186:
299:
198:
213:
641:
254:
20:
658:
619:
228:
676:
270:
231:
234:
233:
230:
680:
596:
has also been observed agitating the eyes of caimans and turtles in order to force tear production, which the male butterflies of the species can drink for minerals. The minerals, which can also be obtained from more typical mud-puddling behavior, are used for the butterfly's
235:
679:
684:
682:
678:
683:
274:
273:
278:
276:
272:
232:
277:
681:
275:
831:
802:
159:
Males seem to benefit from the sodium uptake through mud-puddling behaviour with an increase in reproductive success. The collected sodium and amino acids are often transferred to the female with the
179:, this is released in forced anal jets at 3 second intervals. Fluid of up to 600 times the body mass may pass through and males have a much longer ileum (anterior hindgut) than non-puddling females.
229:
677:
185:
763:
730:
271:
828:
455:
independently in several lineages. Specialist carrion-feeders may even have the ability to smell out and home in on rotting meat over hundreds of meters. In the
Bornean
197:
799:
539:
has been noted to visit and suck tears by inserting their proboscis into the closed eyelids of roosting birds. Similar behaviour has been reported in
113:
are attracted to various kinds of sweat and tears, including that of humans, and other bee species have been recorded as doing so to various degrees.
547:
212:
477:) carrion-feeders tend to have a markedly larger bulk and smaller wings, making them more dashing, maneuverable flyers than fruit specialists like
1019:
541:
170:
When puddling, many butterflies and moths pump fluid through the digestive tract and release fluid from their anus. In some, such as the male
640:
657:
953:
Sculley, Colleen E., and Carol L. Boggs. "Mating systems and sexual division of foraging effort affect puddling behaviour by butterflies."
1348:
Bänziger, Hans; Boongird, Somnuk; Sukumalanand, Prachaval; Bänziger, Sängdao (2009). "Bees (Hymenoptera: Apidae) That Drink Human Tears".
1265:
de Lima Moraes, Leandro JoĂŁo
Carneiro (2018-09-17). "Please, more tears: a case of a moth feeding on antbird tears in central Amazonia".
618:
55:, and sucks up the fluid. Where the conditions are suitable, conspicuous insects such as butterflies commonly form aggregations on wet
501:
are rarely if ever seen on carrion, dung and rotting fruit, though they may be avid mud-puddlers in the strict sense. Altogether, the
1332:
1150:
760:
727:
91:) are diverse in their strategies to gather liquid nutrients. Typically, mud-puddling behaviour takes place on wet soil. But even
298:
253:
1321:(1992): Remarkable new cases of moths drinking human tears in Thailand (Lepidoptera: Thyatiridae, Sphingidae, Notodontidae).
938:
911:
126:
males puddle with a much higher frequency than females. The presence of an assembly of butterflies on the ground acts on
1210:
109:. Again, similar behaviour is not limited to the Lepidoptera; for example, the various species of bees commonly called
1041:(2004): The effect of male sodium diet and mating history on female reproduction in the puddling squinting bush brown
1514:
343:
spp.) or carrion seem to prefer ammonium ions rather than sodium. In rotting, the tissues of fruits release
551:
in Brazil. Other cases of moths drinking human tears have been reported from
Thailand. Some species of the
531:
1071:
Scott R. Smedley in Resh, V. H. & R. T. Cardé (Editors) 2003. Encyclopedia of
Insects. Academic Press.
509:
have numerous mud-puddlers that also frequently visit dung but avoid fruits and carrion (namely the genus
422:
849:(1934): On the sexes of some South American moths attracted to light, human perspiration and damp sand.
719:(1996): Mating systems and sexual division of foraging effort affect puddling behaviour by butterflies.
355:
1087:(Oedipodidae: Orthoptera): Does it detect and prefer salts or nitrogenous compounds from human urine?
1399:
de la Rosa, Carlos L (2014-05-01). "Additional observations of lachryphagous butterflies and bees".
1499:
395:
358:, numerous species of butterflies regularly visit decaying fruit to drink. This behavior is mainly
1158:
578:
875:(2006): Diversity and ecology of carrion- and fruit-feeding butterflies in Bornean rain forest.
