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Madrone butterfly

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382: 65: 593:, the silk nests are maintained on the edge of roof tops of houses. Like other Lepidopteran larvae, they are used in a variety of dishes such as tortilla, omelets, pies and rice. They are perceived as ‘good’ and ‘nutritious’ food due to their high protein content. In the Mixteca region of Oaxaca, excessive consumption of the larvae led to the disappearance of the species from this region. However, it has been reintroduced into this region from the state of Mexico and Durango. 258: 401:
eggs in a clump are full siblings. The number of eggs in a clump can vary from very few to as much as 350-400 eggs. The females also tend to lay these egg clumps near other conspecific eggs. It has been observed that isolated clutches tend to suffer from higher mortality than grouped clutches. It is speculated that the ovipositional behavior of the females has been under strong selection in the past to maximize social interaction among larvae.
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This secondary nest is very tough and made of multiple layers of interlaced double-stranded silk. By the end of the growing season the nest walls can be thick enough to resist tear and hold water. The larvae maintain the nests constantly, and mend any wear and tear. The entrance and exit to the bolsa is at the bottom of the nest. This protects the shelter from rain and predation and facilitates the removal of
496:, but sub-populations are highly differentiated. There is an excess of heterozygotes, and moderate levels of relatedness amongst nest-mates within the sub-populations. The high differentiation among sub-populations is thought to have been caused by weak adult dispersal, and patchiness of madrone habitats due to their restriction to higher elevations. The northern and southern populations of 458:
the trails that are commonly used are much thicker and stronger than trails that are less-frequented. When presented with alternate trail pathways, there is a strong preference to select for newer and stronger trails. The nocturnal foraging is thought to be an evolutionary response to avoid day-active parasitoids, and predators such as birds and social wasps.
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six larval instars. Larval mortality is disproportionately high in males than in females. Despite extremely low temperatures in the winter, the larvae do not undergo diapause, and continue to feed and grow throughout the year. A fully-grown larva is generally between 25 and 30 millimeters long, and pupates by April.
234:. It takes an entire year for this adult butterfly to develop from an egg. The eggs are laid in the month of June and the adults emerge the following May–June. The adults have a black and white pattern on their wings, and the males are generally much smaller and paler than the females. The larvae do not undergo 547:
The species show a general behavioral mechanism to minimize predation and parasitism. The oviposition of the eggs on the underside of the leaves, and nocturnal foraging of larvae decrease exposure to predators and other parasitoids. The final instars of the larvae also exhibit chemical defenses. When
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The later instars are nocturnal and leave their nest for foraging to remote sites an hour or two after sunset. They feed gregariously on the leaves of the host plant until the early hours of morning, and return to the nest before sunrise. The caterpillars in general spin silk whenever they walk, thus
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is dependent on the availability of their preferred host plant, the madrone. In recent times, the madrone trees have been cut and used as firewood and for making furniture and other crafts. The destruction of the madrone habitat threatens the relationship between the plant and the butterfly. However,
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larvae spend more time spinning silk and lesser amount of time foraging as compared to female larvae. Males were also observed to be more active and the first ones to lead a foraging foray. Thus, the males disproportionately bore more of the cost of silk production and exploration for new trails. It
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and leave the venous skeleton of the leaf intact. The larvae then build their first communal nest by folding these consumed leaves and securing them with silken strands. The larvae are a bright green and mildly fuzzy when they hatch, but turn brown and less pubescent as they grow. There are in total
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caterpillars are social, and they construct communal nests. The freshly hatched larvae forage and rest together, and aggregate in a loosely woven tent-like silk structure over the surface of the leaf. This is called the primary bolsa. Encompassing this structure, the larvae build a secondary bolsa.
