{"paper_id":"5967884f-2913-4079-87f6-a78a9d2ea024","body_text":"Temperature regulation of Heterotrigona itama (Cockerell, 1918) in lamp posts nests | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Short Report Temperature regulation of Heterotrigona itama (Cockerell, 1918) in lamp posts nests Florina Anthony, Sze Huei Yek This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4845014/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 08 Nov, 2024 Read the published version in Insectes Sociaux → Version 1 posted 5 You are reading this latest preprint version Abstract The commercial stingless bee Heterotrigona itama naturally nests in tree cavities but was kept in wooden boxes in meliponiculture farms. However, at Universiti Malaysia Sabah (UMS), these bees primarily nest in lamp posts. We conducted a temperature survey to assess lamp posts as potential nesting sites for H. itama , aiming to determine if the preference for occupied lamp posts was related to their distribution. We measured ambient temperatures in occupied lamp posts in shaded areas and unoccupied lamp posts in exposed areas on the UMS campus, calculating the percentage of time these temperatures fell outside the optimal range for H. itama . Additionally, we analyzed the occurrence and timing of temperature regulation in lamp post nests by comparing temperature differences between occupied and unoccupied lamp posts across four-time blocks. Temperature measurements of occupied (shaded) and unoccupied (exposed) lamp posts revealed that shaded lamp posts experienced temperatures outside the bees' optimal range (27°C-32°C) less often than exposed lamp posts (68.4% vs. 78.12%). This suggests that H. itama may prefer shaded lamp posts due to their more stable temperature profile. Additionally, the internal temperature of lamp posts, whether occupied or not, was consistently 1.54–1.76°C warmer than ambient during hotter periods and closer to ambient during cooler periods, indicating inherent insulation properties of the metal lamp posts. However, a notable difference in temperature between occupied and unoccupied posts was observed in the late afternoon and evening, suggesting active thermoregulation by bees to maintain optimal nest temperature. thermoregulation meliponiculture nest sites Universiti Malaysia Sabah main campus Figures Figure 1 Figure 2 Figure 3 Introduction Stingless bees, known as meliponines, are diverse bees in tropical and subtropical regions worldwide (Roubik, 2022 ). These bees are important pollinators of a wide range of crops (Heard, 1999 ; Amano et al., 2000 ; Slaa et al., 2006 ) and play a vital role in the health and functioning of ecosystems (Paz et al., 2021; Campbell et al., 2022 ). In addition to their ecological importance, stingless bees have a long history of use in meliponiculture and are valued for their honey production (Kwapong et al., 2010 ). Malaysia has over 40 known species of stingless bees (Rasmussen, 2008 ). Some common commercial species widely found in meliponiculture farms in Malaysia are Heterotrigona itama , Geniotrigona thoracica , and Lepidotrigona terminata (Msn et al., 2014; Mustafa et al., 2018 ). These species occurred widely in urban areas and forest edges where meliponiculture farms were located (Hamid et al., 2016 ). Due to their survivability in the human-adapted landscape, they are priced as honey producers and pollinators in Malaysia’s meliponiculture industries (Kelly et al., 2014 ; Fahimee et al., 2016 , 2018 ; Ghazi et al., 2018 ). A critical factor determining a stingless bee colony’s success is selecting a suitable nest site (Roubik, 2006 ; Eltz et al., 2003 ; Suderajat et al., 2021 ). Suitable nest sites such as pre-formed tree cavities, active termite or ant nests, and underground cavities up to three meters deep have been documented for stingless bees (Grüter, 2020 ). Some species of stingless bees, especially species kept in meliponiculture farms, are more adaptable to man-made structures, such as cavities in wood walls, iron vessels, brick walls, electrical boxes, and water pipes as their nests (Suderajat et al., 2021 ). Besides nest boxes, the availability of resources within the foraging range and the physical characteristics of the site are also crucial for successful stingless bees keeping (Hubbell & Johnson, 1977 ; Eltz et al., 2003 ; Venturieri, 2009 ; Tornyie & Kwapong, 2005; Soro et al., 2019 ). The optimal temperature inside the nest is essential for the survival and reproduction of stingless bees (Engels et al., 1995 ). Due to their ectothermic nature, nest sites within the optimal temperature and humidity range provide suitable conditions for colony development and growth (Grüter, 2020 ). The ideal temperature range for stingless bees varies depending on the species and the region in which they live (Roubik, 2006 ). Due to their distribution in lowland tropical regions, most species thrive between 25–35°C (Engels et al., 1995 ). For example, H. itama – Malaysia's most common commercial stingless bee species - operates best at 27–32°C (Fahimee et al., 2024 ). At temperatures outside this range, the bees may struggle to regulate their body temperature and be unable to function properly. For example, low temperatures can cause the bees to become sluggish and less effective in foraging, while high temperatures can lead to dehydration and death (Souza-Junior et al., 2020 ; Zhao et al., 2021 ). Stingless bees use various strategies to maintain the temperature in the nest at their optimal range. The most common are modifying or insulating the physical characteristics of the nest space. For example, stingless bees can use wood, bitumen, cerumen, mud, and resin to insulate the nest (Roubik, 2006 ; Grüter, 2020 ; Shanahan & Spivak, 2021). They can also modify the size of the nest entrance, location, and amount of