1190:
Hilgartner, R.; Raoilison, Mamisolo; BĂĽttiker, Willhelm; Lees, David C. & Krenn, Harald W.
969:(2001): Habitat and altitude preferences of butterflies in Aralam Wildlife Sanctuary, Kerala.
928:
901:
752:(1991) Mud puddling by butterflies is not a simple matter Ecological Entomology 16(1):123-127
1519:
1081:
Shen, Ke; Wang, Hao-Jie; Shao, Lin; Xiao, Kai; Shu, Jin-Ping; Xu, Tian-Sen & Li, Guo-Qing
647:
1148:(2001): Species diversity and community structure in neotropical fruit-feeding butterflies.
1504:
1447:
1408:
148:
season. The groups generally include several species, particularly members of the families
19:
1329:
781:(1999): Mud-puddling behavior in tropical butterflies: In search of proteins or minerals?
351:
that result from the metabolic processes of decay organisms, used as fuel by butterflies.
8:
348:
1451:
1412:
1465:
1381:
1373:
1298:
993:
757:
485:
461:
977:
386:
is usually more intentionally utilized. Carrion-feeders seem to represent a different
1509:
1365:
1290:
1282:
1176:
934:
907:
665:
505:
show the highest variety of nutrient-gathering strategies among the butterflies; the
260:
175:
122:
1385:
1302:
359:
1473:
1455:
1416:
1357:
1274:
1245:
1235:
1224:"Save your tears: eye secretions of a Ringed Kingfisher fed upon by an erebid moth"
1202:
1129:
1099:
1095:
1055:
1043:
883:
791:
753:
691:
318:
60:
24:
1460:
1435:
1336:
1194:
873:
Hamer, K.C.; Hill, J.K.; Benedick, S.; Mustaffa, N.; Chey, V.K. & Maryati, M.
835:
806:
767:
734:
556:
437:
376:
243:
128:
43:, but also occurring in other animals, primarily insects. The organism seeks out
820:(1982): Nocturnal occurrences of leafhoppers (Homoptera: Cicadellidae) at soil.
71:, and ecology. This behaviour also has been seen in some other insects like the
1027:
479:
218:
77:
1059:
887:
1493:
1469:
1434:
van den Burg, Matthijs P.; de Rueda, Javier Aznar González (1 January 2021).
1369:
1286:
1250:
598:
506:
387:
371:
160:
1436:"Lachryphagy by cockroaches: reptile tears to increase reproductive output?"
1294:
1206:
626:
590:
visit the eyes of mammals and have been known to cause distress to humans.
494:
285:
171:
164:
149:
1361:
1133:
795:
592:
502:
410:
406:
334:
304:
132:, for example, as a stimulus to join the presumptive mud-puddling flock.
84:
64:
1478:
1377:
1119:) detect and prefer nectar amino acids and other nitrogenous compounds?
991:(1987): Puddling in butterflies: sodium affects reproductive success in
612:. This behaviour might thus be far more common than previously thought.
1240:
1223:
561:
536:
456:
418:
400:
313:
204:
110:
72:
1420:
390:
from "classical" mud-puddlers and fruit-feeders. They include diverse
1278:
1121:
783:
609:
523:
511:
498:
452:
445:
433:
363:
44:
40:
1347:
529:, are somewhat notorious for their blood- and tear-drinking habits.
526:
490:
467:
339:
327:
153:
68:
120:, puddling behaviour is more commonly seen in males. For example,
95:
on human skin may be attractive to butterflies such as species of
1171:
473:
427:
383:
203:
Collective of different butterfly species mud-puddling on a damp
145:
117:
52:
572:(tear drinking) occur in a number of unrelated moths that visit
573:
323:
167:. This nutrition also enhances the survival rate of the eggs.
552:
344:
141:
106:
102:
97:
92:
88:
560:
are called "vampire moths" as they suck blood from sleeping
322:) – are attracted to human urine, specifically to the
1192:(2007): Malagasy birds as hosts for eye-frequenting moths.
605:
519:
391:
56:
515:), and some which are attracted to any pungent substance.
47:
in certain moist substances such as rotting plant matter,
1083:(2009): Mud-puddling in the yellow-spined bamboo locust,
362:, though some are highly attracted to old fruit, notably
330:
48:
1427:
1017:(1996): Sodium: a male nuptial gift to its offspring.