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The eggs are bluish-white and are laid in clumps on the underside of the host plant Madrone in the month of June–July. The larvae hatch out of the eggs after approximately 3–4 weeks, sometimes after up to 60 days, in August. Upon hatching, they communally feed on the leaves and terminal branches of
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spp.). Though there is much variation in the tree quality of the madrone, the females do not show any preferential oviposition with regard to host plant quality. The eggs are laid in a clump on the undersurface of a single leaf of the madrone. The females mate only once, and consequently, all the
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Although mating, flight, and oviposition occurs in the warmest and wettest months of the year, much of the growth of the caterpillar occurs in the coldest months of the year. Even though the days are much warmer than the nights in the winter, the caterpillars remain aggregated in their nests and
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among nest-mates is 0.285, which is much lower than 0.5 that is expected amongst full-siblings. This implies that nest-mates are both kin and non-kin, which can be explained by the proximity of clutches in general. Therefore, it is proposed that communal nesting behavior evolved initially due to
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with heterogametic females. But, the primary sex-ratio i.e., the sex-ratio at conception is extremely male-biased with an average of about 70% males. This bias has been observed in both the eastern and western sub-populations, and is thought to be caused by meiotic instability. Furthermore, the
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The nests are roughly pyramidal, but can show great variation in size. The size of the nest correlates with the size of the population. The number of individuals in a nest can vary from as low as 3 individuals to 528 individuals, with an average of about 112 individuals. The survivorship of the
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The silk nests built by the larvae are believed to have been used in the past for making a paper-like fabric and small boxes. They also served as a base for paintings, and for bandaging wounds. The entire nests have also been recorded to be used as purses and as a container for liquid.
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adapted to escape the nest. The adults have black and white coloration on the wings, and the pattern is more prominent on the males. The males also are usually much paler and smaller than the females. The males have larger eyes and higher wing venation as compared to the females. The
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in the size of pupa. The male pupae are generally much smaller (18–20 mm) than female pupae (21–23 mm). Pupal mortality is much higher in females than in males. The pupal stage lasts for about a month and the adults finally emerge in May–June.
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and all instars are known to be gregarious. It takes nearly an entire year for the adult to emerge from an egg. The eggs are laid in July, and the adults eventually eclose in May–June. Consequently, there is only one generation of eggs laid each year.
449:, facilitated by a single oviposition event leading to an egg mass with high-relatedness. But the maintenance of this behavior among non-kin could be due to high benefits of communal nesting such as predator avoidance and thermodynamic efficiency. 333:
The pupae are initially light-green on pupation and later turn yellow. They pupate head down in the silk nests, and lack silken girdle unlike other Pieridae. The black and white adult wing markings are visible through the pupal case. There exists
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The adults are weak fliers, and display simple sexual behavior. Mating takes place near the communal nests right after the adults emerge from the nests. The females mate only once in their lifetime, and most males fail to find mates.
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venture out to forage strictly after sunset. Even within the nests, the caterpillars choose to cluster in the coolest regions of the nest. This type of voluntary hypothermia is expected to be an adaptation to foraging in the night.
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Underwood, Dessie L. A.; Hussein, Shafinaz; Goodpasture, Carll; Luis, Armando; Bousquets, Jorge Llorente; Shapiro, Arthur M. (2005-03-01). "Geographic Variation in Meiotic Instability in Eucheira socialis (Lepidoptera: Pieridae)".
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has also been observed that nests with male-biased ratios produced heavier male and female pupae than female-biased nests. There seems to be a sexual division of labor, which explains the observation of highly male-biased nests.
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threatened, they regurgitate a droplet of brownish-green fluid. This fluid is assumed to be distasteful to predators, and has been described as having a ‘bitter’ and ‘nutty’ flavor. The fluid also contains alkaloids, such as
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Fitzgerald, T. D.; Underwood, D. L. A. (1998-03-01). "Trail Marking by the Larva of the Madrone Butterfly Eucheira socialis and the Role of the Trail Pheromone in Communal Foraging Behavior".
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by providing a cool shelter for the larvae on sun-intensive days. The quality of the nest i.e., the thickness of the nest wall correlates with survivorship of the larvae.
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larvae is directly proportional to size of the group. Larger groups of larvae tend to forage for longer periods, and gain more weight than larvae from smaller groups.