ventilation to control the in-and-out flow of air (Grüter, 2020 ). Besides that, stingless bees can also regulate nest temperature using brood arrangements. For example, packed cell layers in honeycomb-producing species can store heat better than the loose, insulated cells in cluster-producing species (Grüter, 2020 ). Other than the physical characteristics of the nests and brood arrangements, stingless bees, to some extent, can also behaviorally regulate temperature. For example, wing fanning can achieve ventilation, creating an airflow to reduce temperature (Roubik, 2006 ). Overheating in the brood area triggers accelerated and directed fanning by worker bees and a mass exodus of worker bees from the nest (Engels et al., 1995 ). However, these behaviors are costly, and worker bees engaging in thermoregulation are prevented from foraging. Hence, selecting nest cavities with minimal need for thermoregulation remains a top priority in nest cavity choice. Stingless bees continuously regulate the temperature within their nests throughout the day (Fletcher and Crewer, 1981; Teixeira et al., 2011; Vollet-Neto et al., 2011; Roldão-Sbordoni et al., 2019 ). This ongoing process is essential for brood development and colony survival. While they thermoregulate continuously, certain behaviors are more prevalent at specific times. Fanning and water collection are most common during the hottest part of the day, usually between 10:00 and 14:00 hours (Engles, 1995; Vollet-Neto, 2014). This is supported by research on Scaptotrigona depilis , where these behaviors increase as temperatures rise (Vollet-Neto, 2014). Bee activity in and out of the nests, related to nest temperature regulation, is heightened during the cooler morning hours (7:00–10:00 a.m.) when humidity is higher (Supandi et al., 2020). Within the nests, ‘hot’ nurse bees with elevated thorax temperatures help maintain brood warmth (Roldão-Sbordoni et al. 2019 ). At Universiti Malaysia Sabah's main campus, the commercial stingless bees, H. itama , nest mostly in lamp posts rather than natural tree cavities (Lardizabal and Juris, 2022). We aim to investigate the thermoregulation in lamp posts as nest sites for H. itama . Not all lamp posts were occupied. Hence, we were interested in whether occupied lamp posts were positioned where temperatures were mostly optimum for H. itama (27–32°C). To answer this first research question, we compare the ambient temperature in occupied-shade and unoccupied-exposed lamp posts regions and calculate the percentage of time these are outside of H. itama optimal temperature range. We expect occupied lamp posts to be concentrated in areas where temperatures are mostly optimal, minimizing the effort needed for nest temperature regulation. Secondly, we were interested in whether temperature regulation occurred in lamp post cavities and what time nest temperature regulation occurred. We compare the differential temperature between occupied and unoccupied lamp posts across four-time blocks to answer the second question. We expect most temperature regulation periods to occur when the temperature is outside the optimal temperature for the stingless bee, that is early morning or evening hours. Materials and methods Study sites The Universiti Malaysia Sabah (UMS) main campus covers an area of 4.04 km 2 . Before developing the UMS campus infrastructure in 1995, the area comprised a diverse mix of human settlements, agricultural land, and natural forest tracts. The current campus area features a variety of landscapes, including fragmented secondary forests and open canopy areas with bushes and grasses. The vegetation composition resembles lowland and mangrove forests, particularly in areas extending seaward (Majuakim et al., 2018). The topography is a mix of flat and undulating terrain, with hilly areas having steep slopes. The highest point, UMS Peak, reaches 190 m a.s.l. (Fig. 1 ). The UMS campus experiences an equatorial climate, with consistent temperatures and a high volume of rainfall. The average annual rainfall is 2,700 millimeters, and the mean annual temperature is 28°C (WorldWeatherOnline.com). Please insert Fig. 1 here Heterotrigona itama lamppost nest survey Lardizabal and Juris (2022) conducted a stingless bee diversity survey at the UMS campus in August 2020. They found that H. itama was the most abundant species, with most H. itama nesting inside lamp posts (49%), followed by wall crevices (38%), and a few nested in other substrates. Most nests inside lamp posts (92.6%) are H. itama . To confirm that this is still the case, we conducted H. itama nest survey in lamp posts along the roads of the UMS campus on October 2022. An active nest was evidenced by an entrance tube with bees flying in and out from the protruded entrance tube (Fig. 1 ). We recorded the GPS location of each nest. The Quantum Geographic Information System (QGIS) was utilized to map the distribution of stingless bee nests on lamp posts. To confirm the identification of the stingless bee species, we collected two to five foragers using 10 ml centrifuge tubes. The tubes were placed at a 30-degree angle in front of the nest entrances, effectively blocking the entrances to the target foraging bees. Selected bees are pinned, and the species identification was confirmed based on the stingless bees reference collection at BORNEENSIS, Universiti Malaysia Sabah (Supplementary Table 1). Are occupied lamp posts positioned in an ideal temperature location? The temperature logger measurements were carried out in mid-November 2022. A site with a row of occupied lamp posts (shaded by trees) and a row of unoccupied lamp posts (directly exposed to sunlight) were chosen for ambient temperature measurement (Fig. 1 ). The data logger measured the ambient temperature of both locations – one amid an occupied lamp post and