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This species displays complex defensive behaviors when a breach of the mound occurs. In shallow breaches, a few minor and major soldiers rush out to face any threats while the workers retreat deeper into the nest, a normal behavior common to other Macrotermes species. However in the event of a deep breach, the soldiers assume defensive positions around the perimeter of the breach and begin hammering their heads against the walls of the mound in synchrony, creating a rhythmic rattling noise. It is believed this behavior evolved as a warning due to predation by vertebrates.<ref>{{Cite web |url=https://academic.oup.com/zoolinnean/article-abstract/26/169/358/2732839 |access-date=2022-12-05 |website=academic.oup.com}}</ref> The curving mandibles of the soldiers make them difficult to pry off the skin, with bites commonly occurring when people accidentally step onto the foraging trails. The soldiers also possess a defensive fluid produced from the salivary glands which is composed of lauric acid methyl ester, [[quinone]]s and 3 other unknown compounds. The defensive fluid is believed to be primarily insecticidal via action of the [[lauric acid]], and is thought to aid against predation by [[ant]]s.<ref>{{Cite journal |last=Diyana |first=M. H. Alia |last2=Appalasamy |first2=S. |last3=Arumugam |first3=N. |last4=Boon |first4=J. G. |date=July 2019 |title=A study of a termite chemical defense fluid compound of Macrotermes carbonarius |url=https://dx.doi.org/10.1088/1755-1315/269/1/012009 |journal=IOP Conference Series: Earth and Environmental Science |language=en |volume=269 |issue=1 |pages=012009 |doi=10.1088/1755-1315/269/1/012009 |issn=1755-1315}}</ref>
This species displays complex defensive behaviors when a breach of the mound occurs. In shallow breaches, a few minor and major soldiers rush out to face any threats while the workers retreat deeper into the nest, a normal behavior common to other Macrotermes species. However in the event of a deep breach, the soldiers assume defensive positions around the perimeter of the breach and begin hammering their heads against the walls of the mound in synchrony, creating a rhythmic rattling noise. It is believed this behavior evolved as a warning due to predation by vertebrates.<ref>{{Cite web |url=https://academic.oup.com/zoolinnean/article-abstract/26/169/358/2732839 |access-date=2022-12-05 |website=academic.oup.com}}</ref> The curving mandibles of the soldiers make them difficult to pry off the skin, with bites commonly occurring when people accidentally step onto the foraging trails. The soldiers also possess a defensive fluid produced from the salivary glands which is composed of lauric acid methyl ester, [[quinone]]s and 3 other unknown compounds. The defensive fluid is believed to be primarily insecticidal via action of the [[lauric acid]], and is thought to aid against predation by [[ant]]s.<ref>{{Cite journal |last=Diyana |first=M. H. Alia |last2=Appalasamy |first2=S. |last3=Arumugam |first3=N. |last4=Boon |first4=J. G. |date=July 2019 |title=A study of a termite chemical defense fluid compound of Macrotermes carbonarius |url=https://dx.doi.org/10.1088/1755-1315/269/1/012009 |journal=IOP Conference Series: Earth and Environmental Science |language=en |volume=269 |issue=1 |pages=012009 |doi=10.1088/1755-1315/269/1/012009 |issn=1755-1315}}</ref>


