542:
564:
225:
workers and minor workers) and soldiers (major soldiers and minor soldiers). The royal pair produces workers, soldiers and alates (future royal pairs). The royal pair lives in the "royal chamber" where the queen continuously lays eggs when the king mates with her. Young ones are immature workers, soldier and alates. They live in the royal chamber after they hatch. Workers concentrate on colony process, for example collecting dead plant material, making the fungal comb, brood care, taking care of young ones etc. Soldiers protect the colony.
384:) that are used to asexually propagate the fungus in the colony. The fungus genotype that exhibits the highest production of mycotĂȘtes is generally preferred by the termites, resulting in the high expression of that strain which over time leads to the emergence of a monoculture in the colony. Similar but different looking structures known as primordia also form and are precursors to the sexual fruiting bodies of Termitomyces, although their growth is normally suppressed by the consumption of the primordia by the termites.
530:
55:
515:
31:
348:
307:, back to the mound. This material is chewed up and semi-digested by the termites, fertilised with their faeces and placed in the chambers where it is quickly colonised by the fungus to form a "fungus comb". The termites cultivate these fungus gardens, adding more substrate as required, and removing the older parts of the comb for consumption by all members of the colony.
368:
then molded into the primordial fungus comb. The spores after having survived the passage through the gut of their termite host begin to germinate into homokaryonic hyphae which rapidly colonize the new fungus comb. The resulting homokaryonic hyphae then fuse with unrelated homokaryons of the same species in a process called
273:
often bear fruiting bodies which produce spores, and it is believed that transmission of the fungus to other termites is mainly by horizontal transmission (sibling to sibling) rather than by vertical transmission (mother to daughter). Some species are an exception to this, and in all five species of
254:
Like other termites, Macrotermitinae are soil engineers, mixing their salivary secretions with soil particles to make their strong, hard mounds and galleries. Their mounds are some of the largest built by any species of termite, with volumes of thousands of litres and lasting for many decades. They
224:
Macrotermitinae has a complex colony system. A mature
Macrotermitinae colony consists of a royal pair, sterile caste, winged reproductive called "alates" and young ones. A royal pair are a king and queen which are the only ones capable of reproduction. The sterile caste is made up of workers (major
367:
detritus; broadly consisting of decaying leaf, wood and grass debris with which the spores had settled upon, and which in turn are ingested and partially digested along with the collected detritus. The detritus along with the spores is then formed into round pellets known as mylospheres, which are
375:
Due to this method of sexual reproduction, a mixed culture of many different genotypes exist within the fungus gardens of young
Macrotermitinae colonies. In contrast, mature colonies are known to only have one singularly cloned Termitomyces strain. A monoculture likely arises in a colony through
329:
Shortly after a nuptial flight, the alates quickly remove their wings and set off to form pairs consisting of the male (king) and female (queen) individuals respectively. The fully claustral royal pairs rapidly sequester themselves within the clay rich sandy soils of their environment and form a
195:
and are distinguished by the fact that they cultivate fungi inside their nests to feed the members of the colony. Despite the popular reputation of termites for breaking down and digesting wood, most termite species do not possess the capability to digest the cellulose in wood. Macrotermitinae
289:
They have a rather rigid caste system, with little flexibility after the early instar stage. They also exhibit complex behavioural activities and their presence in an arid or semi-arid area can be dominant over other termite species. As compared to other higher termites however, they show some
376:
positive-frequency dependent selection, in which a genotype outcompetes others via preference by the termites for the most vigorously productive and prolific strain. As the fungus grows, white nutrient rich spheres known as mycotĂȘtes, otherwise known as "nodules" or
303:; it is unclear whether one species of termite is always associated with one species of fungus, and it is probable that several species of termite may utilise a single fungal species. The worker termites bring plant material such as dried grass, decaying wood and
932:
Hsieh, Huei-Mei; Chung, Mei-Chu; Chen, Pao-Yang; Hsu, Fei-Man; Liao, Wen-Wei; Sung, Ai-Ning; Lin, Chun-Ru; Wang, Chung-Ju Rachel; Kao, Yu-Hsin; Fang, Mei-Jane; Lai, Chi-Yung; Huang, Chieh-Chen; Chou, Jyh-Ching; Chou, Wen-Neng; Chang, Bill Chia-Han (2017-09-19).
334:, also known as a claustral chamber. The pairs mate and soon eggs are laid over a couple days, which can take anywhere from 2 - 4 weeks to hatch into several dozen nymphs that can take anywhere between 1 - 3 months to mature into the first workers and soldiers.
204:(fungus-cultivating ants). Worker termites find plant debris and macerate it, chewing and moistening the material. They excrete the resulting fecal pellets inside the mound. Other worker termites use this matter to construct fungal combs. The
407:, in which the royal termite pair are the ones to carry and propagate the fungus every new generation and in which the fungus is always asexually propagated via parent-to-offspring with no sexual reproduction of the symbiont.
245:
analysis places the development of fungiculture several million years after the loss of gut protozoa by the ancestor of the
Termitidae family, which is estimated to have happened between 50 and 80 million years ago.