the other amid an unoccupied lamp post (Onset Company; model U23-002, Onset external data recorder HOBO Pro V2 temp / RH). The data logger was deployed for 24 hours, recording 5- or 10-minute temperature readings. The hourly mean temperature was plotted on a line graph to visualize the temperature variations between the unoccupied and occupied lamp posts over the 24-hour period. The minimal (27°C) and maximal (32°C) temperature for H. itama (Fahimee et al., 2024 ) was added to the line graph for better visualization and calculation of the percentage of the day when temperatures exceeded the optimal range. All graphs and calculations were done in R software (version 4.3.3). Does temperature regulation occur in lamp post nests? We selected nine H. itama -occupied nests on the lamp posts to investigate whether and when temperature regulation occurred. The ambient and internal temperature for the nine selected H. itama lamp post nests were measured in late November 2022. Nine adjacent unoccupied lamp posts were selected as controls. The temperature was measured using an infrared thermometer (Fluke Corporation, model 566 IR thermometer; Everett WA) at four different time blocks of the day: block 1: 0700–0930 hr, block 2: 0930–1300 hr, block 3: 1300–1630 hr and block 4: 1630–2000 hr, to represent the entire activity range of H. itama . To determine if temperature regulation occurred, we used a two-way analysis of variance (ANOVA) with repeated measures. The two independent variables tested were (i) occupied versus unoccupied lamp posts and (ii) time blocks. The dependent variable was the differential temperature (ambient - inside). Our data did not meet the assumption of sphericity. Hence, we applied Greenhouse-Geisser correction (Bathke et al., 2009; Abdi, 2010) to our analyses. Two-way ANOVA was conducted using JASP software (version 0.16.4). A cluster bar chart was created to visualize temperature regulation across four time blocks. A positive value indicates ambient > inside temperature, and a negative value indicates inside > ambient temperature. Zero indicates ambient = inside temperature. Temperature regulation is observed when there is a difference in bar height or direction in differential temperature in occupied lamp posts compared to unoccupied adjacent lamp posts. Result Survey on H. itama nesting distribution on lamp posts. This study found that stingless bees occupied 82 lamp posts. H. itama inhabited approximately 64 lamp posts, making up 78% of all lamp post nests (Supplementary Table 1). The remaining nests belonged to other species: Tetragonula fuscobalteata (10 nests), Tetragonula laeviceps (7 nests), and Tetragonula geissleri (1 nest) (Supplementary Fig. 1). Are occupied lamp posts positioned in an ideal temperature location? The ambient temperature at occupied (shaded) and unoccupied (exposed) locations show a similar pattern in temperature change throughout the day, i.e. , a gradual rise in temperature from the early morning, peaking around midday, and then a gradual decline as the day progresses into the evening (Fig. 2 ). The unoccupied-exposed location reaches a higher peak temperature (33.38°C) than the occupied-shaded location (30.96°C). The occupied-shaded location remains consistently lower than the unoccupied-exposed lamp posts throughout the day (mean occupied-shaded = 25.89°C, mean unoccupied-exposed = 26.73°C). The temperatures in both locations are similar in the early morning and late evening. In this study, the temperature of the unoccupied-exposed lamp posts was outside the optimal range (27°C to 32°C) for 78.12% of the day, which translates to 18.75 hours. In contrast, occupied-shaded lamp posts experienced temperatures outside the optimal range for 68.4% of the day or 16.42 hours. Consequently, unoccupied-exposed lamp posts were within the optimal temperature range for only 5.25 hours, while occupied-shaded lamp posts enjoyed a longer duration of 7.58 hours within this range. This notable difference was particularly pronounced during the peak foraging activity period of 09:30 to 13:00 hours. Please insert Fig. 2 here Does temperature regulation occur in lamp post nests? There is no significant difference in temperatures between occupied and unoccupied lamp posts ( F = 2.709, p = 0.074). There is also no significant effect between the occupied and unoccupied lamp posts over the four-time blocks ( F = 0.829, p = 0.450) (Supplementary Table 2). In general, the inside temperature of the lamp posts was higher than the ambient temperature during the warmer parts of the day (Block 2 and 3: 0930–1630 hr) and closer to ambient temperature in the early morning (Block 1: 0700–0930 hr) and late afternoon to evening (Block 4: 1630–2000 hr), regardless of whether it is occupied or unoccupied. Temperature regulation (albeit not statistically significant) was detected over three of the four-time blocks. Temperature regulation was detected at Block 2 and 3: 0930–1630 hr, with − 0.017°C, indicating inside > ambient temperature between occupied and unoccupied lamp posts. Temperature regulation was detected again at Block 4: 1630–2000 hr, with − 0.889°C, indicating inside > ambient temperature between occupied and unoccupied lamp posts. Please insert Fig. 3 here Discussion Most stingless bees would prefer pre-existing cavities such as tree hollows, brick walls, and termite mounds, regardless of whether these are naturally occurring or man-made structures (Aidar et al., 2013; Choudharya et al., 2021; Hora et al., 2023). In our study, H. itama strongly preferred lamp posts as nesting materials at the Universiti Malaysia Sabah campus (Supplementary Fig. 1; Lardizabal & Juris, 2022). This preference likely stems from decreased suitable natural nesting sites, such as tree cavities in an increasingly urbanized campus environment. Urbanization