899:
1433:
582:is a notable example. Stingless bees in the genera
293:
906:. University of California Press. pp. 457–.
1491:
1179:for being easily attracted with stinking cheese.
1039:Molleman, F.; Zwaan, B.J. & Brakefield, P.M.
59:, dung, or carrion. From the fluids they obtain
1264:
316:– e.g. the yellow-spined bamboo locust (
308:butterfly drinking from the tears of a turtle
67:that play various roles in their physiology,
1323:Natural History Bulletin of the Siam Society
1314:
1312:
1221:
926:
1350:Journal of the Kansas Entomological Society
868:
866:
864:
862:
822:Journal of the Kansas Entomological Society
144:this phenomenon is mostly seen in the post-
28:mud-puddling at the edge of a forest stream
1398:
779:Beck, J.; MĂĽhlenberg, E. & Fiedler, K.
1477:
1459:
1309:
1249:
1239:
920:
712:
710:
608:, but has recently also been observed in
483:and opportunistic fruit visitors such as
221:excreting excess water after mud-puddling
1401:Frontiers in Ecology and the Environment
893:
859:
297:
18:
772:
191:Aggregation of butterflies mud puddling
1492:
967:Sreekumar, P.G. & Balakrishnan, M.
707:
903:Latin American Insects and Entomology
39:, is a behaviour most conspicuous in
347:and other organic compounds such as
1049:Behavioral Ecology and Sociobiology
13:
1113:Erhardt, A. & Rusterholz, H.P.
758:10.1111/j.1365-2311.1991.tb00199.x
14:
1531:
1228:Revista Brasileira de Ornitologia
337:that are attracted to dung (e.g.
1115:(1998): Do Peacock butterflies (
674:
656:
639:
617:
604:Tear-drinking is not limited to
294:Other sources of liquid nutrient
268:
252:
226:
211:
196:
184:
1392:
1341:
1258:
1215:
1182:
1163:
1146:DeVries, P.J. & Walla, T.R.
1139:
1105:
1074:
1065:
1032:
1007:
982:
960:
856:: 769-791.-- Volume 67 pp.81-87
717:Sculley, C.E. & Boggs, C.L.
465:) or opportunistic (some other
101:. More unusual sources include
1100:10.1016/j.jinsphys.2008.10.011
947:
900:Charles Leonard Hogue (1993).
840:
811:
739:
489:. Other butterflies like most
75:, e.g. the potato leafhopper,
1:
1461:10.1080/23766808.2021.1953892
989:Pivnik, K. & McNeil, J.N.
700:
1089:Journal of Insect Physiology
1015:Medley S.R. & Eisner, T.
669:) sipping from a guava fruit
624:Blue lesser purple emperor (
601:during sexual reproduction.
532:Hemiceratoides hieroglyphica
7:
877:Journal of Tropical Ecology
423:gossamer-winged butterflies
10:
1536:
651:feeding on a bird dropping
356:Borneo lowland rain forest
288:) butterflies mud-puddling
135:
1175:species are famous among
1060:10.1007/s00265-004-0789-2
888:10.1017/S0266467405002750
695:drinking from metal fence
1440:Neotropical Biodiversity
999:Physiological Entomology
396:brush-footed butterflies
16:Insect feeding behaviour
933:. Berg. pp. 280–.
579:Mecistoptera griseifusa
1515:Ecological restoration
1207:10.1098/rsbl.2006.0581
930:Insects and Human Life
309:
29:
1362:10.2317/JKES0811.17.1
1222:Sazima, Ivan (2015).
1134:10.1007/s004420050690
957:21.2 (1996): 193-197.
955:Ecological Entomology
927:Brian Morris (2004).
796:10.1007/s004420050770
721:Ecological Entomology
648:Cyclosia papilionaris
568:(eye-attraction) and
301:
22:
564:, including humans.
451:Carrion-feeding has
1452:2021NeBio...7..276V
1413:2014FrEE...12..210D
1151:Biol. J. Linn. Soc.