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Underwood, Dessie L. A. (1994-08-01). "Intraspecific variability in host plant quality and ovipositionaI preferences in Eucheira socialis (Lepidoptera: Pieridae)".
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The other silk producer was the Madrone butterfly... Aztec artisans cut up the large sacs, piecing together the resulting swatches into larger pieces of "fabric."
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Fitzgerald, Terrence D.; Underwood, Dessie L. A. (2000-02-01). "Winter foraging patterns and voluntary hypothermia in the social caterpillar Eucheira socialis".
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is also male-biased. Such a ratio is thought to be evolutionarily maintained because of the selective advantage of male-biased groups in the communal nests.
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in the north. The population distribution is patchy due to the poor dispersal ability of the adults, and the patchy distribution of the host-plant, madrone.
609: 764:"natural history, sociobiology, and ethnobiology of Eucheira socialis Westwood (Lepidoptera: Pieridae), a unique and little-known butterfly from Mexico" 429:
The construction and maintenance of this nest is essential for survival of the larval species into adulthood. The nest plays an important role in
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Factors influencing oviposition behavior in the Mexican Pierid butterfly, Eucheira socialis, on its host plant, Arbutus xalapensis (Ericaceae)
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habitats. They are generally found in pine-oak and arid tropical scrub ecosystems. They are distributed from northern Sonora to Jalisco.
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Ramos-Elorduy, Julieta; Moreno, José MP; Vázquez, Adolfo I.; Landero, Ivonne; Oliva-Rivera, Héctor; Camacho, Víctor HM (2011-01-06).
1373: 1458: 1473: 1463: 1208:"Edible Lepidoptera in Mexico: Geographic distribution, ethnicity, economic and nutritional importance for rural people" 686:
Porter, Adam H.; Geiger, HansjüRg; Underwood, Dessie L. A.; Llorente-Bousquets, Jorge; Shapiro, Arthur M. (1997-03-01).
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Underwood, Dessie L. A.; Shapiro, Arthur M. (1999-09-01). "Evidence for division of labor in the social caterpillar
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the inaccessibility of these habitats due to their confinement in high elevations possibly decreases the risk.
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and go through four distinct developmental stages namely egg, larva, pupa and adult. There are six larval
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and continue to feed and grow communally in the coldest months of the year. There are two subspecies of
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extracted the silk fibres from the nests and wove them into sashes. This practise had ceased by 1997.
64: 866:"A male-biased primary sex ratio and larval mortality in Eucheira socialis (Lepidoptera: Pieridae)" 17: 625: 346:. Their escape is facilitated by soft, pliable wings. The wings also have an atypical shape and 1267: 485: 975: 865: 522: 159: 1430: 1360: 1064: 807: 489: 8: 396:
within 3 hours post-eclosion. They lay their eggs exclusively on the host plant madrone (
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spp.) as a host-plant. The species is of considerable interest to lepidopterists due to
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Fischer, Hanna M.; Wheat, Christopher W.; Heckel, David G.; Vogel, Heiko (2008-05-01).
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Adult butterflies emerge from their nests through the small exits while they are still
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is restricted to the highlands of Mexico at elevations of 1800 meters in
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is one of the many lepidopterans consumed in Mexico. In some part of
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The Unbroken Thread: Conserving the Textile Traditions of Oaxaca
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and dead larvae. The nests are not reused by later generations.
1105: 570: 317: 215: 106: 86: 1365: 1195:. Los Angeles: The Getty Conservation Institute. p. 125. 1121:
10.1603/0013-8746(2005)098[0227:GVIMII]2.0.CO;2
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Underwood*, Dessie L.A.; Shapiro, Arthur M. (1999).