and habitat alteration often reduce natural nesting sites, driving bees to adapt to artificial alternatives resembling preferred sites. For example, a re-evaluation of stingless bee species and abundance over eight years at the Federal University of Juiz de Fora campus in Minas Gerais, southeastern Brazil, recorded increased species richness and abundance in stingless bees to man-made nest sites. This is especially pronounced for Trigona spinipes , which recorded a switch to 100% occupancy in man-made nests (Vieira et al. 2016 ). Not all lamp posts at UMS were occupied with stingless bee nests. The lamp posts that were not occupied generally were situated in an exposed area ( personal observations ). Unoccupied-exposed lamp posts have only 5.25 hours per day that falls within the optimal foraging range of H. itama (27–32°C) (Fahimee et al., 2018 ). In contrast, occupied-shaded lamp posts have an additional 2.33 hours that falls within H. itama optimal foraging range. The periods where the temperature falls outside the optimal range occur during peak foraging hours (1000–1300 hr), supporting the significance of shaded areas for nest site choice (Hoffmann et al., 2016 , Ramos et al., 2023 ). Stingless bees, when overheated, can affect foraging performance and colony survivability (Lopez et al. 2011; Santos et al., 2018). Therefore, stingless bees will prioritize nest sites that ensure longest span of optimal temperatures for offspring development and colony survival (Silve et al., 2017). By selecting shaded lamp posts, H. itama can reduce overheating and the need for active thermoregulation, thus potentially conserving energy for other essential tasks like foraging and brood care (Jones & Oldroyd, 2007 ; Stabentheiner, 2021). We expected thermoregulation to occur when the nest temperature was outside the optimal range for H. itama . Our findings support this hypothesis with internal > ambient temperature from 0930 to 1630 (time blocks 2 to 3; Fig. 3 ). Putting aside the natural insulation of nesting materials, stingless bees are able to further cool their nests with certain behaviors. For example, some stingless bees increase water collection (Engels et al., 1995 ), while other species practice water regurgitation, storing water internally and releasing it inside the nest for additional cooling (Roubik, 2006 ). To further enhance the cooling process, the bees can use wing fanning to increase airflow and pushes hot air out of the nest (Jones and Oldroyd, 2007 ). We do not know which of these behaviors applies in our model system, but we expected behaviors with the least energy expenditure (increase water collection and water regurgitation) would be prioritize over wing fanning. The widespread utilization of lamp posts as nest sites suggests that the metal material can retain some residual heat from the day, radiating warmth into the nest. Coupled with passive thermoregulatory behaviors, such as huddling in tight clusters, reducing their exposed surface area to minimize heat loss and share body warmth (Engels et al., 1995 ; Jones and Oldroyd, 2007 ), occupied lamp post nests have internal > ambient temperature (time block 4: 1630–2000 hrs; Fig. 3 ). The temperature can also be increased with enhanced activity levels that generate metabolic heat that contributes to warming the nest (Roldão-Sbordoni et al., 2019 ). Moreover, stingless bees can reverse the direction of their fanning behavior. Instead of cooling the nest, this draws in warmer air from upper regions and circulates it around the brood (Jones and Oldroyd, 2007 ). All these combined actions maintain a suitable temperature for the optimal functioning of the colony. 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J Comp Physiol A 207(3):337–351 Suderajat A, Riyanto R, Mulawarman M (2021) The Types of Trigona Bee (Apidae: Meliponinae) in Three Different Habitat in South Sumatra. Jurnal Biologi Tropis 21(1):206–212 Supandi, Arkan F, Gusa RF, Jumnahdi M, Kurniawan R (2020) Design of system for setting the temperature and monitoring bees in and out the hive. In IOP Conference Series: Earth and Environmental Science (Vol. 599, No. 1, p. 012050). IOP Publishing Tornyie F, Kwapong PK (2015) Nesting ecology of stingless bees and potential threats to their survival within selected landscapes in the northern Volta region of Ghana. Afr J Ecol 53(4):398–405 Udy KL, Reininghaus H, Scherber C, Tscharntke T (2020) Plant–pollinator interactions along an urbanization gradient from cities and villages to farmland landscapes. Ecosphere, 11(2), e03020 Vieira KM, Netto P, Amaral DL, Mendes SS, Castro LC, Prezoto F (2016) Nesting stingless bees in urban areas: a reevaluation after eight years. Sociobiology 63(3):976–981 Venturieri GC (2009) The impact of forest exploitation on Amazonian stingless bees (Apidae, Meliponini). Genet Mol Res 8(2):684–689 Vollet-Neto A, Menezes C, Imperatriz-Fonseca VL (2015) Behavioural and developmental responses of a stingless bee ( Scaptotrigona depilis ) to nest overheating. Apidologie 46(4):455–464 Wilson ES, Murphy CE, Rinehart JP, Yocum G, Bowsher JH (2020) Microclimate temperatures impact nesting preference in Megachile rotundata (Hymenoptera: Megachilidae). Environ Entomol 49(2):296–303 World Weather Online https://www.worldweatheronline.com/kota-kinabalu-weather-average/sabah/my.aspx Zhao H, Li G, Guo D, Li H, Liu Q, Xu B, Guo X (2021) Response mechanisms to heat stress in bees. Apidologie 52:388–399 Supplementary Files SupplementaryFigure2.png SupplementaryTable1.docx SupplementaryTable2.docx Cite Share Download PDF Status: Published Journal Publication published 08 Nov, 2024 Read the published version in Insectes Sociaux → Version 1 posted Editorial decision: Major Revisions Needed 28 Aug, 2024 Reviewers agreed at journal 08 Aug, 2024 Reviewers invited by journal 04 Aug, 