388:
31:
464:
334:
a whole and has been in place for about 50 million years. An ancient origin of sociality in this group helps explain very sophisticated forms of social communication in some species, such as pheromonal regulation of reproduction and complex forms of kin recognition. The origin of queen and workers castes in allodapine bees is relatively recent, much less than 40 million years ago, compared with the honeybees, bumble bees and stingless bees, where true queen and worker castes evolved about 100 million years ago.
483:
54:
445:
502:
407:
426:
367:
the expanse of the Indian Ocean separating
Australia from Africa. The most likely routes involved were now-submerged island stepping stones across the Indian Ocean, or dispersal from Africa to Antarctica and then overland dispersal from Antarctica to Australia when the two continents were still connected (ref). Both of these scenarios are problematic, but have been suggested for other animal and plant species.
376:
habitat loss in
Madagascar poses a major threat to that island's unique bee fauna, including allodapine bees, many of which are still to be scientifically described; and (ii) the Australian region is likely to contain many undescribed socially parasitic species which are threatened because of their very small populations sizes. Conservation threats to allodapine bees in Asia have not been studied.
358:
Molecular research has revealed nine origins of social parasitism in allodapine bees, more than all other bees and wasp groups combined. These repeated origins of social parasitism are probably due to the allodapine trait of rearing brood in communal tunnels, a trait that might allow other species to surreptitiously lay additional eggs without them being detected.
306:
adults. The appendages, tubercles and setae serve to hold and manipulate food, and may also help larvae move around the nest. These abilities are important because larvae compete with each other to gain food, a situation which is different from all other bees, where individual larvae are isolated in cells and do not have to compete with each other.
366:
Several studies have shown that allodapine bees first evolved in Africa and then spread to
Madagascar, Asia and Australia. The earliest dispersal from Africa to Australia occurred about 30 million years ago and did not appear to involve a route via Asia, leading to a biogeographical puzzle because of
333:
Many allodapine species exhibit very simple forms of social organization, without clear queen or worker castes. For this reason it was long thought that they had only recently evolved forms of social living. However, molecular phylogenetic studies show that social living is ancestral for the tribe as
342:
Most allodapine bee species have strongly female-biased sex ratios, and in many species less than 15% of brood are male. This is very different from the vast majority of animal species where sex ratios are very close to 1:1 males:females. The preponderance of female-biased sex ratios in allodapine
309:
There are over 300 described species of allodapine bees, but many more species are undescribed. They are unique among bees in progressively rearing their larvae in undivided tunnels, so that individual larvae are not physically isolated from each other and are in constant contact with adult females,
357:
Socially parasitic allodapine bees are species that have evolved to exploit the social systems of their hosts (which are other allodapine bees) so that the parasites enter the host colonies and lay their eggs there, and both the parasite adults as well as their larvae are fed by the host species.
305:
The larvae of allodapine bees are remarkable in their complex morphology, and in most species they possess appendages, tubercles and long setae. The strange morphology of allodapine larvae is probably a result of living in open tunnels where they are in constant contact with other larvae and with
375:
Recent studies are marked by the number of species they have involved that have not been formally described (refs). This suggests that there is a large amount of allodapine diversity that is not covered by formal scientific taxonomy. Conservation concerns centre on two regions: (i) large-scale
301:
which, like most other bees, is carried on specialised hairs of the hind pair of legs, but the pollen is fed to the larvae in a progressive fashion and usually placed directly onto their bodies where they then consume it.
1011:
Bull, Nicholas J., et al. "Giving your daughters the edge: bequeathing reproductive dominance in a primitively social bee." Proceedings of the Royal
Society of London B: Biological Sciences 265.1404 (1998):
1101:
Barker, N.P.; Weston, P.H.; Rutschmann, F.R. & Sauquet, H. (2007). "Molecular dating of the 'Gondwanan' plant family
Proteaceae is only partially congruent with the timing of the break-up of Gondwana".
