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Grassroots Journal of Natural Resources

Coordinated and published by the Grassroots Institute, the Grassroots Journal of Natural Resources (GJNR) is an international journal dedicated to the latest advancements in natural resources throughout the world. The goal of this journal is to provide a platform for scientists, social scientists, policy analysts, managers and practitioners (on all academic and professional levels) all over the world to promote, discuss and share various new issues and developments in different arenas of natural resources.


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M – 00244Review Article

Articulating Fragrant Agarwood Formation as an Outcome of the Interaction between the Insect Zeuzera conferta and Aquilaria trees – A Review

Arup Khakhlari*1, Supriyo Sen2

1Department of Biosciences, Assam Don Bosco University, Tapesia Campus, Sonapur, Assam, India.

Email: | ORCID:

2Department of Biosciences, Assam Don Bosco University, Tapesia Campus, Sonapur, Assam, India.

Email:; | ORCID ID: 0000-0002-3800-4942

*Corresponding author

Grassroots Journal of Natural Resources, 4(3): 62-78. Doi:

Received: 17 August 2021

Reviewed: 30 August 2021

Provisionally Accepted: 10 September 2021

Revised: 21 September 2021

Finally Accepted: 24 September 2021

Published: 30 September 2021


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Agarwood is the resinous infected wood obtained from Aquilaria species, which is a highly priced product in the flavour and fragrance market. Its formation is a complex process of interaction between the plant, insect, and microorganisms. Multiple studies concerning the interaction of microorganisms with the Aquilaria tree have been reported. However, the significant interaction between the insect Zeuzera conferta Walker (Lepidoptera: Cossidae) with Aquilaria has been overlooked, and only exiguous studies have been accomplished. Considering the dearth of available literature on this interesting phenomenon a review has been attempted. The taxonomical and morphological descriptions proffered by researchers and the insect life cycle are discussed. The review lays emphasis on the chemical ecology of the interaction between Z. conferta, Aquilaria and associating microorganisms as a possible continuum operating in the form of complex chemical signalling via release and sensing of Volatile Organic Compounds (VOCs), Herbivore Induced Plant Volatiles (HIPVs) and Microbial Volatile Organic Compounds (MVOCs). The review also scrutinizes the future perspectives of understanding the interaction in devising suitable management strategies to prevent uncontrolled infestation and, simultaneously, develop artificial rearing technology for the insect Z. conferta as a strategy for ensuring sustainable livelihood of farmers dependent on agarwood production.


Insecticides; Frass; Taxonomy; Artificial rearing; Interaction; Lepidopteran


Aartsma, Y., Leroy, B., van der Werf, W., Dicke, M., Poelman, E.H. and Bianchi, F.J.J.A. (2018).

Abdel-Moaty, R.M., Hashim, S.M. and Tadros, A.W. (2019). The Impact of the Leopard Moth Zeuzera

Aljbory, Z. and Chen, M.S. (2018). Indirect plant defense against insect herbivores: a review. Insect

Arora, G.S. (1976). A Taxonomic revision of the Indian species of the family Cossidae (Lepidoptera).

Atreya, K., Johnsen, F.H. and Sitaula, B.K. (2012). Health and environmental costs of pesticide use in

Baksha, M.W. and Islam, M.R. (1999). Biology and ecology of Zeuzera conferta Walker (Cossidae:

Bennett, R.N. and Wallsgrove, R.M. (1994). Tansley Review No. 72 Secondary metabolites in plant

Borthakur, N.D., Borah, R.K., Dutta, B.K. and Jayaraj, R.S.C. (2021). Neurozerra conferta Walker.

Breznak, J.A. (1982). Intestinal Microbiota of Termites and other Xylophagous Insects. Annual

Chen, D.Q. and Purcell, A. (1997). Occurrence and Transmission of Facultative Endosymbionts in

Chhipa, H. and Kaushik, N. (2017). Fungal and Bacterial Diversity Isolated from Aquilaria

Davis, T.S., Crippen, T.L., Hofstetter, R.W. and Tomberlin, J.K. (2013). Microbial Volatile Emissions as

Davis, T.S., Hofstetter, R.W., Foster, J.T., Foote, N.E. and Keim, P. (2011). Interactions between the

Dicke, M. and Baldwin, I.T. (2010). The evolutionary context for herbivore-induced plant volatiles:

Dudareva, N., Negre, F., Nagegowda, D.A. and Orlova, I. (2006). Plant volatiles: recent advances and

Ezenwa, V.O., Gerardo, N.M., Inouye, D.W., Medina, M. and Xavier, J.B. (2012). Animal behavior and

Fatouros, N.E., Bukovinszkine’Kiss, G., Kalkers, L.A., Gamborena, R.S., Dicke, M. and Hilker, M.