2024 Editor assigned by journal 04 Aug, 2024 First submitted to journal 01 Aug, 2024 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {\"props\":{\"pageProps\":{\"initialData\":{\"identity\":\"rs-4845014\",\"acceptedTermsAndConditions\":true,\"allowDirectSubmit\":false,\"archivedVersions\":[],\"articleType\":\"Short Report\",\"associatedPublications\":[],\"authors\":[{\"id\":335898434,\"identity\":\"304c6307-ce49-44aa-8a3d-2b2bdcbb8ad5\",\"order_by\":0,\"name\":\"Florina Anthony\",\"email\":\"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABAklEQVRIiWNgGAWjYBAC9nYgkQBhs4EIOSQ2dsBzGK6FGazMmDgtDEhaEhsIamFmPvbh4Q47Bv7Z/ccefNxxOH3DteMPGD6UHWbQnZGAQwtb8ozEM8kMEncOsxvOPHM4d8PtHAPGGecOM5jdwK7FnpnHmCGxjZmB4UYymzRvG1gLAzOQgVMLD0RLPYM8VEu6we30B8x/CWs5zGAA1ZJgcDvBgJkRrxa2ZKCW4zyGN5LNJGe2pRvOBPrlYM+5dB6zMw+wa2FvPsz4s61aTu5G4jOJj23W8ny30x8++FFmLWd2HLstcK1QuhlMHgCLCODXAgN1SGz+A0RpGQWjYBSMgmEPAHoRW85n+Ba/AAAAAElFTkSuQmCC\",\"orcid\":\"https://orcid.org/0000-0003-3045-2801\",\"institution\":\"Universiti Malaysia Sabah\",\"correspondingAuthor\":true,\"prefix\":\"\",\"firstName\":\"Florina\",\"middleName\":\"\",\"lastName\":\"Anthony\",\"suffix\":\"\"},{\"id\":335898435,\"identity\":\"06ef9dde-9502-4947-848c-363d413b009e\",\"order_by\":1,\"name\":\"Sze Huei Yek\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"Universiti Malaysia Sabah\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"Sze\",\"middleName\":\"Huei\",\"lastName\":\"Yek\",\"suffix\":\"\"}],\"badges\":[],\"createdAt\":\"2024-08-02 01:56:44\",\"currentVersionCode\":1,\"declarations\":\"\",\"doi\":\"10.21203/rs.3.rs-4845014/v1\",\"doiUrl\":\"https://doi.org/10.21203/rs.3.rs-4845014/v1\",\"draftVersion\":[],\"editorialEvents\":[{\"content\":\"https://doi.org/10.1007/s00040-024-01006-w\",\"type\":\"published\",\"date\":\"2024-11-08T15:56:59+00:00\"}],\"editorialNote\":\"\",\"failedWorkflow\":false,\"files\":[{\"id\":63643421,\"identity\":\"cde08423-c85a-43ac-827c-86898d14d177\",\"added_by\":\"auto\",\"created_at\":\"2024-08-30 13:25:17\",\"extension\":\"png\",\"order_by\":1,\"title\":\"Figure 1\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":1155001,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eStudy site map at Universiti Malaysia Sabah (UMS) campus, highlighted within the larger Sabah region, showing the locations of \\u003cem\\u003eHeterotrigona itama\\u003c/em\\u003e nests (purple circles) and temperature data loggers (yellow circles). Rectangular boxes indicate the placement of data loggers to record ambient and internal temperatures in both shaded and exposed lamp posts as part of temperature regulation experiments. Inset: Close-up view of an \\u003cem\\u003eH. itama\\u003c/em\\u003e nest entrance on a lamp post.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Figure1.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4845014/v1/44272a3641c46911a8dd5c3b.png\"},{\"id\":63643423,\"identity\":\"c5016879-01c8-45da-a5eb-c41687b0cec8\",\"added_by\":\"auto\",\"created_at\":\"2024-08-30 13:25:17\",\"extension\":\"png\",\"order_by\":2,\"title\":\"Figure 2\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":151960,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eThe ambient temperature in occupied-shaded and unoccupied-exposed lamp posts over 12 hours (November 15, 2022). The two red dotted vertical lines indicate the minimum (27°C) and maximum temperature (32°C) for \\u003cem\\u003eH. itama\\u003c/em\\u003e.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Figure2.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4845014/v1/2ee1c39dc04b8f85fae509a9.png\"},{\"id\":63643426,\"identity\":\"56d998cf-c45f-4ed0-ae50-d86aa5b150a8\",\"added_by\":\"auto\",\"created_at\":\"2024-08-30 13:25:18\",\"extension\":\"png\",\"order_by\":3,\"title\":\"Figure 3\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":183994,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eDifferential temperature (ambient - inside) for occupied versus unoccupied lamp posts across four distinct time blocks. Temperature regulation occurred over three of the four-time blocks. Temperature regulation was detected at Block 2 and 3: 0930 – 1630 hr, with -0.017°C, indicating inside \\u0026gt; ambient temperature between occupied and unoccupied lamp posts. Temperature regulation was detected again at Block 4: 1630 – 2000 hr, with -0.889°C, indicating inside \\u0026gt; ambient temperature between occupied and unoccupied lamp posts.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Figure3.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4845014/v1/79dccb508a185743afc0f488.png\"},{\"id\":68750199,\"identity\":\"2b385def-ae06-4694-b558-27232acaf473\",\"added_by\":\"auto\",\"created_at\":\"2024-11-11 16:11:33\",\"extension\":\"pdf\",\"order_by\":0,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"manuscript-pdf\",\"size\":2115483,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"manuscript.pdf\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4845014/v1/456c0bd6-f78f-420d-9204-733935712ad1.pdf\"},{\"id\":63643895,\"identity\":\"d5fcec2c-6e0d-4c8d-8cb0-8c3e36560321\",\"added_by\":\"auto\",\"created_at\":\"2024-08-30 13:33:18\",\"extension\":\"png\",\"order_by\":1,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"supplement\",\"size\":3735088,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"SupplementaryFigure2.