1066:
Schwarz, M.P.; Fuller, S.; Tierney, S.M. & Cooper, S.J.B. (2006). "Molecular phylogenetics of the exoneurine allodapine bees reveal an ancient and puzzling dispersal from Africa to
Australia".
313:
Allodapine bees vary greatly in their forms of sociality, from subsocial to highly eusocial. There are no known species that are purely solitary. They have been used widely to study
1031:
Smith, J.A.; Tierney, S.M.; Park, Y.C.; Fuller, S. & Schwarz, M.P. (2007). "Origins of social parasitism: The importance of divergence ages in phylogenetic studies".
349:, females provide the useful work in the colony and group living increases colony success, so the sex ratio is almost always female biased in this species.
343:
bees is thought to be due to the benefits of sisters cooperating with each other and involves a theory known as local resource enhancement. For example, in
1365:
620:
Schwarz, M.P.; Richards, M.H.; Danforth, B.N. (2007). "Changing paradigms in insect social evolution: new insights from halictine and allodapine bees".
222:
180:
1314:
471:
452:
210:
204:
192:
186:
509:
490:
234:
216:
174:
240:
228:
293:
Many of the species in the tribe form small social colonies where a group of females cooperatively care for the developing larvae. The
1179:
Smith, J.A.; Schwarz, M.P. (2006). "New species and unexpected diversity of socially parasitic bees in the genus
Inquilina Michener".
1021:
Langer, Philipp, et al. "Reproductive skew in the
Australian allodapine bee Exoneura robusta." Animal behaviour 71.1 (2006): 193–201.
667:"Social complexity and large colony sizes are not sufficient to explain lack of reversions to solitary living over long time-scales"
1301:
854:
O'Keefe, K.J.; Schwarz. M.P. (1990). "Pheromones are implicated in reproductive differentiation in a primitively social bee".
1306:
586:
Michener, C.D.; Syed, I.H. (1962), "Specific characters of the larvae and adults of
Allodapula in the Australian region",
1319:
387:
140:
463:
1262:
482:
444:
53:
1138:
314:
501:
899:"Giving your daughters the edge: bequeathing reproductive dominance in a primitively social bee"
1215:
804:
Chenoweth, L.; Schwarz, M.P. (2011). "Historical biogeography of
Australian allodapine bees".
1332:
406:
1355:
1253:
951:
863:
770:"Cooperative nesting and complex female biased sex allocation in a tropical allodapine bee"
633:
8:
1360:
425:
955:
867:
716:
Schwarz, M.P.; Tierney, S.M.; Rehan, S.M.; Chenoweth, L.B. & Cooper, S.B.J. (2007).
665:
Chenoweth, L.B.; Tierney, S.M.; Smith, J.A.; Cooper, S.B.J. & Schwarz, M.P. (2007).
30:
1196:
1161:
1119:
974:
939:
923:
898:
879:
821:
742:
717:
693:
666:
603:
599:
48:
1139:"Bee Conservation in Sub-Saharan Africa and Madagascar: Diversity, Status and Threats"
1327:
1275:
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1115:
1083:
1048:
979:
817:
786:
769:
747:
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664:
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1200:
1165:
1123:
883:
825:
715:
607:
1280:
1188:
1153:
1111:
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1075:
1040:
969:
959:
918:
910:
871:
813:
781:
737:
729:
688:
678:
637:
629:
595:
345:
1267:
1065:
964:
897:
Bull, N.J.; Mibus, A.C.; Norimatsu, Y.; Jarmyn, B.L. & Schwarz, M.P. (1998).
267:
1044:
1238:
318:
1079:
896:
274:. There is also a rare genus, Exoneuridia, that occurs in isolated regions of
1349:
1157:
1030:
683:
1087:
1052:
983:
914:
751:
733:
702:
651:
1288:
1247:
619:
271:
255:
125:
105:
875:
414:
395:
322:
168:
162:
39:
1293:
642:
85:
65:
1209:
1232:
1136:
940:"The antiquity and evolutionary history of social behavior in bees"
552:
Terzo, M. (1999). "Revision du genre Exoneuridia Cockerell, 1911".
