170:, which is poisonous (sequesters toad and/or firefly toxins in its nuchal gland tissues that are toxic if consumed by a predator), toxungenous (the nuchal glands are pressurized and can spray the toxins when ruptured), and venomous (toxic oral gland secretions can be injected via the teeth). Even humans can be considered facultatively poisonous, toxungenous, and venomous because they sometimes make use of toxins by all three means for research and development (e.g., biomedical purposes), agriculture (e.g., spraying insecticides), and nefarious reasons (to kill other animals, including humans).
17:
178:
Toxungen deployment offers a key evolutionary advantage compared to poisons and venoms. Poisons and venoms require direct contact with the target animal, which puts the toxin-possessing animal at risk of injury and death from a potentially dangerous enemy. Evolving the capacity to spit or spray a
118:
near their armpit) that possesses apparent toxicity. When the secretion is licked and combined with saliva, their bite introduces the secretion into a wound, which can cause sometimes severe tissue injury to conspecifics and other aggressors, thereby functioning as a venom. They can also rub the
717:
Nelsen, D. R., Nisani, Z., Cooper, A. M., Fox, G. A., Gren, E. C., Corbit, A. G., & Hayes, W. K. (2014). "Poisons, toxungens, and venoms: redefining and classifying toxic biological secretions and the organisms that employ them". Biological
Reviews, 89(2), 450-465. doi:
854:
Nelsen, D. R., Nisani, Z., Cooper, A. M., Fox, G. A., Gren, E. C., Corbit, A. G., & Hayes, W. K. (2014). "Poisons, toxungens, and venoms: redefining and classifying toxic biological secretions and the organisms that employ them". Biological
Reviews, 89(2), 450-465.
735:
Nelsen, D. R., Nisani, Z., Cooper, A. M., Fox, G. A., Gren, E. C., Corbit, A. G., & Hayes, W. K. (2014). "Poisons, toxungens, and venoms: redefining and classifying toxic biological secretions and the organisms that employ them". Biological
Reviews, 89(2), 450-465.
288:
Nelsen, D. R., Nisani, Z., Cooper, A. M., Fox, G. A., Gren, E. C., Corbit, A. G., & Hayes, W. K. (2014). "Poisons, toxungens, and venoms: redefining and classifying toxic biological secretions and the organisms that employ them". Biological
Reviews, 89(2), 450-465.
270:
Nelsen, D. R., Nisani, Z., Cooper, A. M., Fox, G. A., Gren, E. C., Corbit, A. G., & Hayes, W. K. (2014). "Poisons, toxungens, and venoms: redefining and classifying toxic biological secretions and the organisms that employ them". Biological
Reviews, 89(2), 450-465.
252:
Nelsen, D. R., Nisani, Z., Cooper, A. M., Fox, G. A., Gren, E. C., Corbit, A. G., & Hayes, W. K. (2014). "Poisons, toxungens, and venoms: redefining and classifying toxic biological secretions and the organisms that employ them". Biological
Reviews, 89(2), 450-465.
35:
that is transferred by one animal to the external surface of another animal via a physical delivery mechanism. Toxungens can be delivered through spitting, spraying, or smearing. As one of three categories of biological toxins, toxungens can be distinguished from
182:
Toxins used as toxungens can be acquired by several means. Many species synthesize their own toxins and store them within glands, but others acquire their toxins exogenously from other species. Two examples illustrate exogenous acquisition. Snakes of the genus
919:
Ismail,M., Al-Bekairi, A.M., El-Bedaiwy, A.M. & Abd-El Salam,M. A. (1993). "The ocular effect of spitting cobras: II. Evidence that cardiotoxins are responsible for the corneal opacification syndrome". Clinical
Toxicology 31, 45–62.
506:
Nekaris, K., Moore, R. S., Rode, E. J., & Fry, B. G. (2013). Mad, bad and dangerous to know: the biochemistry, ecology and evolution of slow loris venom. Journal of
Venomous Animals and Toxins including Tropical Diseases, 19, 1-10.
532:
Nekaris, K. A. I., Campera, M., Nijman, V., Birot, H., Rode-Margono, E. J., Fry, B. G., ... & Imron, M. A. (2020). Slow lorises use venom as a weapon in intraspecific competition. Current
Biology, 30(20), R1252-R1253.
224:
possess a secretion used as a venom (injected for predation and/or defense) that can also be sprayed to communicate alarm among nestmates, to mark a trail used for food gathering, or to keep their brood free of parasites.
