136:
dry savanna and semi-arid environments, particularly the hotter, open areas of sub-Saharan Africa. It uses its venom primarily as a means of defense. The spitting cobra has the ability to direct venom up to three metres away from its location. The trajectory of the venom that the cobra sprays is not at random. The spitting cobra has evolved to aim the venom that it spits into or as close as possible to the antagonistâs face and eyes. The cobras can measure and adjust the venom dosage being delivered, based on the size and relative distance of their target, in order to ensure the highest envenomation potential possible.
113:
76:
121:
157:
actions may assist in propelling the venom, but research does not support the hypothesis that they play any major functional part except possibly enhancing the threatening effect of the behavior. When cornered, some species "spit" their toxungen as far as 2 m (6.6 ft). While spitting is typically their primary form of defense, all spitting cobras can also deliver their toxin as a venom by biting.
104:, which can be sprayed on the target surface. Their ability to target and shoot venom is utilised in several different ways, self-defense being the most common instance. Studies have shown that the targets (at which the cobras shoot) are far from random; rather, spitting cobras consciously take aim, directing their spray as close as possible to the eyes and face of an aggressor.
65:
31:
189:
snakes in the Elapid clade, spitting cobras inject their venom through a bite in order to kill their prey. Spitting was evolved as a defense mechanism to deter predators, even if a
Spitting cobra blinds a threat, that is not enough to kill the attacker; therefore spitting cobras can also inject venom directly.
769:
Kazandjian, T. D.; Petras, D.; Robinson, S. D.; van Thiel, J.; Greene, H. W.; Arbuckle, K.; Barlow, A.; Carter, D. A.; Wouters, R. M.; Whiteley, G.; Wagstaff, S. C.; Arias, A. S.; Albulescu, L.-O.; Plettenberg Laing, A.; Hall, C.; Heap, A.; Penrhyn-Lowe, S.; McCabe, C. V.; Ainsworth, S.; da Silva, R.
184:
There are many reasons why an organism undergoes evolution. The spitting cobra predominantly uses its venom for defensive purposes. It has evolved the ability to spit in order to minimize the risk of contact during an altercation. Even though venomous snakes are very dangerous, they often times are
135:
Spitting cobras belong to the
Elapidae, which comprises snakes including cobras, mambas, coral snakes, kraits, taipans, death adders and sea snakes. Many snakes use their venom as both a defensive and predatory mechanism, and this includes the spitting cobras. The spitting cobra typically inhabits
188:
The spitting cobra also has the ability to inject venom through a bite. In fact, a spitting cobra ejects more venom during a bite than spitting venom. Even though the spitting cobra has the ability to spray venom at potential threats, spitting is not the way they kill their prey. Just like most
156:
The toxungen sprays out in distinctive geometric patterns when muscles squeeze the glands to squirt it out through forward-facing holes near the tips of the fangs. Individuals of some species of spitting cobras make hissing exhalations/lunging movements of their heads when "spitting", and such
838:
Hayes, William K.; Herbert, Shelton S.; Harrison, James R.; Wiley, Kristen L. (September 2008). "Spitting versus Biting: Differential Venom Gland
Contraction Regulates Venom Expenditure in the Black-Necked Spitting Cobra, Naja nigricollis nigricollis".
176:. In each of these three events, the venom convergently evolved to be more effective at creating pain in mammals to serve as a better deterrent, with each of the three evolutions roughly correlating with the evolution and/or arrival of early hominins.
144:
The spat toxungen is generally harmless on intact mammalian skin (although contact can result in delayed blistering of the area), but can cause permanent blindness if introduced to the eye; if left untreated it may cause
666:
Panagides, Nadya; Jackson, Timothy N. W.; Ikonomopoulou, Maria P.; Arbuckle, Kevin; Pretzler, Rudolf; Yang, Daryl C.; Ali, Syed A.; Koludarov, Ivan; Dobson, James; Sanker, Brittany; Asselin, Angelique (2017-03-13).
99:
directly from their fangs. This substance has two functions, with the first being as venom that can be absorbed via the victimâs eyes, mouth, or nose (or any mucous membrane or existing wound), and secondly as a
185:
not left injury free after a battle with a predator or larger animal. Being able to spit a toxin from afar dramatically decreases the chances of a spitting cobra getting injured in a squabble.
