{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T00:50:00Z","timestamp":1760057400982,"version":"build-2065373602"},"reference-count":40,"publisher":"MDPI AG","issue":"2","license":[{"start":{"date-parts":[[2025,2,11]],"date-time":"2025-02-11T00:00:00Z","timestamp":1739232000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Graduate Competitive Research Grants Program","award":["2019071"],"award-info":[{"award-number":["2019071"]}]},{"name":"state and federal funds appropriated to the OSU CFAES Wooster Campus","award":["2019071"],"award-info":[{"award-number":["2019071"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Insects"],"abstract":"<jats:p>The eastern subterranean termite, Reticulitermes flavipes, is challenged by the significant energy expenditures of tunnel construction for resource discovery. Subterranean termites use idiothetic mechanisms to explore large spaces, while the use of resource-specific cues for localized search is disputed. Here, termite response to wood mulch, termite workers, extracts of wood mulch, and CO2 alone were tested using a bioassay design that distinguished between attraction and arrestment. Termites showed significant attraction to wood mulch with workers or to wood mulch alone. They did not respond to workers alone at the initial dose tested, but were attracted to workers at higher densities. Termites did not respond to water or the acetone extracts of wood mulch, but did show a partial response to hexane extract compared to intact wood mulch. More significantly, when CO2 was removed from the emissions of wood mulch and workers using soda lime, attraction was eliminated. Furthermore, termites showed a quadratic response to CO2 concentration that peaked at ca. 14,000 ppm. The response to CO2 alone predicted by the model matched termite response to mulch + workers when compared at the level of CO2 they emitted. The results suggest that CO2 is both necessary and sufficient to explain the attraction response of R. flavipes to mulch and workers we observed. It is argued that orientation to food cues complements the previously demonstrated idiothetic program to maximize the efficiency of resource location.<\/jats:p>","DOI":"10.3390\/insects16020194","type":"journal-article","created":{"date-parts":[[2025,2,11]],"date-time":"2025-02-11T09:36:05Z","timestamp":1739266565000},"page":"194","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Reticulitermes flavipes (Blattodea: Rhinotermitidae) Response to Wood Mulch and Workers Mediated by Attraction to Carbon Dioxide"],"prefix":"10.3390","volume":"16","author":[{"given":"Tae Young Henry","family":"Lee","sequence":"first","affiliation":[{"name":"Department of Entomology, College of Food, Agriculture and Environmental Sciences, The Ohio State University, Wooster, OH 44691, USA"},{"name":"Department of Entomology and Nematology, University of Florida, 1881 Natural Area Drive, Gainesville, FL 32611, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-1346-3576","authenticated-orcid":false,"given":"P. Larry","family":"Phelan","sequence":"additional","affiliation":[{"name":"Department of Entomology, College of Food, Agriculture and Environmental Sciences, The Ohio State University, Wooster, OH 44691, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2025,2,11]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1016\/j.jinsphys.2014.09.005","article-title":"Burrowing energetics of the giant burrowing cockroach Macropanesthia rhinoceros: An allometric study","volume":"70","author":"Xu","year":"2014","journal-title":"J. Insect Physiol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"802","DOI":"10.1016\/j.biosystems.2007.04.004","article-title":"Optimal length distribution of termite tunnel branches for efficient food search and resource transportation","volume":"90","author":"Lee","year":"2007","journal-title":"Biosystems"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"526","DOI":"10.1007\/BF01134490","article-title":"Non-random search geometry in subterranean termites","volume":"82","author":"Robson","year":"1995","journal-title":"Naturwissenchaften"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1016\/j.ecoinf.2009.04.003","article-title":"The influence of branching tunnels on subterranean termites\u2019 foraging efficiency: Consideration for simulations","volume":"4","author":"Lee","year":"2009","journal-title":"Ecol. Inform."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1016\/j.aspen.2010.01.003","article-title":"Simulation study on the tunnel networks of subterranean termites and their foraging behaviour","volume":"13","author":"Lee","year":"2010","journal-title":"J. Asia-Pac. Entomol."