''M. carbonarius'' is amongst the most free-ranging of Macrotermes species found in the indomalayan tropics. It forages above the ground from numerous large subterranean tunnels that can extend between 33.8 – 112.8 metres from the nest before emerging onto the surface.<ref>{{Cite journal |last1=Inoue |first1=T. |last2=Kirtibutr |first2=N. |last3=Abe |first3=T. |date=2001-12-01 |title=Underground passage system of Macrotermes carbonarius (Isoptera, Termitidae) in a dry evergreen forest of northeast Thailand |url=https://doi.org/10.1007/PL00001794 |journal=Insectes Sociaux |language=en |volume=48 |issue=4 |pages=372–377 |doi=10.1007/PL00001794 |issn=1420-9098 |s2cid=44923083}}</ref> Foraging trails may also then extend several metres above the surface and can be readily recognized by the positioning of a large amount of major and minor soldiers at the flanks. The trails are paved with soil pellets around the exit holes for some distance, and most notably exhibit a trunk structure with branching forays upstream. Workers primarily forage for leaf litter which constitutes the principal diet of this species, although they will readily take decaying hardwood when available.<ref name=":0" /> Other sources such as palm fronds or dried blades of grass are also occasionally taken. These termites are more readily tolerant of desiccation due to their more sclerotized exoskeleton, which allows them to forage for longer periods in areas that experience harsher conditions that other sympatric Macrotermes species would find intolerable.<ref name=":1" /> They are primarily active during the night and twilight hours with foraging activities continuing longer in the day in more favorable weather such as humid and overcast conditions.<ref>{{Cite journal |last1=Matsumoto |first1=Tadao |last2=Abe |first2=Takuya |date=1979-01-01 |title=The role of termites in an equatorial rain forest ecosystem of West Malaysia |url=https://doi.org/10.1007/BF00345187 |journal=Oecologia |language=en |volume=38 |issue=3 |pages=261–274 |doi=10.1007/BF00345187 |pmid=28309487 |s2cid=25643181 |issn=1432-1939}}</ref><ref name=":1">{{Cite journal |last1=Hu |first1=Jian |last2=Neoh |first2=Kok-Boon |last3=Appel |first3=Arthur G. |last4=Lee |first4=Chow-Yang |date=February 2012 |title=Subterranean termite open-air foraging and tolerance to desiccation: Comparative water relation of two sympatric Macrotermes spp. (Blattodea: Termitidae) |url=https://pubmed.ncbi.nlm.nih.gov/22085890/ |journal=Comparative Biochemistry and Physiology. Part A, Molecular & Integrative Physiology |volume=161 |issue=2 |pages=201–207 |doi=10.1016/j.cbpa.2011.10.028 |issn=1531-4332 |pmid=22085890}}</ref>
''M. carbonarius'' is amongst the most free-ranging of Macrotermes species found in the indomalayan tropics. It forages above the ground from numerous large subterranean tunnels that can extend between 33.8 – 112.8 metres from the nest before emerging onto the surface.<ref>{{Cite journal |last1=Inoue |first1=T. |last2=Kirtibutr |first2=N. |last3=Abe |first3=T. |date=2001-12-01 |title=Underground passage system of Macrotermes carbonarius (Isoptera, Termitidae) in a dry evergreen forest of northeast Thailand |url=https://doi.org/10.1007/PL00001794 |journal=Insectes Sociaux |language=en |volume=48 |issue=4 |pages=372–377 |doi=10.1007/PL00001794 |issn=1420-9098 |s2cid=44923083}}</ref> Foraging trails may also then extend several metres above the surface and can be readily recognized by the positioning of a large amount of major and minor soldiers at the flanks. The trails are paved with soil pellets around the exit holes for some distance, and most notably exhibit a trunk structure with branching forays upstream. Workers primarily forage for leaf litter which constitutes the principal diet of this species, although they will readily take decaying hardwood when available.<ref name=":0" /> Other sources such as palm fronds or dried blades of grass are also occasionally taken. These termites are more readily tolerant of desiccation due to their more sclerotized [[Arthropod cuticle|exoskeleton]], which enables them to forage for longer periods in conditions that other sympatric ''Macrotermes'' species would find intolerable.<ref name=":1" /> They are primarily active during the night and twilight hours with foraging activities continuing longer in the day in more favorable weather such as humid and overcast conditions.<ref>{{Cite journal |last1=Matsumoto |first1=Tadao |last2=Abe |first2=Takuya |date=1979-01-01 |title=The role of termites in an equatorial rain forest ecosystem of West Malaysia |url=https://doi.org/10.1007/BF00345187 |journal=Oecologia |language=en |volume=38 |issue=3 |pages=261–274 |doi=10.1007/BF00345187 |pmid=28309487 |s2cid=25643181 |issn=1432-1939}}</ref><ref name=":1">{{Cite journal |last1=Hu |first1=Jian |last2=Neoh |first2=Kok-Boon |last3=Appel |first3=Arthur G. |last4=Lee |first4=Chow-Yang |date=February 2012 |title=Subterranean termite open-air foraging and tolerance to desiccation: Comparative water relation of two sympatric Macrotermes spp. (Blattodea: Termitidae) |url=https://pubmed.ncbi.nlm.nih.gov/22085890/ |journal=Comparative Biochemistry and Physiology. Part A, Molecular & Integrative Physiology |volume=161 |issue=2 |pages=201–207 |doi=10.1016/j.cbpa.2011.10.028 |issn=1531-4332 |pmid=22085890}}</ref>