44:, which are delivered through a wound generated by a bite, sting, or other such action. Toxungen use offers the evolutionary advantage of delivering toxins into the target's tissues without the need for physical contact.
207:, the highly toxic non-proteinaceous component of their salivary glands that can be ejected into the water to subdue nearby prey, via accumulation from food resources and/or symbiotic tetrodotoxin-producing bacteria.
157:
Animals that deploy toxungens are referred to as toxungenous. Some animals use their toxins in multiple ways, and can be classified as poisonous, toxungenous, and/or venomous. Examples include the scorpion
131:
spray a noxious and potentially injurious secretion from their anal sac when threatened. High concentrations of the spray can be toxic, with rare accounts of spray victims suffering injury and even death.
625:
Fierro, B. R., Agnew, D. W., Duncan, A. E., Lehner, A. F., & Scott, M. A. (2013). Skunk musk causes methemoglobin and Heinz body formation in vitro. Veterinary
Clinical Pathology, 42(3), 291-300.
753:
Hutchinson, D. A., Mori, A., Savitzky, A. H., Burghardt, G. M., Wu, X., Meinwald, J., & Schroeder, F. C. (2007). "Dietary sequestration of defensive steroids in nuchal glands of the Asian snake
600:
Zaks, K. L., Tan, E. O., & Thrall, M. A. (2005). Heinz body anemia in a dog that had been sprayed with skunk musk. Journal of the American Veterinary Medical Association, 226(9), 1516-1518.
416:
Berthé, R. A., De Pury, S., Bleckmann, H., & Westhoff, G. (2009). "Spitting cobras adjust their venom distribution to target distance". Journal of Comparative Physiology A, 195(8), 753-757.
441:
J.P. Dumbacher and Pruett-Jones, S. (1996). "Avian chemical defense". In: Nolan, V., Jr., and Ketterson, E. D. (Eds.), Current Ornithology, vol. 13, Plenum Press, New York (1996), pp. 137-174.
704:
Mori, A., Burghardt, G. M., Savitzky, A. H., Roberts, K. A., Hutchinson, D. A., & Goris, R. C. (2012). "Nuchal glands: a novel defensive system in snakes". Chemoecology, 22(3), 187-198.
220:
that cooperatively subdue their prey by seizing, spread-eagling, and then smearing their toxins onto the prey's surface. Toxungens can also be used for communication and hygiene. Many
403:
Brodie, E. D. & Smatresk,N. J. (1990). "The antipredator arsenal of fire salamanders: spraying of secretions from highly pressurized dorsal skin glands". Herpetologica 46, 1–7.
872:
Eisner, T., Meinwald, J., Monro, A. & Ghent, R. (1961). "Defence mechanisms of Arthropods. I. The composition and function of the spray of the whipscorpion,
344:
Eisner, T., Aneshansley, D. J., Eisner, M., Attygalle, A. B., Alsop, D. W. & Meinwald, J. (2000a). "Spray mechanism of the most primitive bombardier beetle (
820:
Sutherland, S. & Lane, W. (1969). "Toxins and mode of envenomation of the common ringed or blue-banded octopus". Medical Journal of Australia 1, 893–898.
787:
Sutherland, S. & Lane, W. (1969). "Toxins and mode of envenomation of the common ringed or blue-banded octopus". Medical Journal of Australia 1, 893–898.
386:
Sutherland, S. & Lane, W. (1969). "Toxins and mode of envenomation of the common ringed or blue-banded octopus". Medical Journal of Australia 1, 893–898.
558:
Stankowich, T., Caro, T., & Cox, M. (2011). Bold coloration and the evolution of aposematism in terrestrial carnivores. Evolution, 65(11), 3090-3099.
232:
that spray or smear their secretion onto insect prey enhance toxin penetration by including a spreading agent that additionally enhances toxicity. Some
210:
Toxungens are most commonly used for defensive purposes, but can be used in other contexts as well. Examples of toxungen use for predation include the
481:". In Alterman, L.; Doyle, G.A.; Izard, M.K (eds.). Creatures of the Dark: The Nocturnal Prosimians. New York, New York: Plenum Press. Pp. 413–424.
486:
119:
secretion on their fur or lick their offspring before stashing them in a secure location, thereby functioning potentially as a toxungen.
149:
movies as capable of spitting a toxic secretion, no evidence exists to suggest that any dinosaur possessed either a toxungen or venom.
16:
327:
caterpillars (Lep., Saturniidae) spray an irritant secretion from defensive glands. Journal of Chemical Ecology 20, 2127–2138.