163:during mating as a
1335:2011-07-21 at the
1241:10.1007/BF03544314
1177:lepidopterologists
994:Thymelicus lineola
834:2011-05-20 at the
805:2011-07-07 at the
766:2012-10-21 at the
733:2012-10-18 at the
486:Charaxes durnfordi
462:Charaxes bernardus
415:Charaxes bernardus
310:
30:
1421:10.1890/14.wb.006
1251:20.500.12733/1251
940:978-1-84520-075-6
913:978-0-520-07849-9
685:
666:Euthalia aconthea
279:
261:Monarch butterfly
236:
176:Gluphisia crenata
123:Speyeria mormonia
87:(butterflies and
1527:
1484:
1483:
1481:
1463:
1431:
1425:
1424:
1396:
1390:
1389:
1345:
1339:
1320:
1316:
1307:
1306:
1279:10.1002/ecy.2518
1262:
1256:
1255:
1253:
1243:
1219:
1213:
1209:(HTML abstract)
1191:
1186:
1180:
1167:
1161:
1147:
1143:
1137:
1114:
1109:
1103:
1085:Ceracris kiangsu
1082:
1078:
1072:
1069:
1063:
1044:Bicyclus anynana
1040:
1036:
1030:
1016:
1011:
1005:
990:
986:
980:
971:Tropical Ecology
968:
964:
958:
951:
945:
944:
924:
918:
917:
897:
891:
874:
870:
857:
848:
847:Collenette, C.L.
844:
838:
819:
815:
809:
798:(HTML abstract)
780:
776:
770:
751:
747:
743:
737:
718:
714:
692:Catagramma pygas
687:
686:
660:
643:
621:
545:in Colombia and
522:, mainly of the
319:Ceracris kiangsu
284:Cabbage white (
281:
280:
256:
238:
237:
215:
200:
188:
25:Parthenos sylvia
1535:
1534:
1530:
1529:
1528:
1526:
1525:
1524:
1500:Insect behavior
1490:
1489:
1488:
1487:
1432:
1428:
1397:
1393:
1346:
1342:
1337:Wayback Machine
1318:
1317:
1310:
1263:
1259:
1220:
1216:
1189:
1187:
1183:
1168:
1164:
1145:
1144:
1140:
1136:(HTML abstract)
1112:
1110:
1106:
1102:(HTML abstract)
1080:
1079:
1075:
1070:
1066:
1062:(HTML abstract)
1047:(Lepidoptera).
1038:
1037:
1033:
1014:
1012:
1008:
988:
987:
983:
966:
965:
961:
952:
948:
941:
925:
921:
914:
898:
894:
890:(HTML abstract)
872:
871:
860:
846:
845:
841:
836:Wayback Machine
817:
816:
812:
807:Wayback Machine
778:
777:
773:
768:Wayback Machine
749:
745:
744:
740:
735:Wayback Machine
716:
715:
708:
703:
696:
688:
675:
670:
661:
652:
644:
635:
622:
548:Gorgone macarea
438:common imperial
377:Bassarona dunya
296:
289:
282:
269:
264:
257:
248:
244:Papilio glaucus
239:
227:
222:
216:
207:
201:
192:
189:
138:
129:Battus philenor
17:
12:
11:
5:
1533:
1523:
1522:
1517:
1512:
1507:
1502:
1486:
1485:
1446:(1): 276–278.
1426:
1391:
1356:(2): 135–150.
1340:
1308:
1257:
1234:(4): 392–394.
1214:
1201:(2): 117–120.
1181:
1162:
1138:
1128:(4): 536-542.
1104:
1073:
1064:
1054:(4): 404–411.
1031:
1026:(2): 809–813.
1006:
981:
976:(2): 277-281.
959:
946:
939:
919:
912:
892:
858:
839:
810:
790:(1): 140–148.
771:
738:
726:(2): 193-197.