The fungus comb of this species is notably less complex than that of its relatives, with a less defined airy grooved pattern common to other ''Macrotermitinae''. The termites, rather than continuously build upon old fungus comb as it is gradually eaten, instead consume the entire comb once it has matured and been completed. After being fully consumed, a new fungus comb is constructed in its place.<ref>{{Cite journal |last1=Hyodo |first1=Fujio |last2=Azuma |first2=Jun Ichi |last3=Abe |first3=Takuya |date=1999-12-01 |title=A new pattern of lignin degradation in the fungus comb of Macrotermes carbonarius (Isoptera, Termitidae Macrotermitinae) |url=https://okayama.pure.elsevier.com/en/publications/a-new-pattern-of-lignin-degradation-in-the-fungus-comb-of-macrote |journal=Sociobiology |language=English |volume=34 |issue=3 |pages=591–596 |issn=0361-6525}}</ref>
The fungus comb of this species is notably less complex than that of its relatives, with a less defined airy grooved pattern common to other ''Macrotermitinae''. The termites, rather than continuously build upon old fungus comb as it is gradually eaten, instead consume the entire comb once it has matured and been completed. After being fully consumed, a new fungus comb is constructed in its place.<ref>{{Cite journal |last1=Hyodo |first1=Fujio |last2=Azuma |first2=Jun Ichi |last3=Abe |first3=Takuya |date=1999-12-01 |title=A new pattern of lignin degradation in the fungus comb of Macrotermes carbonarius (Isoptera, Termitidae Macrotermitinae) |url=https://okayama.pure.elsevier.com/en/publications/a-new-pattern-of-lignin-degradation-in-the-fungus-comb-of-macrote |journal=Sociobiology |language=English |volume=34 |issue=3 |pages=591–596 |issn=0361-6525}}</ref>

Revision as of 08:02, 12 April 2023

Macrotermes carbonarius
Soldiers and workers
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Blattodea
Infraorder: Isoptera
Family: Termitidae
Genus: Macrotermes
Species:
M. carbonarius
Binomial name
Macrotermes carbonarius
(Hagen, 1858)

Macrotermes carbonarius,[1] also known as Kongkiak in Malay, is a large black species of fungus-growing termite in the genus Macrotermes. It is one of the most conspicuous species of Macrotermes found in the Indomalayan tropics, forming large foraging trails in the open that can extend several metres in distance. M. carbonarius is a highly aggressive species with the soldiers possessing large curving mandibles that easily break skin. It is found in Cambodia, Indonesia, Malaysia, Myanmar, Singapore, Thailand and Vietnam.[2]

Identification

Soldiers, workers and imagoes of Macrotermes carbonarius are easily distinguished from other species in its range due to the almost jet-black coloration of their exoskeleton.

Like other Macrotermes species, M. carbonarius has a dimorphic worker and soldier caste, with the largest soldiers commonly reaching lengths of 15 – 18 mm.

The imagoes can closely resemble the imagoes of some species of Odontotermes, however can be distinguished by the proportionally smaller eyes and overall larger size of M. carbonarius imagoes which can attain a body length of 15 – 16 mm.

Biology

M. carbonarius is confined to elevations below 160 metres and primarily inhabits the tropical to subtropical lowland forests of Southeast Asia. It can be found across a wide variety of forested flat lands, most commonly in dipterocarp and coastal forests as well as rural areas and plantations.[3]

The nests are large and conspicuous, broad and irregular in shape. No obvious ventilation system exists within the mound walls. The walls of the mound are thick and compact, varying 11-18 in width, and the height of the mound ranges from 30–48 cm. Majority of the colony and the fungus gardens resides either just slightly below or above ground level or entirely within the mound itself.[4]

This species displays complex defensive behaviors when a breach of the mound occurs. In shallow breaches, a few minor and major soldiers rush out to face any threats while the workers retreat deeper into the nest, a normal behavior common to other Macrotermes species. However in the event of a deep breach, the soldiers assume defensive positions around the perimeter of the breach and begin hammering their heads against the walls of the mound in synchrony, creating a rhythmic rattling noise. It is believed this behavior evolved as a warning due to predation by vertebrates.[5] The curving mandibles of the soldiers make them difficult to pry off the skin, with bites commonly occurring when people accidentally step onto the foraging trails. The soldiers also possess a defensive fluid produced from the salivary glands which is composed of lauric acid methyl ester, quinones and 3 other unknown compounds. The defensive fluid is believed to be primarily insecticidal via action of the lauric acid, and is thought to aid against predation by ants.[6]