310:
In addition, some species feed on various types of living and dead plant material including wood, but not on decomposing vegetation; these termites have a similar microbial gut flora to other species of termite.
255:
are probably the most complex mound colonies of any insect group. There are 11 accepted genera in the
Macrotermitinae and about 330 species, with the greatest diversity being in Africa. About 40 species of
563:
216:
The labrum of the imago and worker have a sclerotized transverse band at the posterior section; labrum appears divided by a "hinge" of hyaline tissue. Mesentero-proctodeal junction with four lobes.
541:
208:
then spreads through the comb and digests the plant material into a form that makes for nutritious food for the colony. The mounds are kept humid as possible to encourage rapid fungal growth.
586:
Treatise on the
Isoptera of the world. (Bulletin of the American Museum of Natural History, no. 377) Krishna, Kumar.; Grimaldi, David A.; Krishna, Valerie.; Engel, Michael S. Page: 74
326:
is coordinated with neighboring colonies and triggered by seasonal rainfalls. The nuptial flights of most species are nocturnal in nature although some are crepuscular or diurnal.
355:
Around the time of the maturity and emergence of the first foraging workers, the appearance of fruiting bodies or mushrooms can be observed sprouting from the mounds of mature
141:
488:
380:, begin to grow on the comb and are primarily what the termites eat. The mycotĂȘtes contain substructures known as conidiophores that form asexual spores (
280:
tested, the symbiont fungi did not bear sexual fruiting bodies, and transmission was through the maternal route. Another exception was the single species
322:
like most eusocial insects primarily reproduce through a mass-swarming event known as a nuptial flight, of which the releasing of mature winged sexuals (
495:
391:, the above described is how their Termitomyces symbiont is propagated generation-to-generation. Few known exceptions exist, the most prominent being
644:"Oligocene Termite Nests with In Situ Fungus Gardens from the Rukwa Rift Basin, Tanzania, Support a Paleogene African Origin for Insect Agriculture"
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514:
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418:
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467:
400:
276:
446:
529:
481:
1061:
857:
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363:) which are wind dispersed into the environment. The nanitic workers pick up these spores incidentally as they forage for
569:
Worker termites (Macrotermitinae) closing a newly exposed shaft inside a termite mound to prevent the entry of predators
241:. Fossil evidence from Tanzania show that the Macrotermitinae had developed agriculture about 31 million years ago.
1079:
642:
Roberts EM, Todd CN, Aanen DK, Nobre T, Hilbert-Wolf HL, O'Connor PM, Tapanila L, Mtelela C, Stevens NJ (2016).
54:
237:
of Africa, the Middle East, and southern and southeastern Asia, but it is not present in
Australia or the
785:
Aanen DK, Eggleton P, Rouland-Lefevre C, Guldberg-Froslev T, Rosendahl S, Boomsma JJ (November 2002).
1039:
1092:
1001:
392:
359:, typically between 1/2 - 3 months after nuptial flights. The mushrooms release sexual spores (
282:
40:
935:"A termite symbiotic mushroom maximizing sexual activity at growing tips of vegetative hyphae"
754:
701:
Bucek A, Ć obotnĂk J, He S, Shi M, McMahon DP, Holmes E, Roisin Y, Lo N, Bourguignon T (2019).
616:
1087:
845:
1048:
946:
798:
714:
655:
8:
875:"The Longevity of Colonies of Fungus-Growing Termites and the Stability of the Symbiosis"
262:
950:
802:
718:
659:
977:
934:
909:
874:
678:
643:
49:
821:
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1115:
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732:
683:
622:
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954:
904:
886:
816:
806:
722:
673:
663:
590:
1024:
787:"The evolution of fungus-growing termites and their mutualistic fungal symbionts"
668:
201:
791:
Proceedings of the
National Academy of Sciences of the United States of America
554:
521:
364:
959:
727:
702:
372:, resulting in a heterokaryon with multiple genetically distinguished nuclei.
293:
The mound contains galleries and chambers in which the termites grow fungi as
1109:
968:
900:
242:
891:
784:
703:"Evolution of Termite Symbiosis Informed by Transcriptome-Based Phylogenies"
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918:
830:
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where again the fungus did not fruit, and where transmission was paternal.
257:
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453:
396:
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233:
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177:
126:
30:
595:
238:
197:
181:
173:
106:
86:
66:
995:
873:
Wisselink, Margo; Aanen, Duur K.; van ât Padje, Anouk (2020-08-13).
1018:
381:
205:
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116:
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primitive features and have failed to evolve soil consumption.
150:
96:
76:
756:
347:
188:
621:. Springer Science & Business Media. pp. 736â745.
520:
A Macrotermitinae mound in the
Okavango Delta just outside
872:
535:
A termite soldier (Macrotermitinae) in the Okavango Delta
553:
with a captured worker termite (Macrotermitinae) in the
759:. Springer Science & Business Media. p. 202.
261:
have been identified as symbionts. In contrast to the
641:
700:
196:instead use their mounds to cultivate fungus in a
931:
403:. In the case of the exceptions, the symbiont is
1107:
172:, the fungus-growing termites, constitute a
852:. Cambridge University Press. p. 56.