114:), which comprise several species of nocturnal primates in Southeast Asia, produce a secretion in their brachial glands (a
804:
Yamate, Y., Takatani, T., & Takegaki, T. (2021). "Levels and distribution of tetrodotoxin in the blue-lined octopus
837:
Richard, F. J., Fabre, A. & Dejean, A. (2001). "Predatory behavior in dominant arboreal ant species: the case of
944:
890:
228:
Because of their unique delivery system, toxungens may be chemically designed to better penetrate body surfaces.
808:
in Japan, with special reference to within-body allocation." Journal of Molluscan Studies, 87(1), eyaa042. doi:
99:, plant materials, and even manufactured pesticides. Some of the described substances may be toxic, at least to
821:
788:
387:
160:
925:
146:
583:
Wood, W. F. (1999). The history of skunk defensive secretion research. The Chemical Educator, 4(2), 44-50.
164:, which is both toxungenous (can spray its toxins) and venomous (can inject its toxins), and the snake
692:
142:
889:
Prestwich, G. D. (1984). "Defense-mechanisms of termites". Annual Review of Entomology 29, 201–232.
877:
671:
370:
311:
211:
199:
166:
214:, which can squirt its secretion into water to immobilize or kill its prey, and ants of the genus
40:, which are passively transferred via ingestion, inhalation, or absorption across the skin, and
778:, have chemical preference for a skin toxin of toads?" Current Herpetology, 40(1), 1-9. doi: .
634:
609:
567:
542:
516:
490:
482:
457:
Morozov, N. S. (2015). Why do birds practice anting? Biology Bulletin Reviews, 5(4), 353-365.
425:
921:
705:
306:
Koopowitz, H. (1970). "Feeding behaviour and the role of the brain in the polyclad flatworm,
719:
626:
601:
584:
559:
534:
508:
458:
442:
417:
236:
have modified their secretion so that the cardiotoxins are more injurious to eye membranes.
650:
949:
84:
842:
758:
473:
Alterman, L. (1995). "Toxins and toothcombs: potential allospecific chemical defenses in
349:
233:
938:
907:
563:
216:
137:
87:, or cloacas, and some anoint themselves with heterogenously acquired chemicals from
204:
902:
Eisner, T., Rossini, C. & Eisner, M. (2000b). "Chemical defense of an earwig (
691:
scorpions: risk assessment and venom metering". Animal Behaviour, 81(3), 627-633.
666:
Nisani, Z., & Hayes, W. K. (2015). "Venom-spraying behavior of the scorpion
605:
446:
365:
Nisani, Z., & Hayes, W. K. (2015). "Venom-spraying behavior of the scorpion
221:
115:
92:
65:
328:
179:
toxic secretion can reduce this risk by delivering the toxins from a distance.
538:
462:
421:
229:
185:
128:
124:
107:
100:
69:
856:
737:
290:
272:
254:
860:
762:
741:
723:
675:
512:
374:
353:
332:
294:
276:
258:
88:
61:
53:
876:(Lucas)(Arachnida, Pedipalpida)". Journal of Insect Physiology 6, 272–298.
825:
792:
638:
613:
571:
546:
520:
429:
391:
588:
83:, as a number of species deploy defensive secretions from their stomachs,
809:
494:
73:
841:
sp. (Hymenoptera: Formicidae)". Journal of Insect Behavior 14, 271–282.
757:". Proceedings of the National Academy of Sciences, 104(7), 2265-2270.
404:
194:
57:
630:
96:
37:
774:
Fukuda, M., & Mori, A. (2021). "Does an Asian natricine snake,
120:
41:
32:
15:
190:
80:
687:
Nisani, Z., & Hayes, W. K. (2011). "Defensive stinging by
670:(Arachnida: Buthidae)". Behavioural Processes, 115, 46-52.
369:(Arachnida: Buthidae)". Behavioural Processes, 115, 46-52.
52:
Toxungens have evolved in a variety of animals, including
31:
comprises a secretion or other body fluid of one or more
20:
Experimentally induced toxungen spraying by the scorpion
106:
Toxungen use might also exist in several mammal groups.
189:
sequester their nuchal gland toxins from their diet of
348:)". Journal of Experimental Biology 203, 1265–1275.
655:." At Blogosaur, Phillip J. Curie Dinosaur Museum
8:
103:, which would qualify them as toxungens.