M. carbonarius is amongst the most free-ranging of Macrotermes species found in the indomalayan tropics. It forages above the ground from numerous large subterranean tunnels that can extend between 33.8 – 112.8 metres from the nest before emerging onto the surface.[7] Foraging trails may also then extend several metres above the surface and can be readily recognized by the positioning of a large amount of major and minor soldiers at the flanks. The trails are paved with soil pellets around the exit holes for some distance, and most notably exhibit a trunk structure with branching forays upstream. Workers primarily forage for leaf litter which constitutes the principal diet of this species, although they will readily take decaying hardwood when available.[3] Other sources such as palm fronds or dried blades of grass are also occasionally taken. These termites are more readily tolerant of desiccation due to their more sclerotized exoskeleton, which enables them to forage for longer periods in conditions that other sympatric Macrotermes species would find intolerable.[8] They are primarily active during the night and twilight hours with foraging activities continuing longer in the day in more favorable weather such as humid and overcast conditions.[9][8]

The fungus comb of this species is notably less complex than that of its relatives, with a less defined airy grooved pattern common to other Macrotermitinae. The termites, rather than continuously build upon old fungus comb as it is gradually eaten, instead consume the entire comb once it has matured and been completed. After being fully consumed, a new fungus comb is constructed in its place.[10]

Reproduction

Nuptial flights occur during daylight in the late afternoon, usually ~30 minutes before sunset. Windless, hot and dry conditions in the absence of rain following a day of heavy rain is preferred.

Horizontally elongated openings known as flight holes are constructed and opened around midday. If favorable conditions persist up to the late afternoon, massive nuptial flights consisting of hundreds or thousands alates are released by colonies. Flights are short in duration and last around 4–10 minutes. Flights occur as early as mid October, peaking in November, with smaller sporadic flights occurring from December to January.[11]

References

  1. ^ Hagen HA 1858. Monographie der Termiten. Linnaea Entomologica 12: i-iii + 4-342 + 459.
  2. ^ Termite catalogue
  3. ^ a b Effects of quality and quantity of food on the foraging area of Macrotermes carbonarius in tropical rain forest. Siew Chin Chua, Juei Ling Hsu INTERNATIONAL FIELD BIOLOGY COURSE 2005, 2005
  4. ^ McCOMIE, L. D.; Dhanarajan, G. (September 1993). "The physical and chemical composition of mounds of Macrotermes carbonarius (Hagen) (Termitidae, Macrotermitinae), in Penang, Malaysia". Journal of Soil Science. 44 (3): 427–433. doi:10.1111/j.1365-2389.1993.tb00465.x.
  5. ^ academic.oup.com https://academic.oup.com/zoolinnean/article-abstract/26/169/358/2732839. Retrieved 2022-12-05. {{cite web}}: Missing or empty |title= (help)
  6. ^ Diyana, M. H. Alia; Appalasamy, S.; Arumugam, N.; Boon, J. G. (July 2019). "A study of a termite chemical defense fluid compound of Macrotermes carbonarius". IOP Conference Series: Earth and Environmental Science. 269 (1): 012009. doi:10.1088/1755-1315/269/1/012009. ISSN 1755-1315.
  7. ^ Inoue, T.; Kirtibutr, N.; Abe, T. (2001-12-01). "Underground passage system of Macrotermes carbonarius (Isoptera, Termitidae) in a dry evergreen forest of northeast Thailand". Insectes Sociaux. 48 (4): 372–377. doi:10.1007/PL00001794. ISSN 1420-9098. S2CID 44923083.
  8. ^ a b Hu, Jian; Neoh, Kok-Boon; Appel, Arthur G.; Lee, Chow-Yang (February 2012). "Subterranean termite open-air foraging and tolerance to desiccation: Comparative water relation of two sympatric Macrotermes spp. (Blattodea: Termitidae)". Comparative Biochemistry and Physiology. Part A, Molecular & Integrative Physiology. 161 (2): 201–207. doi:10.1016/j.cbpa.2011.10.028. ISSN 1531-4332. PMID 22085890.
  9. ^ Matsumoto, Tadao; Abe, Takuya (1979-01-01). "The role of termites in an equatorial rain forest ecosystem of West Malaysia". Oecologia. 38 (3): 261–274. doi:10.1007/BF00345187. ISSN 1432-1939. PMID 28309487. S2CID 25643181.
  10. ^ Hyodo, Fujio; Azuma, Jun Ichi; Abe, Takuya (1999-12-01). "A new pattern of lignin degradation in the fungus comb of Macrotermes carbonarius (Isoptera, Termitidae Macrotermitinae)". Sociobiology. 34 (3): 591–596. ISSN 0361-6525.
  11. ^ Neoh, Kok-Boon; Lee, Chow-Yang (December 2009). "Flight activity of two sympatric termite species, Macrotermes gilvus and Macrotermes carbonarius (Termitidae: Macrotermitinae)". Environmental Entomology. 38 (6): 1697–1706. doi:10.1603/022.038.0623. ISSN 1938-2936. PMID 20021766.
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