1054: 891: 1186:de Avila, Alejandro (1997). Klein, Kathryn (ed.). 863: 1445: 1004: 1109:Annals of the Entomological Society of America 692:Annals of the Entomological Society of America 230:in the larval stages, and heavily male-biased 973: 325:the plant. They consume the epidermis and 42: 1241: 1223: 1212:Journal of Ethnobiology and Ethnomedicine 937: 797: 641: 1185: 761: 380: 256: 14: 1446: 436:Although all the eggs in a clutch are 1266: 1265: 1152:Latin American insects and entomology 1149: 1050: 1048: 974:Sun, J.J.; Underwood, D.L.A. (2011). 757: 755: 753: 751: 749: 747: 745: 743: 741: 580: 555: 308:Characteristic of all lepidopterans, 969: 967: 965: 963: 961: 959: 957: 933: 931: 887: 885: 883: 859: 857: 855: 853: 851: 849: 847: 845: 793: 791: 789: 787: 785: 739: 737: 735: 733: 731: 729: 727: 725: 723: 721: 681: 679: 677: 675: 673: 671: 669: 560: 479: 461: 452: 214:, it is endemic to the highlands of 1011:Behavioral Ecology and Sociobiology 533: 409: 284:are found in central Mexico, while 24: 1045: 820:10.1111/j.1365-2311.1994.tb00416.x 404: 25: 1485: 954: 938:Underwood, Dessie LeeAnn (1992). 928: 880: 842: 782: 718: 666: 1077:10.1046/j.1365-2311.2000.00236.x 218:, and exclusively relies on the 63: 1199: 1179: 1150:Hogue, Charles Leonard (1993). 1143: 1099: 998: 762:P.G., Kevan; R.A., Bye (1991). 630:Molecular Biology and Evolution 466:It has been observed that male 362: 252: 1459:Taxa named by John O. Westwood 617: 603: 376: 355:does not anneal properly post- 13: 1: 1474:Butterflies described in 1834 980:Evolutionary Ecology Research 870:Evolutionary Ecology Research 596: 528: 303: 1464:Butterflies of North America 512:The determination of sex in 507: 202:. It was first described by 7: 542: 474: 206:in 1834. Locally known as 198:that belongs to the family 10: 1490: 1009:(Lepidoptera: Pieridae)". 894:Journal of Insect Behavior 1274: 573:and other communities in 367: 165: 158: 60:Scientific classification 58: 50: 41: 34: 942:(Thesis). Davis, Calif. 228:gregarious nest-building 190:, commonly known as the 906:10.1023/A:1021000108290 359:and is non-functional. 290:Sierra Madre Occidental 262:Sierra Madre Occidental 389: 269: 248:E. socialis westwoodi. 1469:Lepidoptera of Mexico 1225:10.1186/1746-4269-7-2 1057:Ecological Entomology 1023:10.1007/s002650050614 800:Ecological Entomology 704:10.1093/aesa/90.2.230 643:10.1093/molbev/msn014 523:operational sex-ratio 384: 286:E. socialis westwoodi 260: 490:genetic architecture 282:E. socialis socialis 272:The distribution of 244:E. socialis socialis 208:Mariposa del madroño 27:Species of butterfly 1069:2000EcoEn..25...35F 812:1994EcoEn..19..245U 581:Use as food source 556:Human interactions 390: 386:Arbutus xalapensis 270: 1441: 1440: 1426:Open Tree of Life 1268:Taxon identifiers 1007:Eucheira socialis 561:Use of silk nests 504:differentiation. 