770:
R.; Dorrestein, P. C.; Richardson, M. K.; GutiĂ©rrez, J. M.; Calvete, J. J.; Harrison, R. A.; Vetter, I.; Undheim, E. A. B.; WĂŒster, W.; Casewell, N. R. (2021).
535:
Berthé, Ruben Andres; de Pury, Stéphanie; Bleckmann, Horst; Westhoff, Guido (2009-08-01). "Spitting cobras adjust their venom distribution to target distance".
626:
Berthé, Ruben Andres. Spitting behaviour and fang morphology of spitting cobras. Doctoral thesis, Rheinischen
Friedrich-Wilhelms-UniversitÀt, Bonn, April 2011.
476:
Hus, Konrad Kamil; Buczkowicz, Justyna; Petrilla, VladimĂr; PetrillovĂĄ, Monika; Ćyskowski, Andrzej; LegĂĄth, Jaroslav; Bocian, Aleksandra (March 2018).
669:"How the Cobra Got Its Flesh-Eating Venom: Cytotoxicity as a Defensive Innovation and Its Co-Evolution with Hooding, Aposematic Marking, and Spitting"
417:
Hus, Konrad; Buczkowicz, Justyna; Petrilla, VladimĂr; PetrillovĂĄ, Monika; Ćyskowski, Andrzej; LegĂĄth, Jaroslav; Bocian, Aleksandra (2018-03-08).
943:
636:
Rasmussen, Sara; Young, B.; Krimm, Heather (September 1995). "On the 'spitting' behaviour in cobras (Serpentes: Elapidae)".
366:
Westhoff, G.; TzschÀtzsch, K.; Bleckmann, H. (October 2005). "The spitting behavior of two species of spitting cobras".
172:
effects typical of other cobra species. The ability to spit likely evolved in cobras three times independently through
242:
727:
317:
883:
964:
331:
219:
197:âĄ: Not a âtrue spitting cobraâ, although these species have the ability to âejectâ venom, they rarely do so.
969:
296:
249:
786:
324:
233:
205:
173:
884:"Dentitional phenomena in cobras revisited: Spitting and fang structure in the Asiatic species of
902:
864:
751:
568:
399:
256:
112:
959:
856:
820:
802:
743:
708:
690:
609:
560:
552:
517:
499:
458:
440:
391:
383:
75:
771:
848:
810:
794:
735:
698:
680:
645:
599:
544:
507:
489:
448:
430:
375:
303:
226:
790:
120:
815:
703:
668:
649:
512:
477:
453:
418:
282:
53:
953:
772:"Convergent evolution of pain-inducing defensive venom components in spitting cobras"
755:
289:
275:
868:
572:
403:
310:
169:
129:
96:
17:
937:
931:
548:
494:
435:
379:
212:
165:
860:
806:
747:
694:
556:
503:
444:
387:
798:
739:
342:
161:
824:
712:
613:
564:
521:
462:
395:
685:
588:"The buccal buckle: The functional morphology of venom spitting in cobras"
64:
263:
146:
101:
921:. University of California Press, Berkeley and Los Angeles, California.
906:
88:
30:
604:
587:
116:
An alert, ready to attack
Mandalay spitting cobra (Naja mandalayensis)
852:
150:
586:
Young, B. A.; Dunlap, K.; Koenig, K.; Singer, M. (September 2004).
119:
111:
92:
74:
63:
29:
58:
2: Horizontal section through the fang at the discharge orifice.
160:
Most spitting cobras' venom/toxungen is significantly
882:
WĂŒster, Wolfgang; Thorpe, Roger S. (December 1992).
934:
of an
African red spitting cobra spraying its venom
728:"Spitting cobras' venom evolved to inflict pain"
95:that can intentionally, defensively shoot their
8:
60:3: Frontal view of the discharge orifices.
919:Snakes: The Evolution of Mystery in Nature
946:'Spitting Cobras' Sharp-Shooting Secrets"
814:
702:
684:
603:
511:
493:
452:
434:
345:have been reported to spit occasionally.