433:
198:
156:
310:
who provide them with food, groom them, and remove their faeces.
283:
718:"The evolution of eusociality in bees: Workers began by waiting"
298:
275:
263:
259:
115:
95:
75:
853:
294:
287:
279:
937:
803:
767:
1178:
585:
541:, Baltimore & London: Johns Hopkins University Press
370:
1137:
Eardley, C.; Gikungu, M. & Schwarz, M.P. (2009).
532:
530:
528:
1347:
799:
797:
763:
761:
525:
794:
758:
575:, Harvard University Press, pp. 307–309
1366:Taxa named by Theodore Dru Alison Cockerell
938:Cardinal, S. & Danforth, B.N. (2011).
361:
29:
973:
963:
922:
785:
774:Biological Journal of the Linnean Society
741:
692:
682:
641:
551:
1001:, New Jersey: Princeton University Press
838:
570:
536:
1348:
1214:
1213:
1033:Molecular Phylogenetics and Evolution
843:, Cambridge: Harvard University Press
634:10.1146/annurev.ento.51.110104.150950
996:
768:Thompson, S.; Schwarz, M.P. (2006).
371:Conservation issues and biodiversity
352:
262:. They occur throughout sub-Saharan
254:is a tribe of bees in the subfamily
328:
13:
600:10.1111/j.1440-6055.1962.tb00168.x
14:
1377:
337:
1193:10.1111/j.1744-7917.2009.01266.x
1116:10.1111/j.1365-2699.2007.01749.x
903:Proceedings of the Royal Society
818:10.1111/j.1365-2699.2011.02488.x
787:10.1111/j.1095-8312.2006.00679.x
588:Australian Journal of Entomology
500:
481:
462:
443:
424:
405:
386:
52:
1172:
1130:
1094:
1059:
1024:
1015:
1005:
990:
931:
890:
847:
841:The Social Behavior of the Bees
832:
573:The Social Behavior of the Bees
325:, and historical biogeography.
709:
658:
613:
579:
564:
545:
1:
554:Belgian Journal of Entomology
519:
965:10.1371/journal.pone.0021086
7:
1045:10.1016/j.ympev.2006.12.028
622:Annual Review of Entomology
10:
1382:
379:
1222:
1080:10.1080/10635150500431148
153:
148:
49:Scientific classification
47:
37:
28:
23:
671:BMC Evolutionary Biology
1104:Journal of Biogeography
839:Michener, C.D. (1984),
806:Journal of Biogeography
684:10.1186/1471-2148-7-246
571:Michener, C.D. (1974),
537:Michener, C.D. (2007),
362:Historical biogeography
915:10.1098/rspb.1998.0450
734:10.1098/rsbl.2010.0757
1158:10.1051/apido/2009016
43:sp. in South Africa
997:West, S.A. (2009),
956:2011PLoSO...621086C
868:1990NW.....77...83O
856:Naturwissenschaften
1068:Systematic Biology
876:10.1007/bf01131780
1343:
1342:
1328:Open Tree of Life
1216:Taxon identifiers
1110:(12): 2012–2027.
909:(1404): 221–225.
539:Bees of the World
353:Social parasitism
323:social parasitism
248:
247:
144:
1373:
1336:
1335:
1323:
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1143:
1134:
1128:
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1063:
1057:
1056:
1039:(3): 1131–1137.
1028:
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1002:
994:
988:
987:
977:
967:
935:
929:
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894:
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851:
845:
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836:
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829:
812:(8): 1471–1483.
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713:
707:
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346:Exoneura robusta
329:Social evolution
315:social evolution
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21:
20:
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722:Biology Letters
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268:South East Asia
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51:
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1298:
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1272:
1259:
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1228:
1226:
1220:
1219:
1207:
1206:
1187:(4): 343–350.
1181:Insect Science
1171:
1152:(3): 355–366.