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4845014/v1/c97261f7897ecdba9c7570cf.png\"},{\"id\":63643422,\"identity\":\"3b9856da-79ed-4e9c-ba3d-c0f40a0bccca\",\"added_by\":\"auto\",\"created_at\":\"2024-08-30 13:25:17\",\"extension\":\"docx\",\"order_by\":2,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"supplement\",\"size\":19643,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"SupplementaryTable1.docx\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4845014/v1/31a4b1d2a88ebbcaab94b6ba.docx\"},{\"id\":63643425,\"identity\":\"ebdb4097-0f51-4aeb-9526-8fc491879db5\",\"added_by\":\"auto\",\"created_at\":\"2024-08-30 13:25:17\",\"extension\":\"docx\",\"order_by\":3,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"supplement\",\"size\":15014,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"SupplementaryTable2.docx\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-4845014/v1/8db491c9cb73c64500fa4897.docx\"}],\"financialInterests\":\"\",\"formattedTitle\":\"Temperature regulation of Heterotrigona itama (Cockerell, 1918) in lamp posts nests\",\"fulltext\":[{\"header\":\"Introduction\",\"content\":\"\\u003cp\\u003eStingless bees, known as meliponines, are diverse bees in tropical and subtropical regions worldwide (Roubik, \\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e). These bees are important pollinators of a wide range of crops (Heard, \\u003cspan citationid=\\\"CR22\\\" class=\\\"CitationRef\\\"\\u003e1999\\u003c/span\\u003e; Amano et al., \\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e2000\\u003c/span\\u003e; Slaa et al., \\u003cspan citationid=\\\"CR50\\\" class=\\\"CitationRef\\\"\\u003e2006\\u003c/span\\u003e) and play a vital role in the health and functioning of ecosystems (Paz et al., 2021; Campbell et al., \\u003cspan citationid=\\\"CR8\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e). In addition to their ecological importance, stingless bees have a long history of use in meliponiculture and are valued for their honey production (Kwapong et al., \\u003cspan citationid=\\\"CR32\\\" class=\\\"CitationRef\\\"\\u003e2010\\u003c/span\\u003e). Malaysia has over 40 known species of stingless bees (Rasmussen, \\u003cspan citationid=\\\"CR45\\\" class=\\\"CitationRef\\\"\\u003e2008\\u003c/span\\u003e). Some common commercial species widely found in meliponiculture farms in Malaysia are \\u003cem\\u003eHeterotrigona itama\\u003c/em\\u003e, \\u003cem\\u003eGeniotrigona thoracica\\u003c/em\\u003e, and \\u003cem\\u003eLepidotrigona terminata\\u003c/em\\u003e (Msn et al., 2014; Mustafa et al., \\u003cspan citationid=\\\"CR41\\\" class=\\\"CitationRef\\\"\\u003e2018\\u003c/span\\u003e). These species occurred widely in urban areas and forest edges where meliponiculture farms were located (Hamid et al., \\u003cspan citationid=\\\"CR21\\\" class=\\\"CitationRef\\\"\\u003e2016\\u003c/span\\u003e). Due to their survivability in the human-adapted landscape, they are priced as honey producers and pollinators in Malaysia\\u0026rsquo;s meliponiculture industries (Kelly et al., \\u003cspan citationid=\\\"CR27\\\" class=\\\"CitationRef\\\"\\u003e2014\\u003c/span\\u003e; Fahimee et al., \\u003cspan citationid=\\\"CR15\\\" class=\\\"CitationRef\\\"\\u003e2016\\u003c/span\\u003e,\\u003cspan citationid=\\\"CR16\\\" class=\\\"CitationRef\\\"\\u003e2018\\u003c/span\\u003e; Ghazi et al., \\u003cspan citationid=\\\"CR19\\\" class=\\\"CitationRef\\\"\\u003e2018\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003eA critical factor determining a stingless bee colony\\u0026rsquo;s success is selecting a suitable nest site (Roubik, \\u003cspan citationid=\\\"CR47\\\" class=\\\"CitationRef\\\"\\u003e2006\\u003c/span\\u003e; Eltz et al., \\u003cspan citationid=\\\"CR12\\\" class=\\\"CitationRef\\\"\\u003e2003\\u003c/span\\u003e; Suderajat et al., \\u003cspan citationid=\\\"CR55\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e). Suitable nest sites such as pre-formed tree cavities, active termite or ant nests, and underground cavities up to three meters deep have been documented for stingless bees (Gr\\u0026uuml;ter, \\u003cspan citationid=\\\"CR20\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). Some species of stingless bees, especially species kept in meliponiculture farms, are more adaptable to man-made structures, such as cavities in wood walls, iron vessels, brick walls, electrical boxes, and water pipes as their nests (Suderajat et al., \\u003cspan citationid=\\\"CR55\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e). Besides nest boxes, the availability of resources within the foraging range and the physical characteristics of the site are also crucial for successful stingless bees keeping (Hubbell \\u0026amp; Johnson, \\u003cspan citationid=\\\"CR25\\\" class=\\\"CitationRef\\\"\\u003e1977\\u003c/span\\u003e; Eltz et al., \\u003cspan citationid=\\\"CR12\\\" class=\\\"CitationRef\\\"\\u003e2003\\u003c/span\\u003e; Venturieri, \\u003cspan citationid=\\\"CR60\\\" class=\\\"CitationRef\\\"\\u003e2009\\u003c/span\\u003e; Tornyie \\u0026amp; Kwapong, 2005; Soro et al., \\u003cspan citationid=\\\"CR52\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003eThe optimal temperature inside the nest is essential for the survival and reproduction of stingless bees (Engels et al., \\u003cspan citationid=\\\"CR13\\\" class=\\\"CitationRef\\\"\\u003e1995\\u003c/span\\u003e). Due to their ectothermic nature, nest sites within the optimal temperature and humidity range provide suitable conditions for colony development and growth (Gr\\u0026uuml;ter, \\u003cspan citationid=\\\"CR20\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). The ideal temperature range for stingless bees varies depending on the species and the region in which they live (Roubik, \\u003cspan citationid=\\\"CR47\\\" class=\\\"CitationRef\\\"\\u003e2006\\u003c/span\\u003e). Due to their distribution in lowland tropical regions, most species thrive between 25\\u0026ndash;35\\u0026deg;C (Engels et al., \\u003cspan citationid=\\\"CR13\\\" class=\\\"CitationRef\\\"\\u003e1995\\u003c/span\\u003e). For example, \\u003cem\\u003eH. itama\\u003c/em\\u003e \\u0026ndash; Malaysia's most common commercial stingless bee species - operates best at 27\\u0026ndash;32\\u0026deg;C (Fahimee et al., \\u003cspan