34:Schematic comparison between sections of
940:of an African red spitting cobra feeding
478:"First Look at the Venom of Naja ashei"
419:"First Look at the Venom of Naja ashei"
353:
361:
359:
357:
7:
661:
659:
52:1: Section of the whole fang in the
537:Journal of Comparative Physiology A
368:Journal of Comparative Physiology A
650:10.1111/j.1469-7998.1995.tb02743.x
25:
592:Journal of Experimental Biology
27:Common name for several snakes
1:
68:Juvenile red spitting cobra,
726:Leslie, Mitch (2021-01-21).
986:
549:10.1007/s00359-009-0451-6
495:10.3390/molecules23030609
436:10.3390/molecules23030609
380:10.1007/s00359-005-0010-8
917:Greene, Harry W. (1997)
799:10.1126/science.abb9303
740:10.1126/science.abg6859
888:(Serpentes: Elapidae)"
841:Journal of Herpetology
264:Hemachatus haemachatus
234:Naja nigricincta woodi
132:
117:
80:
72:
61:
686:10.3390/toxins9030103
123:
115:
78:
67:
33:
174:convergent evolution
130:full-face protection
791:2021Sci...371..386K
304:Naja philippinensis
638:Journal of Zoology
297:Naja mandalayensis
133:
118:
87:is any of several
81:
79:Red spitting cobra
73:
62:
36:non-spitting cobra
785:(6527): 386â390.
605:10.1242/jeb.01170
598:(20): 3483â3494.
164:, apart from the
16:(Redirected from
977:
965:Snakes of Africa
911:
910:
892:
879:
873:
872:
853:10.1670/07-076.1
835:
829:
828:
818:
776:
766:
760:
759:
723:
717:
716:
706:
688:
663:
654:
653:
633:
627:
624:
618:
617:
607:
583:
577:
576:
532:
526:
525:
515:
497:
473:
467:
466:
456:
438:
414:
408:
407:
363:
290:Naja sagittifera
243:Naja nigricollis
227:Naja nigricincta
200:African cobras:
85:"spitting" cobra
21:
985:
984:
980:
979:
978:
976:
975:
974:
950:
949:
928:
914:
890:
881:
880:
876:
837:
836:
832:
774:
768:
767:
763:
725:
724:
720:
665:
664:
657:
635:
634:
630:
625:
621:
585:
584:
580:
534:
533:
529:
475:
474:
470:
416:
415:
411:
374:(10): 873â881.
365:
364:
355:
351:
338:
311:Naja samarensis
220:Naja mossambica
195:
182:
142:
110:
59:
57:
51:
28:
23:
22:
18:Spitting cobras
15:
12:
11:
5:
983:
981:
973:
972:
970:Snakes of Asia
967:
962:
952:
951:
948:
947:
944:Discovery News
941:
935:
927:
926:External links
924:
923:
922:
913:
912:
901:(4): 424â434.
874:
847:(3): 453â460.
830:
761:
718:
655:
628:
619:
578:
543:(8): 753â757.
527:
468:
409:
352:
350:
347:
336:
335:
332:Naja sumatrana
328:
325:Naja sputatrix
321:
318:Naja siamensis
314:
307:
300:
293:
286:
279:
270:Asian cobras:
268:
267:
260:
253:
246:
239:
238:
237:
223:
216:
213:Naja katiensis
209:
194:
191:
181:
178:
141:
138:
126:Naja siamensis
109:
106:
54:sagittal plane
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
982:
971:
968:
966:
963:
961:
958:
957:
955:
945:
942:
939:
936:
933:
930:
929:
925:
920:
916:
915:
908:
904:
900:
896:
895:Herpetologica
889:
887:
878:
875:
870:
866:
862:
858:
854:
850:
846:
842:
834:
831:
826:
822:
817:
812:
808:
804:
800:
796:
792:
788:
784:
780:
773:
765:
762:
757:
753:
749:
745:
741:
737:
733:
729:
722:
719:
714:
710:
705:
700:
696:
692:
687:
682:
678:
674:
670:
662:
660:
656:
651:
647:
643:
639:
632:
629:
623:
620:
615:
611:
606:
601:
597:
593:
589:
582:
579:
574:
570:
566:
562:
558:
554:
550:
546:
542:
538:
531:
528:
523:
519:
514:
509:
505:
501:
496:
491:
487:
483:
479:
472:
469:
464:
460:
455:
450:
446:
442:
437:
432:
428:
424:
420:
413:
410:
405:
401:
397:
393:
389:
385:
381:
377:
373:
369:
362:
360:
358:
354:
348:
346:
344:
339:
334:
333:
329:
327:
326:
322:
320:
319:
315:
313:
312:
308:
306:
305:
301:
299:
298:
294:
292:
291:
287:
285:
284:
283:Naja kaouthia
280:
278:
277:
273:
272:
271:
266:
265:
261:
259:
258:
254:
252:
251:
247:
245:
244:
240:
236:
235:
231:
230:
229:
228:
224:
222:
221:
217:
215:
214:
210:
208:
207:
203:
202:
201:
198:
192:
190:
186:
179:
177:
175:
171:
167:
163:
158:
154:
152:
148:
139:
137:
131:
127:
122:
114:
107:
105:
103:
98:
94:
90:
86:
77:
71:
66:
55:
49:
45:
41:
37:
32:
19:
918:
898:
894:
885:
877:
844:
840:
833:
782:
778:
764:
731:
721:
676:
672:
644:(1): 27â35.