705:
704:
702:
699:
698:
697:
689:
673:
671:
663:Common baron (
662:
655:
653:
645:
638:
636:
623:
616:
599:spermatophores
566:Ophthalmotropy
480:Prothoe franck
459:, specialist (
442:Cheritra freja
295:
292:
291:
290:
283:
267:
265:
258:
251:
249:
240:
225:
223:
219:Spot swordtail
217:
210:
208:
202:
195:
193:
190:
183:
137:
134:
78:Empoasca fabae
15:
9:
6:
4:
3:
2:
1532:
1521:
1518:
1516:
1513:
1511:
1508:
1506:
1503:
1501:
1498:
1497:
1495:
1480:
1475:
1471:
1467:
1462:
1457:
1453:
1449:
1445:
1441:
1437:
1430:
1422:
1418:
1414:
1410:
1406:
1402:
1395:
1387:
1383:
1379:
1375:
1371:
1367:
1363:
1359:
1355:
1351:
1344:
1338:
1334:
1331:
1327:
1324:
1315:
1313:
1304:
1300:
1296:
1292:
1288:
1284:
1280:
1276:
1273:(2): e02518.
1272:
1268:
1261:
1252:
1247:
1242:
1237:
1233:
1229:
1225:
1218:
1212:
1208:
1204:
1200:
1197:
1196:
1185:
1178:
1174:
1173:
1166:
1160:
1156:
1153:
1152:
1142:
1135:
1131:
1127:
1124:
1123:
1118:
1108:
1101:
1097:
1093:
1090:
1086:
1077:
1068:
1061:
1057:
1053:
1050:
1046:
1045:
1035:
1029:
1025:
1022:
1021:
1010:
1004:(4): 461–472.
1003:
1000:
996:
995:
985:
979:
975:
972:
963:
956:
950:
942:
936:
932:
931:
923:
915:
909:
905:
904:
896:
889:
885:
881:
878:
869:
867:
865:
863:
855:
852:
843:
837:
833:
830:
829:HTML abstract
826:
823:
814:
808:
804:
801:
797:
793:
789:
786:
785:
775:
769:
765:
762:
759:
755:
742:
736:
732:
729:
725:
722:
713:
711:
706:
694:
693:
672:
668:
667:
659:
654:
650:
649:
642:
637:
633:
629:
628:
620:
615:
614:
613:
611:
607:
602:
600:
595:
594:
589:
585:
581:
580:
575:
571:
567:
563:
559:
558:
554:
550:
549:
544:
543:
542:Azeta melanea
538:
534:
533:
528:
525:
521:
516:
514:
513:
508:
507:Limenitidinae
504:
500:
496:
492:
488:
487:
482:
481:
476:
475:
470:
469:
464:
463:
458:
454:
449:
447:
443:
439:
435:
431:
429:
424:
421:, as well as
420:
416:
412:
408:
404:
402:
397:
393:
389:
388:feeding guild
385:
381:
379:
378:
373:
372:Limenitidinae
369:
368:Neorina lowii
365:
361:
360:opportunistic
357:
352:
350:
346:
342:
341:
336:
333:in it. Those
332:
329:
325:
321:
320:
315:
307:
306:
300:
287:
266:
262:
255:
250:
246:
245:
224:
220:
214:
209:
206:
199:
194:
187:
182:
181:
180:
178:
177:
173:
168:
166:
162:
161:spermatophore
157:
155:
151:
147:
143:
133:
131:
130:
125:
124:
119:
114:
112:
108:
104:
100:
99:
94:
90:
86:
82:
80:
79:
74:
70:
66:
62:
58:
54:
50:
46:
42:
38:
34:
27:
26:
21:
1520:Hydrogeology
1479:10261/249057
1443:
1439:
1429:
1404:
1400:
1394:
1353:
1349:
1343:
1330:PDF fulltext
1325:
1322:
1319:Bänziger, H.
1270:
1266:
1260:
1231:
1227:
1217:
1211:PDF fulltext
1198:
1193:
1184:
1170:
1165:
1159:PDF fulltext
1154:
1149:
1141:
1125:
1120:
1116:
1107:
1094:(1): 78-84.
1091:
1088:
1084:
1076:
1067:
1051:
1048:
1042:
1034:
1028:PDF fulltext
1023:
1018:
1009:
1001:
998:
992:
984:
978:PDF fulltext
973:
970:
962:
954:
949:
929:
922:
902:
895:
882:(1): 25–33.
879:
876:
853:
851:Entomologist
850:
842:
827:(1): 73–74.