1129:
1093:
1058:
1023:
1014:
1004:
999:Sex Allocation
989:
930:
889:
846:
831:
793:
780:(2): 355–364.
757:
728:(2): 277–280.
708:
657:
612:
578:
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338:Sex allocation
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319:sex allocation
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1027:
1018:
1008:
1000:
993:
985:
981:
976:
971:
966:
961:
957:
953:
950:(6): e21086.
949:
945:
941:
934:
925:
920:
916:
912:
908:
904:
900:
893:
885:
881:
877:
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827:
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764:
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427:
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269:
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261:
257:
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244:
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236:
231:
230:
225:
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223:Hasinamelissa
219:
218:
213:
212:
207:
206:
201:
200:
195:
194:
189:
188:
183:
182:
181:Compsomelissa
177:
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137:
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67:
64:
61:
60:
55:
50:
46:
42:
41:
36:
32:
27:
22:
19:
16:Tribe of bees
1223:
1184:
1180:
1174:
1149:
1145:
1132:
1107:
1103:
1096:
1074:(1): 31–45.
1071:
1067:
1061:
1036:
1032:
1026:
1017:
1007:
998:
992:
947:
943:
933:
906:
902:
892:
862:(2): 83–86.
859:
855:
849:
840:
834:
809:
805:
777:
773:
725:
721:
711:
674:
670:
660:
625:
621:
615:
594:(1): 30–41,
591:
587:
581:
572:
566:
557:
553:
547:
538:
508:
489:
470:
451:
432:
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344:
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209:
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197:
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185:
179:
173:
167:
161:
155:
154:
135:
38:
18:
1356:Xylocopinae
1289:iNaturalist
1248:Wikispecies
628:: 127–150.
494:sp., female
472:Exoneuridia
456:sp., female
453:Exoneurella
297:are fed on
272:Australasia
256:Xylocopinae
211:Exoneuridia
205:Exoneurella
193:Eucondylops
187:Effractapis
126:Xylocopinae
122:Subfamily:
106:Hymenoptera
24:Allodapini
1361:Bee tribes
1350:Categories
1268:Allodapini
1254:Allodapini
1224:Allodapini
1146:Apidologie
1012:1411–1415.
560:: 137–152.
520:References
510:Macrogalea
491:Halterapis
415:Braunsapis
396:Allodapula
252:Allodapini
235:Macrogalea
217:Halterapis
175:Brevineura
169:Braunsapis
163:Allodapula
136:Allodapini
86:Arthropoda
40:Allodapula
643:2328/9446
513:sp., male
258:, family
241:Nasutapis
229:Inquilina
141:Cockerell
72:Kingdom:
66:Eukaryota
1239:Q4733037
1233:Wikidata
1201:85586108
1166:23555870
1124:86156197
1088:16507522
1053:17433725
984:21695157
944:PLOS ONE
884:43776819
826:83205237
752:20943679
703:18154646
652:16866635
608:83715378
434:Exoneura
199:Exoneura
157:Allodape
112:Family:
82:Phylum:
76:Animalia
62:Domain:
975:3113908
952:Bibcode
924:1689225
864:Bibcode
743:3061166
694:2231370
677:: 246.
380:Gallery
284:Lebanon
149:Genera
132:Tribe:
102:Order:
96:Insecta
92:Class:
1333:707246
1307:633980
1294:487220
1199:
1164:
1122:
1086:
1051:
982:
972:
921:
882:
824:
750:
740:
701:
691:
650:
606:
299:pollen
295:larvae
276:Turkey
270:, and
264:Africa
260:Apidae
143:, 1902
116:Apidae
1320:78174
1281:628CD
1197:S2CID
1162:S2CID
1142:(PDF)
1120:S2CID
880:S2CID
822:S2CID
604:S2CID
1315:NCBI
1302:ITIS
1084:PMID
1049:PMID
980:PMID
748:PMID
699:PMID
648:PMID
288:Iran
286:and
280:Iraq
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