates, and Minyobates, the mantellid genus Mantella and the myobatrachid genus Pseudophryne; and in toads of the bufonid genus Melanophryniscus. More than 70 other genera from 11 amphibian families do not have skin alkaloids. The distribution of various lipophilic alkaloids in amphibians is given in Table 1 and structures are shown in Figure 1.

The origin and function of poisons and noxious substances found in amphibians are only partially known. The high levels of amines, including such well-known biogenic amines as serotonin, histamine, and tyramine and derivatives thereof, found in skin of various toads and frogs (8), undoubtedly are synthesized by the amphibian itself. They are stored in granular skin glands for secretion upon attack by a predator, whereupon their well-known irritant properties on buccal tissue would serve well in chemical defense. The high levels of vasoactive peptides, such as bradykinin, sauvagine, physaelaemin, caerulein, bombesin, dermorphins, etc., presumably also serve in defense against predators, although many, including the magainins, have high activity as antimicrobials (13) and thus might also serve as a chemical defense against microorganisms. Skin secretions from one hylid frog are used in "hunting magic" folk rituals by Amazonian Indians; such secretions contain many vasoactive peptides (10) and a peptide, adenoregulin, that can affect central adenosine receptors (14). The peptides of frog skin are synthesized by the amphibian and indeed additional peptides are being deduced based on cDNAs for their precursors (15). The various hemolytic proteins of certain amphib-



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