citationid=\\\"CR17\\\" class=\\\"CitationRef\\\"\\u003e2024\\u003c/span\\u003e). At temperatures outside this range, the bees may struggle to regulate their body temperature and be unable to function properly. For example, low temperatures can cause the bees to become sluggish and less effective in foraging, while high temperatures can lead to dehydration and death (Souza-Junior et al., \\u003cspan citationid=\\\"CR53\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e; Zhao et al., \\u003cspan citationid=\\\"CR64\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003eStingless bees use various strategies to maintain the temperature in the nest at their optimal range. The most common are modifying or insulating the physical characteristics of the nest space. For example, stingless bees can use wood, bitumen, cerumen, mud, and resin to insulate the nest (Roubik, \\u003cspan citationid=\\\"CR47\\\" class=\\\"CitationRef\\\"\\u003e2006\\u003c/span\\u003e; Gr\\u0026uuml;ter, \\u003cspan citationid=\\\"CR20\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e; Shanahan \\u0026amp; Spivak, 2021). They can also modify the size of the nest entrance, location, and amount of ventilation to control the in-and-out flow of air (Gr\\u0026uuml;ter, \\u003cspan citationid=\\\"CR20\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). Besides that, stingless bees can also regulate nest temperature using brood arrangements. For example, packed cell layers in honeycomb-producing species can store heat better than the loose, insulated cells in cluster-producing species (Gr\\u0026uuml;ter, \\u003cspan citationid=\\\"CR20\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003eOther than the physical characteristics of the nests and brood arrangements, stingless bees, to some extent, can also behaviorally regulate temperature. For example, wing fanning can achieve ventilation, creating an airflow to reduce temperature (Roubik, \\u003cspan citationid=\\\"CR47\\\" class=\\\"CitationRef\\\"\\u003e2006\\u003c/span\\u003e). Overheating in the brood area triggers accelerated and directed fanning by worker bees and a mass exodus of worker bees from the nest (Engels et al., \\u003cspan citationid=\\\"CR13\\\" class=\\\"CitationRef\\\"\\u003e1995\\u003c/span\\u003e). However, these behaviors are costly, and worker bees engaging in thermoregulation are prevented from foraging. Hence, selecting nest cavities with minimal need for thermoregulation remains a top priority in nest cavity choice.\\u003c/p\\u003e \\u003cp\\u003eStingless bees continuously regulate the temperature within their nests throughout the day (Fletcher and Crewer, 1981; Teixeira et al., 2011; Vollet-Neto et al., 2011; Rold\\u0026atilde;o-Sbordoni et al., \\u003cspan citationid=\\\"CR46\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e). This ongoing process is essential for brood development and colony survival. While they thermoregulate continuously, certain behaviors are more prevalent at specific times. Fanning and water collection are most common during the hottest part of the day, usually between 10:00 and 14:00 hours (Engles, 1995; Vollet-Neto, 2014). This is supported by research on \\u003cem\\u003eScaptotrigona depilis\\u003c/em\\u003e, where these behaviors increase as temperatures rise (Vollet-Neto, 2014). Bee activity in and out of the nests, related to nest temperature regulation, is heightened during the cooler morning hours (7:00\\u0026ndash;10:00 a.m.) when humidity is higher (Supandi et al., 2020). Within the nests, \\u0026lsquo;hot\\u0026rsquo; nurse bees with elevated thorax temperatures help maintain brood warmth (Rold\\u0026atilde;o-Sbordoni et al. \\u003cspan citationid=\\\"CR46\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003eAt Universiti Malaysia Sabah's main campus, the commercial stingless bees, \\u003cem\\u003eH. itama\\u003c/em\\u003e, nest mostly in lamp posts rather than natural tree cavities (Lardizabal and Juris, 2022). We aim to investigate the thermoregulation in lamp posts as nest sites for \\u003cem\\u003eH. itama\\u003c/em\\u003e. Not all lamp posts were occupied. Hence, we were interested in whether occupied lamp posts were positioned where temperatures were mostly optimum for \\u003cem\\u003eH. itama\\u003c/em\\u003e (27\\u0026ndash;32\\u0026deg;C). To answer this first research question, we compare the ambient temperature in occupied-shade and unoccupied-exposed lamp posts regions and calculate the percentage of time these are outside of \\u003cem\\u003eH. itama\\u003c/em\\u003e optimal temperature range. We expect occupied lamp posts to be concentrated in areas where temperatures are mostly optimal, minimizing the effort needed for nest temperature regulation. Secondly, we were interested in whether temperature regulation occurred in lamp post cavities and what time nest temperature regulation occurred. We compare the differential temperature between occupied and unoccupied lamp posts across four-time blocks to answer the second question. We expect most temperature regulation periods to occur when the temperature is outside the optimal temperature for the stingless bee, that is early morning or evening hours.\\u003c/p\\u003e\"},{\"header\":\"Materials and methods\",\"content\":\"\\u003cdiv id=\\\"Sec3\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eStudy sites\\u003c/h2\\u003e \\u003cp\\u003eThe Universiti Malaysia Sabah (UMS) main campus covers an area of 4.04 km\\u003csup\\u003e2\\u003c/sup\\u003e. Before developing the UMS campus infrastructure in 1995, the area comprised a diverse mix of human settlements, agricultural land, and natural forest tracts. The current campus area features a variety of landscapes, including fragmented secondary forests and open canopy areas with bushes and grasses. The vegetation composition resembles lowland and mangrove forests, particularly in areas extending seaward (Majuakim et al., 2018). The topography is a mix of flat and undulating terrain, with hilly areas having steep slopes. The highest point, UMS Peak, reaches 190 m a.s.l. (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e). The UMS campus experiences an equatorial climate, with consistent temperatures and a high volume of rainfall. The average annual rainfall is 2,700 millimeters, and the mean annual temperature is 28\\u0026deg;C (WorldWeatherOnline.com).