610:
608:
29:
976:
958:
908:
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820:
810:
726:
677:
667:
614:
346:
850:Production Ecology of Ants and Termites
780:
778:
776:
748:
746:
618:Symbiosis: Mechanisms and Model Systems
605:
1108:
1000:
999:
752:
351:General life cycle of Macrotermitinae
297:. The fungi concerned are species of
843:
837:
773:
743:
694:
846:"Food and Feeds Habits of Termites"
399:and species belonging to the genus
219:
13:
14:
1127:
211:
562:
540:
528:
513:
53:
228:
925:
866:
635:
580:
187:This subfamily consists of 12
180:that is only found within the
1:
574:
314:
669:10.1371/journal.pone.0156847
615:Seckbach J (11 April 2006).
7:
159:
10:
1132:
506:
249:
1008:
960:10.1186/s40529-017-0191-9
728:10.1016/j.cub.2019.08.076
410:
200:relationship, similar to
156:
149:
50:Scientific classification
48:
37:
28:
23:
357:Macrotermitinae colonies
343:Horizontal transmission
892:10.3390/insects11080527
812:10.1073/pnas.222313099
405:vertically transmitted
393:Macrotermes bellicosus
352:
283:Macrotermes bellicosus
41:Macrotermes natalensis
1088:Paleobiology Database
848:. In Brian MG (ed.).
350:
16:Subfamily of termites
713:(21): 3728â3734.e4.
547:An ant of the genus
951:2017BotSt..58...39H
803:2002PNAS...9914887A
719:2019CBio...29E3728B
660:2016PLoSO..1156847R
338:Symbiont life cycle
263:fungus-growing ants
489:Pseudacanthotermes
353:
1103:
1102:
1002:Taxon identifiers
939:Botanical Studies
859:978-0-521-21519-0
766:978-3-540-28180-1
628:978-0-306-48173-4
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870:
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844:Wood TG (1978).
841:
835:
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824:
814:
797:(23): 14887â92.
782:
771:
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753:König H (2006).
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681:
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496:Synacanthotermes
220:Colony structure
202:leaf-cutter ants
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58:
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33:
24:Macrotermitinae
21:
20:
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1004:
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707:Current Biology
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654:(6): e0156847.
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389:Macrotermitinae
320:Macrotermitinae
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170:Macrotermitinae
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137:Macrotermitinae
52:
38:Structure of a
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433:Ancistrotermes
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365:lignocellulose
316:
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212:Identification
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191:and about 350
176:of the family
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440:Euscaiotermes
437:
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430:
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426:Allodontermes
423:
421:
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419:Acanthotermes
416:
415:
408:
406:
402:
398:
395:of the genus
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379:
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361:basidiospores
358:
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475:Odontotermes
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378:sporodochium
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300:Termitomyces
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271:Termitomyces
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258:Termitomyces
256:
253:
243:Phylogenetic
232:
229:Distribution
223:
215:
186:
169:
167:
157:
136:
113:Infraorder:
39:
18:
1034:Wikispecies
468:Microtermes
454:Macrotermes
401:Microtermes
397:Macrotermes
305:leaf litter
277:Microtermes
133:Subfamily:
885:(8): 527.
602:2013-04-25
575:References
557:, Botswana
550:Megaponera
447:Hypotermes
370:plasmogamy
332:copularium
315:Life cycle
274:the genus
184:tropics.
178:Termitidae
127:Termitidae
87:Arthropoda
1025:Q10936976
969:1817-406X
945:(1): 39.
901:2075-4450
596:2246/6430
482:Protermes
239:New World
198:symbiotic
182:Old World
174:subfamily
107:Blattodea
73:Kingdom:
67:Eukaryota
1116:Termites
1110:Category
1019:Wikidata
987:28929370
919:32823564
831:12386341
737:31630948
688:27333288
648:PLOS ONE
387:In most
206:mycelium
158:14, see
123:Family:
117:Isoptera
83:Phylum:
77:Animalia
63:Domain:
1054:2636686
978:5605481
947:Bibcode
910:7469218
879:Insects
799:Bibcode
715:Bibcode
679:4917219
656:Bibcode
507:Gallery
382:conidia
324:alates)
250:Ecology
235:tropics
193:species
103:Order:
97:Insecta
93:Class:
1093:277643
1067:650449
985:
975:
967:
917:
907:
899:
856:
829:
822:137514
819:
763:
735:
686:
676:
625:
411:Genera
269:, the
267:Attini
189:genera
151:Genera
144:, 1934
142:Kemner
44:mound
1080:62955
600:Date:
1075:NCBI
1062:ITIS
983:PMID
965:ISSN
915:PMID
897:ISSN
854:ISBN
827:PMID
761:ISBN
733:PMID
684:PMID
623:ISBN
588:URI:
168:The
160:text
1049:EoL
973:PMC
955:doi
905:PMC
887:doi
817:PMC
807:doi
723:doi
674:PMC
664:doi
591:hdl
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