484:There is limited 480:Genetic structure 462:Division of labor 453:Communal foraging 348:venation patterns 336:sexual dimorphism 268:habitat in Sonora 192:Madrone butterfly 187:Eucheira socialis 183: 182: 169:Eucheira socialis 53:Eucheira socialis 36:Eucheira socialis 16:(Redirected from 1481: 1434: 1433: 1421: 1420: 1408: 1407: 1395: 1394: 1382: 1381: 1369: 1368: 1356: 1355: 1343: 1342: 1333: 1332: 1320: 1319: 1310: 1309: 1308: 1295: 1294: 1293: 1263: 1262: 1256: 1255: 1245: 1227: 1203: 1197: 1196: 1194: 1183: 1177: 1176: 1147: 1141: 1140: 1103: 1097: 1096: 1052: 1043: 1042: 1002: 996: 995: 971: 952: 951: 935: 926: 925: 889: 878: 877: 861: 840: 839: 795: 780: 779: 759: 716: 715: 683: 664: 663: 645: 621: 615: 607: 534:Thermoregulation 431:thermoregulation 410:Communal nesting 204:John O. Westwood 171: 151:E. socialis 68: 67: 46: 32: 31: 21: 1489: 1488: 1484: 1483: 1482: 1480: 1479: 1478: 1444: 1443: 1442: 1437: 1429: 1424: 1416: 1411: 1403: 1398: 1390: 1385: 1377: 1372: 1364: 1359: 1351: 1346: 1338: 1336: 1328: 1323: 1315: 1313: 1304: 1303: 1298: 1289: 1288: 1283: 1270: 1260: 1259: 1204: 1200: 1192: 1184: 1180: 1162: 1148: 1144: 1104: 1100: 1053: 1046: 1003: 999: 972: 955: 936: 929: 890: 881: 862: 843: 796: 783: 760: 719: 684: 667: 622: 618: 608: 604: 599: 583: 563: 558: 545: 536: 531: 510: 482: 477: 464: 455: 412: 407: 405:Social behavior 379: 370: 365: 306: 255: 212:tzauhquiocuilin 179: 173: 167: 154: 62: 28: 23: 22: 15: 12: 11: 5: 1487: 1477: 1476: 1471: 1466: 1461: 1456: 1439: 1438: 1436: 1435: 1422: 1409: 1396: 1383: 1370: 1357: 1344: 1334: 1321: 1311: 1296: 1280: 1278: 1272: 1271: 1258: 1257: 1198: 1178: 1161:978-0520078499 1160: 1142: 1115:(2): 227–235. 1098: 1044: 1017:(4): 228–236. 997: 953: 927: 900:(2): 247–263. 879: 841: 806:(3): 245–256. 781: 717: 698:(2): 230–236. 665: 636:(5): 809–820. 616: 601: 600: 598: 595: 582: 579: 562: 559: 557: 554: 544: 541: 535: 532: 530: 527: 509: 506: 481: 478: 476: 473: 463: 460: 454: 451: 411: 408: 406: 403: 388:, a host plant 378: 375: 369: 366: 364: 361: 314:holometabolous 305: 302: 254: 251: 181: 180: 174: 163: 162: 156: 155: 148: 146: 142: 141: 134: 130: 129: 124: 120: 119: 114: 110: 109: 104: 100: 99: 94: 90: 89: 84: 80: 79: 74: 70: 69: 56: 55: 48: 47: 39: 38: 26: 9: 6: 4: 3: 2: 1486: 1475: 1472: 1470: 1467: 1465: 1462: 1460: 1457: 1455: 1452: 1451: 1449: 1432: 1427: 1423: 1419: 1414: 1410: 1406: 1401: 1397: 1393: 1388: 1384: 1380: 1375: 1371: 1367: 1362: 1358: 1354: 1349: 1345: 1341: 1335: 1331: 1326: 1322: 1318: 1312: 1307: 1301: 1297: 1292: 1286: 1282: 1281: 1279: 1277: 1273: 1269: 1264: 1253: 1249: 1244: 1239: 1235: 1231: 1226: 1221: 1217: 1213: 1209: 1202: 1191: 1190: 1182: 1175: 1171: 1167: 1163: 1157: 1153: 1146: 1138: 1134: 1130: 1126: 1122: 1118: 1114: 1110: 1102: 1094: 1090: 1086: 1082: 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587:E. socialis 518:chromosomal 514:E. socialis 498:E. socialis 494:E. socialis 468:E. socialis 