Fatouros, N.E., Dicke, M., Mumm, R., Meiners, T. and Hilker, M. (2008). Foraging behavior of egg

Feeny, P., Rosenberry, L. and Carter, M. (1983). Chemical aspects of oviposition behavior in

Field, L.M., Pickett, J.A. and Wadhams, L.J. (2008). Molecular studies in insect olfaction. Insect

Forbes, A.A., Bagley, R.K., Beer, M.A., Hippee, A.C. and Widmayer, H.A. (2018). Quantifying the

Giron, D., Dubreuil, G., Bennett, A., Dedeine, F., Dicke, M., Dyer, L.A., Erb, M., Harris, M.O., Huguet, E.,

Haine, E.R., Moret, Y., Siva-Jothy, M.T. and Rolff, J. (2008). Antimicrobial defense and persistent

Hilker, M. and Meiners, T. (2010). How do plants “notice” attack by herbivorous arthropods?

Hilker, M., Kobs, C., Varama, M. and Schrank, K. (2002). Insect egg deposition induces Pinus to

Hilker, M., Stein, C., Schroeder, R., Varama, M. and Mumm, R. (2005). Insect egg deposition induces

Hogenhout, S.A. and Bos, J.I. (2011). Effector proteins that modulate plant-insect interactions.

Holloway, J.D. (1986). The moths of Borneo part 1, key to families, Families Cossidae, Metarbelidae,

Hoque, M.N., Khan, M.M.H. and Mondal, M.F. (2019). Insect infested agarwood: A newly prized

Howe, G.A. and Jander, G. (2008). Plant immunity to insect herbivores. Annual Review of Plant

Ibrahim, R., Alhamadi, S., Binnaser, Y.S. and Shawer, D. (2019). Seasonal prevalence and

Irianto, R.S.B., Santoso, E. and Sitepu, I.R. (2011). Pests that attack gaharu-yielding plants. In

Islam, M.A. (2004). A monograph on Keora (Sonneratia apetala). Forestry and Wood technology

Kalita, J., Bhattacharyya, P.R., Deka Boruah, H.P., Unni, B.G., Lekhak, H. and Nath, S.C. (2015).

Kaloshian, I. and Walling, L.L. (2016). Plant Immunity: Connecting the Dots Between Microbial

Kessler, A. and Heil, M. (2011). The multiple faces of indirect defences and their agents of natural

Leal, W.S. (2013). Odorant reception in insects: roles of receptors, binding proteins, and degrading

Lemke, T., Stingl, U., Egert, M., Friedrich, M.W. and Brune, A. (2003). Physicochemical Conditions

Loon, J.J.A. (1996). Chemosensory basis of feeding and oviposition behaviour in herbivorous

Lowery, C.A., Dickerson, T.J. and Janda, K.D. (2008). Interspecies and interkingdom communication

Ma, Q., Fonseca, A., Liu, W., Fields, A.T., Pimsler, M.L., Spindola, A.F., Tarone, A.M., Crippen, T.L.,

Mithofer, A. and Boland, W. (2012). Plant defense against herbivores: chemical aspects. Annual

Mohamed, R., Jong, P.L. and Zali, M.S. (2010). Fungal diversity in wounded stems of Aquilaria

Moore, I. and Navon, A. (1966). The rearing and some bionomics of the leopard moth, Zeuzera

Morris, W.F. and Kareiva, P.M. (1991). How insect herbivores find suitable host plants: the interplay

Nago, H. and Matsumoto, M. (1994). An Ecological Role of Volatiles Produced by Lasiodiplodia

Nath, S.C. and Saikia, N. (2002). Indigenous knowledge on utility and Utilitarian aspects of Aquilaria

Ong, S.P., Cheng, S., Chong, V.C. and Tan, Y.S. (2010). Pests of Planted Mangroves in Peninsular

Ong, S.P., Mohd Farid, A. and Lee, S.S. (2014). Pest and disease survey of Aquilaria sp. (karas)

Paré, P.W. and Tumlinson, J.H. (1999). Plant volatiles as a defense against insect herbivores. Plant

Penaflor, M.F.G.V. and Bento, J.M.S. (2013). Herbivore-Induced Plant Volatiles to Enhance Biological

Pichersky, E. and Gershenzon, J. (2002). The formation and function of plant volatiles: perfumes for