891:doi:10.1146/annurev.en.29.010184.001221
245:
822:doi:10.5694/j.1326-5377.1969.tb49778.x
789:doi:10.5694/j.1326-5377.1969.tb49778.x
388:doi:10.5694/j.1326-5377.1969.tb49778.x
7:
405:https://www.jstor.org/stable/3892595
323:Deml, R. & Dettner, K. (1994).
153:Classification of toxin deployment
14:
693:doi:10.1016/j.anbehav.2010.12.010
878:doi:10.1016/0022-1910(61)90054-3
672:doi:10.1016/j.beproc.2015.03.002
564:10.1111/j.1558-5646.2011.01334.x
371:doi:10.1016/j.beproc.2015.03.002
312:doi:10.1016/0003-3472(70)90066-7
79:Toxungen use possibly exists in
197:, Blue-ringed octopuses (genus
651:Carter, N. Undated. "The real
310:". Animal Behavior 18, 31–35.
141:was portrayed in the original
135:Although the extinct theropod
1:
922:doi:10.3109/15563659309000373
706:doi:10.1007/s00049-011-0086-2
123:and several other members of
906:)". Chemoecology 10, 81–87.
843:doi:10.1023/A:1007845929801
759:doi:10.1073/pnas.0610785104
606:10.2460/javma.2005.226.1516
447:10.1007/978-1-4615-5881-1_4
966:
350:doi:10.1242/jeb.203.8.1265
908:doi:10.1007/s000490050011
539:10.1016/j.cub.2020.08.084
463:10.1134/S2079086415040076
422:10.1007/s00359-009-0451-6
874:Mastigoproctus giganteus
689:Parabuthus transvaalicus
668:Parabuthus transvaalicus
367:Parabuthus transvaalicus
161:Parabuthus transvaalicus
22:Parabuthus transvaalicus
513:10.1186/1678-9199-19-21
147:Jurassic World Dominion
806:Hapalochlaena fasciata
329:doi:10.1007/BF02066249
174:Evolution and function
48:Taxonomic distribution
25:
857:doi:10.1111/brv.12062
738:doi:10.1111/brv.12062
589:10.1007/s00897990286a
291:doi:10.1111/brv.12062
273:doi:10.1111/brv.12062
255:doi:10.1111/brv.12062
19:
308:Planocera gilchristi
776:Rhabdophis tigrinus
755:Rhabdophis tigrinus
212:blue-ringed octopus
167:Rhabdophis tigrinus
346:Metrius contractus
26:
945:Animal physiology
720:10.1111/brv.12062
631:10.1111/vcp.12074
487:978-0-306-45183-6
33:biological toxins
957:
929:
917:
911:
900:
894:
887:
881:
870:
864:
852:
846:
835:
829:
818:
812:
802:
796:
785:
779:
772:
766:
751:
745:
733:
727:
715:
709:
702:
696:
685:
679:
664:
658:
648:
642:
623:
617:
598:
592:
581:
575:
556:
550:
530:
524:
504:
498:
471:
465:
455:
449:
439:
433:
414:
408:
401:
395:
384:
378:
363:
357:
342:
336:
321:
315:
304:
298:
286:
280:
268:
262:
250:
85:uropygial glands
965:
964:
960:
959:
958:
956:
955:
954:
935:
934:
933:
932:
918:
914:
901:
897:
888:
884:
871:
867:
853:
849:
836:
832:
819:
815:
803:
799:
786:
782:
773:
769:
752:
748:
734:
730:
716:
712:
703:
699:
686:
682:
665:
661:
649:
645:
624:
620:
599:
595:
582:
578:
557:
553:
531:
527:
505:
501:
472:
468:
456:
452:
440:
436:
415:
411:
402:
398:
385:
381:
364:
360:
343:
339:
322:
318:
305:
301:
287:
283:
269:
265:
251:
247:
242:
234:Spitting cobras
176:
155:
50:
12:
11:
5:
963:
961:
953:
952:
947:
937:
936:
931:
930:
912:
904:Doru taeniatum
895:
882:
865:
847:
830:
813:
797:
780:
767:
746:
728:
710:
697:
680:
659:
643:
618:
593:
576:
551:
525:
499:
466:
450:
434:
409:
396:
379:
358:
337:
316:
299:
281:
263:
244:
243:
241:
238:
200:Hapalochlaeana
175:
172:
154:
151:
49:
46:
13:
10:
9:
6:
4:
3:
2:
962:
951:
948:
946:
943:
942:
940:
927:
923:
916:
913:
909:
905:
899:
896:
892:
886:
883:
879:
875:
869:
866:
862:
858:
851:
848:
844:
840:
839:Crematogaster
834:
831:
827:
823:
817:
814:
810:
807:
801:
798:
794:
790:
784:
781:
777:
771:
768:
764:
760:
756:
750:
747:
743:
739:
732:
729:
725:
721:
714:
711:
707:
701:
698:
694:
690:
684:
681:
677:
673:
669:
663:
660:
656:
654:
653:Dilophosaurus
647:
644:
640:
636:
632:
628:
622:
619:
615:
611:
607:
603:
597:
594:
590:
586:
580:
577:
573:
569:
565:
561:
555:
552:
548:
544:
540:
536:
529:
526:
522:
518:
514:
510:
503:
500:
496:
492:
488:
484:
480:
476:
470:
467:
464:
460:
454:
451:
448:
444:
438:
435:
431:
427:
423:
419:
413:
410:
406:
400:
397:
393:
389:
383:
380:
376:
372:
368:
362:
359:
355:
351:
347:
341:
338:
334:
330:
326:
325:Attacus atlas
320:
317:
313:
309:
303:
300:
296:
292:
285:
282:
278:
274:
267:
264:
260:
256:
249:
246:
239:
237:
235:
231:
226:
223:
222:hymenopterans
219:
218:
217:Crematogaster
213:
208:
206:
202:
201:
196:
192:
188:
187:
180:
173:
171:
169:
168:
163:
162:
152:
150:
148:
144:
143:Jurassic Park
140:
139:
138:Dilophosaurus
133:
130:
126:
122:
117:
113:
109:
104:
102:
101:ectoparasites
98:
94:
90:
86:
82:
77:
75:
71:
67:
63:
59:
55:
47:
45:
43:
39:
34:
30:
23:
18:
915:
903:
898:
885:
873:
868:
850:
838:
833:
816:
805:
800:
783:
775:
770:
754:
749:
731:
713:
700:
688:
683:
667:
662:
652:
646:
621:
596:
579:
554:
528:
502:
479:Perodicticus
478:
474:
469:
453:
437:
412:
399:
382:
366:
361:
345:
340:
324:
319:
307:
302:
284:
266:
248:
227:
215:
209:
205:tetrodotoxin
198:
184:
181:
177:
165:
159:
156:
136:
134:
111:
108:Slow lorises
105:
93:caterpillars
78:
51:
28:
27:
21:
116:scent gland
66:cephalopods
939:Categories
475:Nycticebus
240:References
230:Arthropods
203:) acquire
186:Rhabdophis
129:Mustelidae
125:Mephitidae
112:Nycticebus
89:millipedes
70:amphibians
195:fireflies
62:arachnids
54:flatworms
924:. PMID:
861:24102715
859:. PMID:
824:. PMID:
791:. PMID:
763:17284596
761:. PMID:
742:24102715
740:. PMID:
724:24102715
722:. PMID:
676:25748565
674:. PMID:
639:24033800
614:15882003
572:22023577
547:33080192
521:31074351
495:33441731
430:19462171
390:. PMID:
375:25748565
373:. PMID:
354:10729276
352:. PMID:
333:24242735
331:. PMID:
295:24102715
293:. PMID:
277:24102715
275:. PMID:
259:24102715
257:. PMID:
74:reptiles
29:Toxungen
926:8433415
826:4977737
793:4977737
392:4977737
193:and/or
110:(genus
97:beetles
58:insects
38:poisons
950:Toxins
637:
612:
570:
545:
519:
493:
485:
428:
121:Skunks
72:, and
42:venoms
191:toads
81:birds
635:PMID
610:PMID
568:PMID
543:PMID
517:PMID
491:OCLC
483:ISBN
477:and
426:PMID
145:and
127:and
627:doi
602:doi
585:doi
560:doi
535:doi
509:doi
459:doi
443:doi
418:doi
941::
633:.
608:.
566:.
541:.
515:.
489:.
424:.
95:,
91:,
76:.
68:,
64:,
60:,
56:,
928:.
910:.
893:.
880:.
863:.
845:.
828:.
811:.
795:.
765:.
744:.
726:.
708:.
695:.
678:.
657:.
641:.
629::
616:.
604::
591:.
587::
574:.
562::
549:.
537::
523:.
511::
497:.
461::
445::
432:.
420::
407:.
394:.
377:.
356:.
335:.
314:.
297:.
279:.
261:.
24:.
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