},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Gao, Y., Wen, P., Card\u00e9, R.T., Xu, H., and Huang, Q. (2021). In addition to cryptochrome 2, magnetic particles with olfactory co-receptor are important for magnetic orientation in termites. Commun. Biol., 4.","DOI":"10.1038\/s42003-021-02661-6"},{"key":"ref_7","first-page":"471","article-title":"Characterization of tunneling geometry of subterranean termites (lsoptera: Rhinotermitidae) by computer","volume":"44","author":"Su","year":"2004","journal-title":"Sociobiology"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1299","DOI":"10.1111\/ele.12660","article-title":"Foraging success under uncertainty: Search tradeoffs and optimal space use","volume":"19","author":"Bartumeus","year":"2016","journal-title":"Ecol. Lett."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"2076","DOI":"10.1111\/brv.12883","article-title":"A guide to area-restricted search: A foundational foraging behaviour","volume":"97","author":"Dorfman","year":"2022","journal-title":"Biol. Rev. Camb. Philos. Soc."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Bell, W.J., and Card\u00e9, R.T. (1984). Odor dispersion. Chemical Ecology of Insects, Chapman and Hall, Ltd.. [1st ed.].","DOI":"10.1007\/978-1-4899-3368-3"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1603\/0022-0493-95.1.121","article-title":"Responses of Coptotermes formosanus and Reticulitermes flavipes (Isoptera: Rhinotermitidae) to three types of wood rot fungi cultured on different substrates","volume":"95","author":"Cornelius","year":"2002","journal-title":"J. Econ. Entomol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"476","DOI":"10.1093\/jee\/98.2.476","article-title":"Attraction of subterranean termites (Isoptera) to carbon dioxide","volume":"98","author":"Bernklau","year":"2005","journal-title":"J. Econ. Entomol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1093\/jee\/98.2.471","article-title":"Directional change in tunneling of subterranean termites (Isoptera: Rhinotermitidae) in response to decayed wood attractants","volume":"98","author":"Su","year":"2005","journal-title":"J. Econ. Entomol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"502","DOI":"10.1093\/jee\/98.2.502","article-title":"Effect of summon preferred food source on feeding, tunneling, and bait station discovery by the Formosan subterranean termite (Isoptera: Rhinotermitidae)","volume":"98","author":"Cornelius","year":"2005","journal-title":"J. Econ. Entomol."},{"key":"ref_15","first-page":"1","article-title":"Effect of Chemical Cues on the Foraging and Tunneling Behavior of Formosan Subterranean Termites (Isoptera: Rhinotermitidae)","volume":"8","author":"Cornelius","year":"2015","journal-title":"Midsouth Entomol."},{"key":"ref_16","unstructured":"Suoja, S.B., Lewis, V.R., Wood, D.L., and Wilson, M. (1999). Comparisons of single and group bioassays on attraction and arrestment of Reticulitermes sp. (Isoptera: Rhinotermitidae) to selected cellulosic materials. Sociobiology, 33."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1193","DOI":"10.1603\/0022-0493-94.5.1193","article-title":"Tunnel orientation and search pattern sequence of the Formosan subterranean termite (Isoptera: Rhinotermitidae)","volume":"94","author":"Campora","year":"2001","journal-title":"J. Econ. Entomol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1398","DOI":"10.1603\/0022-0493-94.6.1398","article-title":"Tunnel formation by Reticulitermes flavipes and Coptotermes formosanus (Isoptera: Rhinotermitidae) in response to wood in sand","volume":"94","author":"Puche","year":"2001","journal-title":"J. Econ. Entomol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"511","DOI":"10.1111\/j.1744-7917.2007.00180.x","article-title":"Tunnel geometry of the subterranean termite Reticulitermes grassei (Isoptera: Rhinotermitidae) in response to sand bulk density and the presence of food","volume":"14","author":"Nobre","year":"2007","journal-title":"Insect Sci."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1111\/j.1365-3032.1978.tb00138.x","article-title":"The concepts of olfactory \u2018arrestment\u2019 and \u2018attraction\u2019","volume":"3","author":"Kennedy","year":"1979","journal-title":"Physiol. Entomol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1515\/hfsg.1998.52.1.7","article-title":"Attraction of Steamed Japanese Larch (Larix leptolepis (Sieb, et Zucc.) Gord.) Heartwood to the Subterranean Termite Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae)","volume":"52","author":"Doi","year":"1998","journal-title":"Holzforschung"},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Mitaka, Y., Matsuyama, S., Mizumoto, N., Matsuura, K., and Akino, T. (2020). Chemical identification of an aggregation pheromone in the termite Reticulitermes speratus. Sci. Rep., 10.","DOI":"10.1038\/s41598-020-64388-4"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1007\/s000400050037","article-title":"Systematic search for food in the subterranean termite Reticulitermes santonensis De Feytaud (Isoptera, Rhinotermitidae)","volume":"44","author":"Reinhard","year":"1997","journal-title":"Insectes Sociaux"},{"key":"ref_24","first-page":"159","article-title":"Cues Used by Subterranean Termites during Foraging and Food Assessment","volume":"Volume 1","author":"Khan","year":"2018","journal-title":"Termites and Sustainable Management. Sustainability in Plant and Crop Protection"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1007\/s10682-007-9220-6","article-title":"Consequences of food distribution for optimal searching behavior: An evolutionary model","volume":"23","author":"Scharf","year":"2009","journal-title":"Evol. Ecol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1016\/j.cois.2016.02.009","article-title":"The neurobiological basis of orientation in insects: Insights from the silkmoth mating dance","volume":"15","author":"Namiki","year":"2016","journal-title":"Curr. Opin. Insect Sci."