824:
821:
813:
800:PDF fulltext
787:
782:
774:
761:PDF fulltext
741:
728:PDF fulltext
723:
720:
690:
664:
646:
631:
627:Apatura ilia
625:
603:
591:
588:Pariotrigona
587:
583:
577:
569:
565:
555:
546:
540:
530:
517:
510:
495:Papilionidae
484:
478:
472:
466:
460:
450:
441:
426:
414:
399:
382:
375:
367:
353:
338:
317:
311:
303:
286:Pieris rapae
263:mud-puddling
247:mud-puddling
242:
174:
169:
165:nuptial gift
158:
150:Papilionidae
140:In tropical
139:
127:
121:
115:
96:
83:
76:
36:
35:, or simply
33:Mud-puddling
32:
31:
23:
1505:Butterflies
1328:: 101–102.
1195:Biol. Lett.
818:Adler, P.H.
610:cockroaches
593:Dryas iulia
584:Lisotrigona
570:lachryphagy
562:vertebrates
503:Nymphalidae
411:tawny rajah
407:Nymphalinae
335:Lepidoptera
305:Dryas iulia
85:Lepidoptera
73:leafhoppers
65:amino acids
41:butterflies
1494:Categories
1407:(4): 210.
1117:Inachis io
750:LA Jackson
701:References
537:Madagascar
457:Charaxinae
419:Charaxinae
401:Cirrochroa
314:Orthoptera
205:stream bed
172:notodontid
111:sweat bees
1470:2376-6808
1370:0022-8567
1287:0012-9658
1122:Oecologia
784:Oecologia
746:Boggs, CL
634:) on dung
524:subfamily
512:Limenitis
499:Morphinae
446:Theclinae
444:) of the
434:Curetinae
417:) of the
364:Satyrinae
45:nutrients
1510:Habitats
1386:84853663
1378:20621944
1333:Archived
1303:52290796
1295:30222853
1169:Several
1157:: 1–15.
832:Archived
803:Archived
764:Archived
731:Archived
557:Calyptra
527:Calpinae
518:Certain
491:Pieridae
468:Charaxes
430:tagalica
398:such as
374:such as
349:alcohols
340:Zeuxidia
328:ammonium
154:Pieridae
116:In many
69:ethology
37:puddling
1448:Bibcode
1409:Bibcode
1267:Ecology
1172:Apatura
574:mammals
474:Polyura
453:evolved
436:or the
432:of the
428:Curetis
409:or the
405:of the
394:, e.g.
384:Carrion
146:monsoon
136:On soil
118:species
53:carrion
1468:
1384:
1376:
1368:
1301:
1293:
1285:
937:
910:
748:&
403:emalea
370:) and
366:(e.g.
345:sugars
324:sodium
51:, and
1466:eISSN
1382:S2CID
1374:JSTOR
1299:S2CID
606:moths
553:genus
520:moths
425:like
312:Some
259:Male
241:Male
142:India
107:tears
103:blood
98:Halpe
93:sweat
89:moths
61:salts
1366:ISSN
1291:PMID
1283:ISSN
1020:PNAS
935:ISBN
908:ISBN
632:ilia
586:and
497:and
471:and
392:taxa
331:ions
326:and
152:and
105:and
63:and
57:soil
1474:hdl
1456:doi
1417:doi
1358:doi
1275:doi
1271:100
1246:hdl
1236:doi
1203:doi
1130:doi
1126:117
1096:doi
1056:doi
884:doi
854:102
792:doi
788:119
754:doi
630:f.
535:of
354:In
49:mud
1496::
1472:.
1464:.
1454:.
1442:.
1438:.
1415:.
1405:12
1403:.
1380:.
1372:.
1364:.
1354:82
1352:.
1326:40
1311:^
1297:.
1289:.
1281:.
1269:.
1244:.
1232:23
1230:.
1226:.
1188:*
1155:74
1111:*
1092:55
1052:56
1024:93
1013:*
1002:12
997:.
974:42
880:22
861:^
825:55
724:21
709:^
576:.
493:,
448:.
380:.
302:A
156:.
81:.
1482:.
1476::
1458::
1450::
1444:7
1423:.
1419::
1411::
1388:.
1360::
1305:.
1277::
1254:.
1248::
1238::
1205::
1199:3
1132::
1098::
1058::
943:.
916:.
886::
794::
756::
440:(
413:(
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