Price, P.W., Denno, R.F., Eubanks, M.D., Finke, D.L. and Kaplan, I. (2011). Insect ecology: behavior,

Qiao, H.L., Lu, P.F., Chen, J., Ma, W.S., Qin, R.M. and Li, X.M. (2012). Antennal and behavioural

Renwick, J.A.A. (1989). Chemical ecology of oviposition in phytophagous insects. Experientia, 45(3):

Renwick, J.A.A. and Chew, F.S. (1994). Oviposition behaviour in Lepidoptera. Annual Review on

Reynolds, B.C. and Hunter, M.D. (2004). Nutrient Cycling. In MD Lowman, In HB Rinker (eds),

Reynolds, B.C., Crossley, D.A. and Hunter, M.D. (2003). Responses of soil invertebrates to forest

Roepke, W. (1955). Notes and description of Cossidae from New Guinea (Lepidoptera: Heterocera).

Roepke, W. (1957). The Cossidae of the Malayan Region (Lepidoptera: Heterocera). Verhandel

Rozen, D.E., Engelmoer, D.J.P. and Smiseth, P.T. (2008). Antimicrobial strategies in burying beetles

Ryu, C-M., Farag, M.A., Hu, C-H., Reddy, M.S., Kloepper, J.W. and Pare, P.W. (2004). Bacterial volatiles

Ryu, C-M., Faragt, M.A., Hu, C-H., Reddy, M.S., Wei, H-X., Paré, P.W. and Kloepper, J.W. (2003).

Salerno, G., De Santis, F., Iacovone, A., Bin, F. and Conti, E. (2013). Short-range cues mediate

Schiestl, F.P. (2010). The evolution of floral scent and insect chemical communication. Ecology

Schoorl, J.W. (Pim) Jr. (1990). Phylogenetic study on Cossidae (Lepidoptera: Ditrysia) based on

Sen, S., Dehingia, M., Talukdar, N.C. and Khan, M. (2017). Chemometric analysis reveals links in the

Senthilkumar, N. and Murugesan, S. (2015). Insect pests of important trees species in South India

Shamseldean, M.M., Hasanain, S.A. and Rezk, M.Z.A. (2009). Virulence of Entomopathogenic

Sutrisno, H. (2015). Molecular phylogeny of Indonesian Zeuzera (Lepidoptera: Cossidae) wood

Syazwan, S.A., Lee, S.Y., Ong, S.P. and Mohamed, R. (2019). Damaging Insect Pests and Diseases and

Tomberlin, J.K., Byrd, J.H., Wallace, J.R. and Benbow, M.E. (2012). Assessment of decomposition

Vannucci, M. (2002). Indo-West Pacific Mangroves. In: de Lacerda L.D. (eds), Mangrove

Walker, F. (1856). List of the specimens of Lepidopterous insects in the collection of the British

Yakovlev, R.V. (2011). Catalogue of the Family Cossidae of the Old World. Neue Entomologische

Zhang, X.L., Liu, Y.Y., Wei, J.H., Yang, Y., Zhang, Z., Huang, J.Q., Chen, H.Q. and Liu, Y.J. (2012).

Harvard Style

Khakhlari, A. and Sen, S. (2021). Articulating Fragrant Agarwood Formation as an Outcome of the Interaction between the Insect Zeuzera conferta and Aquilaria trees – A Review. Grassroots Journal of Natural Resources, 4(3): 62-78. Doi:

APA Style

Khakhlari, A., & Sen, S. (2021). Articulating Fragrant Agarwood Formation as an Outcome of the Interaction between the Insect Zeuzera conferta and Aquilaria trees – A Review. Grassroots Journal of Natural Resources, 4(3), 62-78.

Chicago/Turabian Style

KaKhakhlari, Arup, Sen, Supriyo. 2021. “Articulating Fragrant Agarwood Formation as an Outcome of the Interaction between the Insect Zeuzera conferta and Aquilaria trees – A Review. Grassroots Journal of Natural Resources, 4 no. 3: 62-78.

ACS Style

Khakhlari A., Sen S. Articulating Fragrant Agarwood Formation as an Outcome of the Interaction between the Insect Zeuzera conferta and Aquilaria trees – A Review. Grassroots Journal of Natural Resources, 2021, 4 (3), 62-78.

AAA Style

Khakhlari, Arup and Supriyo Sen. 2021. “Articulating Fragrant Agarwood Formation as an Outcome of the Interaction between the Insect Zeuzera conferta and Aquilaria trees – A Review. Grassroots Journal of Natural Resources, 4 (3): 62-78.

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