},{"key":"ref_27","unstructured":"Grace, J.K., and Campora, C.E. (2005, January 11\u201313). Food Location and Discrimination by Subterranean Termites (Isoptera: Rhinotermitidae). Proceedings of the Fifth International Conference on Urban Pests, Singapore."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Su, N.-Y., and Lee, C.-Y. (2023). Ecology and foraging behavior. Biology and Management of the Formosan Subterranean Termite and Related Species, CABI. [1st ed.].","DOI":"10.1079\/9781800621596.0000"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"493","DOI":"10.1016\/j.jtbi.2006.07.026","article-title":"Food encounter rates of simulated termite tunnels with variable food size\/distribution pattern and tunnel branch length","volume":"243","author":"Lee","year":"2006","journal-title":"J. Theor. Biol."},{"key":"ref_30","first-page":"203","article-title":"Interlaboratory studies on termite wood decay fungi associations. 2. Response of termites to Gloeophyllum trabeum grown on different species of wood (Isoptera: Mastotermitidae, Termopsidae, Rhinotermitidae, Termitidae)","volume":"18","author":"Lenz","year":"1991","journal-title":"Sociobiology"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.1365-3032.2005.00487.x","article-title":"Chemically-mediated host-plant location and selection by root-feeding insects","volume":"31","author":"Johnson","year":"2006","journal-title":"Physiol. Entomol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"604","DOI":"10.1007\/s10886-012-0106-x","article-title":"Foraging in the dark-chemically mediated host plant location by belowground insect herbivores","volume":"38","author":"Johnson","year":"2012","journal-title":"J. Chem. Ecol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/B978-0-12-417165-7.00003-9","article-title":"Nature, evolution and characterisation of rhizospheric chemical exudates affecting root herbivores","volume":"45","author":"Hiltpold","year":"2013","journal-title":"Adv. Insect Physiol."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Kojima, W. (2015). Attraction to carbon dioxide from feeding resources and conspecific neighbours in larvae of the rhinoceros beetle Trypoxylus dichotomus. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0141733"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1007\/978-3-642-61132-2_5","article-title":"Sensory and behavioural responses of terrestrial invertebrates to biogenic carbon dioxide gradients","volume":"4","author":"Stange","year":"1996","journal-title":"Adv. Bioclimatol."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Paul, E.A. (2015). The soil habitat. Soil Microbiology, Ecology, and Biochemistry, Elsevier Academic Press. [4th ed.].","DOI":"10.1016\/B978-0-12-415955-6.00001-3"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1080\/03650340.2010.506482","article-title":"Estimation of CO2 diffusion coefficient at 0\u201310 cm depth in undisturbed and tilled soils","volume":"58","author":"Jabro","year":"2012","journal-title":"Arch. Agron. Soil Sci."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"716","DOI":"10.1016\/j.tree.2020.04.001","article-title":"Plant secondary compounds in soil and their role in belowground species interactions","volume":"35","author":"Ehlers","year":"2020","journal-title":"Trends Ecol. Evol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"e65575","DOI":"10.7554\/eLife.65575","article-title":"Plant-associated CO2 mediates long-distance host location and foraging behavior of a root herbivore","volume":"10","author":"Arce","year":"2021","journal-title":"eLife"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1016\/j.ecocom.2015.03.002","article-title":"Composite random search strategies based on non-directional sensory cues","volume":"22","author":"Nolting","year":"2015","journal-title":"Ecol. Complex."}],"container-title":["Insects"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2075-4450\/16\/2\/194\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T16:31:00Z","timestamp":1760027460000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2075-4450\/16\/2\/194"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,2,11]]},"references-count":40,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2025,2]]}},"alternative-id":["insects16020194"],"URL":"https:\/\/doi.org\/10.3390\/insects16020194","relation":{},"ISSN":["2075-4450"],"issn-type":[{"type":"electronic","value":"2075-4450"}],"subject":[],"published":{"date-parts":[[2025,2,11]]}}}