641:
637:
631:
622:
595:
591:
581:
540:
536:
530:
485:
481:
471:
426:
422:
412:
371:
367:
341:Some of the
340:
337:
330:
323:
316:
309:
302:
295:
288:
281:
274:
269:
262:
257:Naja pallida
255:
248:
241:
232:
225:
218:
211:
204:
199:
196:
187:
183:
159:
155:
143:
134:
125:
124:Handling of
84:
82:
70:Naja pallida
69:
47:
43:
39:
35:
679:(3): E103.
250:Naja nubiae
170:cardiotoxic
954:Categories
488:(3): 609.
429:(3): 609.
349:References
206:Naja ashei
166:neurotoxic
153:swelling.
108:Background
861:0022-1511
807:0036-8075
756:234134648
748:0036-8075
695:2072-6651
557:1432-1351
504:1420-3049
482:Molecules
445:1420-3049
423:Molecules
388:0340-7594
343:Viperidae
276:Naja atra
162:cytotoxic
960:Elapidae
869:86154588
825:33479150
713:28335411
614:15339944
573:26202435
565:19462171
522:29518026
463:29518026
404:39976553
396:16007458
180:Benefits
147:chemosis
128:, using
102:toxungen
44:spitting
907:3892862
816:7610493
787:Bibcode
779:Science
732:Science
704:5371858
513:6017371
454:6017371
193:Species
151:corneal
89:species
38:fangs (
905:
867:
859:
823:
813:
805:
754:
746:
711:
701:
693:
673:Toxins
612:
571:
563:
555:
520:
510:
502:
461:
451:
443:
402:
394:
386:
42:) and
938:Video
932:Video
903:JSTOR
891:(PDF)
865:S2CID
775:(PDF)
752:S2CID
569:S2CID
400:S2CID
140:Venom
97:venom
93:cobra
48:right
886:Naja
857:ISSN
821:PMID
803:ISSN
744:ISSN
709:PMID
691:ISSN
610:PMID
561:PMID
553:ISSN
518:PMID
500:ISSN
459:PMID
441:ISSN
392:PMID
384:ISSN
168:and
149:and
40:left
849:doi
811:PMC
795:doi
783:371
736:doi
699:PMC
681:doi
646:doi
642:237
600:doi
596:207
545:doi
541:195
508:PMC
490:doi
449:PMC
431:doi
376:doi
372:191
91:of
956::
899:48
897:.
893:.
863:.
855:.
845:42
843:.
819:.
809:.
801:.
793:.
781:.
777:.
750:.
742:.
734:.
730:.
707:.
697:.
689:.
675:.
671:.
658:^
640:.
608:.
594:.
590:.
567:.
559:.
551:.
539:.
516:.
506:.
498:.
486:23
484:.
480:.
457:.
447:.
439:.
427:23
425:.
421:.
398:.
390:.
382:.
370:.
356:^
83:A
56:.
50:).
909:.
871:.
851::
827:.
797::
789::
758:.
738::
715:.
683::
677:9
652:.
648::
616:.
602::
575:.
547::
524:.
492::
465:.
433::
406:.
378::
46:(
20:)
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