442:relatedness 415:E. socialis 377:Oviposition 310:E. socialis 297:E. socialis 274:E. socialis 266:E. socialis 240:E. socialis 117:Lepidoptera 1448:Categories 614:, funet.fi 597:References 529:Physiology 502:karyotypic 304:Life cycle 97:Arthropoda 1337:ButMoth: 1234:1746-4269 1129:0013-8746 1085:1365-2311 1031:0340-5443 992:1522-0613 914:0892-7553 828:1365-2311 776:0013-8878 712:0013-8746 652:0737-4038 508:Sex-ratio 353:proboscis 327:mesophyll 232:sex ratio 145:Species: 83:Kingdom: 77:Eukaryota 1400:LepIndex 1314:BioLib: 1306:Eucheira 1291:Q1768392 1285:Wikidata 1276:Eucheira 1252:21211040 1170:25164105 1137:86117866 1093:84930808 1039:11984074 948:29716202 922:23710704 836:85119308 660:18296701 611:Eucheira 591:Huasteca 543:Defenses 475:Genetics 394:oviposit 357:eclosion 242:, named 236:diapause 200:Pieridae 176:Westwood 138:Eucheira 127:Pieridae 123:Family: 93:Phylum: 87:Animalia 73:Domain: 18:Eucheira 1454:Pierini 1379:1920812 1340:10671.0 1243:3034662 1065:Bibcode 808:Bibcode 550:arbutin 488:in the 420:excreta 398:Arbutus 344:teneral 318:instars 278:madrone 224:Arbutus 220:Madrone 133:Genus: 113:Order: 107:Insecta 103:Class: 51:Female 1431:205906 1418:320212 1405:177273 1392:247789 1317:531567 1250:  1240:  1232:  1168:  1158:  1135:  1127:  1091:  1083:  1037:  1029:  990:  946:  920:  912:  834:  826:  774:  710:  658:  650:  575:Oaxaca 571:Mixtec 440:, the 368:Mating 216:Mexico 178:, 1834 1366:20314 1330:80856 1218:: 2. 1193:(PDF) 1133:S2CID 1089:S2CID 1035:S2CID 986:(6). 918:S2CID 832:S2CID 194:is a 1413:NCBI 1374:GBIF 1353:4F5V 1325:BOLD 1248:PMID 1230:ISSN 1166:OCLC 1156:ISBN 1125:ISSN 1081:ISSN 1027:ISSN 988:ISSN 944:OCLC 910:ISSN 876:(6). 824:ISSN 772:ISSN 708:ISSN 656:PMID 648:ISSN 569:The 312:are 246:and 1361:EoL 1348:CoL 1238:PMC 1220:doi 1117:doi 1073:doi 1019:doi 902:doi 816:doi 700:doi 638:doi 516:is 492:of 210:or 1450:: 1428:: 1415:: 1402:: 1389:: 1376:: 1363:: 1350:: 1327:: 1302:: 1287:: 1246:. 1236:. 1228:. 1214:. 1210:. 1172:. 1164:. 1131:. 1123:. 1113:98 1111:. 1087:. 1079:. 1071:. 1061:25 1059:. 1047:^ 1033:. 1025:. 1015:46 1013:. 984:13 982:. 978:. 956:^ 930:^ 916:. 908:. 898:11 896:. 882:^ 872:. 868:. 844:^ 830:. 822:. 814:. 804:19 802:. 784:^ 770:. 766:. 720:^ 706:. 696:90 694:. 690:. 668:^ 654:. 646:. 634:25 632:. 628:. 552:. 264:, 1254:. 1222:: 1216:7 1139:. 1119:: 1095:. 1075:: 1067:: 1041:. 1021:: 994:. 950:. 924:. 904:: 874:1 838:. 818:: 810:: 778:. 714:. 702:: 662:. 640:: 222:( 20:)

Index

Eucheira

Scientific classification
Edit this classification
Eukaryota
Animalia
Arthropoda
Insecta
Lepidoptera
Pieridae
Eucheira
Binomial name
Westwood
lepidopteran
Pieridae
John O. Westwood
Mexico
Madrone
gregarious nest-building
sex ratio
diapause

Sierra Madre Occidental
madrone
Sierra Madre Occidental
holometabolous
instars
mesophyll
sexual dimorphism
teneral

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