\\u003c/p\\u003e \\u003cp\\u003e \\u003c/p\\u003e \\u003cp\\u003e \\u003cem\\u003ePlease insert\\u003c/em\\u003e Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e \\u003cem\\u003ehere\\u003c/em\\u003e\\u003c/p\\u003e \\u003cp\\u003e \\u003cb\\u003eHeterotrigona itama\\u003c/b\\u003e \\u003cb\\u003elamppost nest survey\\u003c/b\\u003e\\u003c/p\\u003e \\u003cp\\u003eLardizabal and Juris (2022) conducted a stingless bee diversity survey at the UMS campus in August 2020. They found that \\u003cem\\u003eH. itama\\u003c/em\\u003e was the most abundant species, with most \\u003cem\\u003eH. itama\\u003c/em\\u003e nesting inside lamp posts (49%), followed by wall crevices (38%), and a few nested in other substrates. Most nests inside lamp posts (92.6%) are \\u003cem\\u003eH. itama\\u003c/em\\u003e. To confirm that this is still the case, we conducted \\u003cem\\u003eH. itama\\u003c/em\\u003e nest survey in lamp posts along the roads of the UMS campus on October 2022. An active nest was evidenced by an entrance tube with bees flying in and out from the protruded entrance tube (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e). We recorded the GPS location of each nest. The Quantum Geographic Information System (QGIS) was utilized to map the distribution of stingless bee nests on lamp posts. To confirm the identification of the stingless bee species, we collected two to five foragers using 10 ml centrifuge tubes. The tubes were placed at a 30-degree angle in front of the nest entrances, effectively blocking the entrances to the target foraging bees. Selected bees are pinned, and the species identification was confirmed based on the stingless bees reference collection at BORNEENSIS, Universiti Malaysia Sabah (Supplementary Table\\u0026nbsp;1).\\u003c/p\\u003e \\u003c/div\\u003e \\u003cdiv id=\\\"Sec4\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eAre occupied lamp posts positioned in an ideal temperature location?\\u003c/h2\\u003e \\u003cp\\u003eThe temperature logger measurements were carried out in mid-November 2022. A site with a row of occupied lamp posts (shaded by trees) and a row of unoccupied lamp posts (directly exposed to sunlight) were chosen for ambient temperature measurement (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e). The data logger measured the ambient temperature of both locations \\u0026ndash; one amid an occupied lamp post and the other amid an unoccupied lamp post (Onset Company; model U23-002, Onset external data recorder HOBO Pro V2 temp / RH). The data logger was deployed for 24 hours, recording 5- or 10-minute temperature readings.\\u003c/p\\u003e \\u003cp\\u003eThe hourly mean temperature was plotted on a line graph to visualize the temperature variations between the unoccupied and occupied lamp posts over the 24-hour period. The minimal (27\\u0026deg;C) and maximal (32\\u0026deg;C) temperature for \\u003cem\\u003eH. itama\\u003c/em\\u003e (Fahimee et al., \\u003cspan citationid=\\\"CR17\\\" class=\\\"CitationRef\\\"\\u003e2024\\u003c/span\\u003e) was added to the line graph for better visualization and calculation of the percentage of the day when temperatures exceeded the optimal range. All graphs and calculations were done in R software (version 4.3.3).\\u003c/p\\u003e \\u003c/div\\u003e \\u003cdiv id=\\\"Sec5\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eDoes temperature regulation occur in lamp post nests?\\u003c/h2\\u003e \\u003cp\\u003eWe selected nine \\u003cem\\u003eH. itama\\u003c/em\\u003e-occupied nests on the lamp posts to investigate whether and when temperature regulation occurred. The ambient and internal temperature for the nine selected \\u003cem\\u003eH. itama\\u003c/em\\u003e lamp post nests were measured in late November 2022. Nine adjacent unoccupied lamp posts were selected as controls. The temperature was measured using an infrared thermometer (Fluke Corporation, model 566 IR thermometer; Everett WA) at four different time blocks of the day: block 1: 0700\\u0026ndash;0930 hr, block 2: 0930\\u0026ndash;1300 hr, block 3: 1300\\u0026ndash;1630 hr and block 4: 1630\\u0026ndash;2000 hr, to represent the entire activity range of \\u003cem\\u003eH. itama\\u003c/em\\u003e.\\u003c/p\\u003e \\u003cp\\u003eTo determine if temperature regulation occurred, we used a two-way analysis of variance (ANOVA) with repeated measures. The two independent variables tested were (i) occupied \\u003cem\\u003eversus\\u003c/em\\u003e unoccupied lamp posts and (ii) time blocks. The dependent variable was the differential temperature (ambient - inside). Our data did not meet the assumption of sphericity. Hence, we applied Greenhouse-Geisser correction (Bathke et al., 2009; Abdi, 2010) to our analyses. Two-way ANOVA was conducted using JASP software (version 0.16.4). A cluster bar chart was created to visualize temperature regulation across four time blocks. A positive value indicates ambient\\u0026thinsp;\\u0026gt;\\u0026thinsp;inside temperature, and a negative value indicates inside \\u0026gt;\\u0026thinsp;ambient temperature. Zero indicates ambient\\u0026thinsp;=\\u0026thinsp;inside temperature. Temperature regulation is observed when there is a difference in bar height or direction in differential temperature in occupied lamp posts compared to unoccupied adjacent lamp posts.\\u003c/p\\u003e \\u003c/div\\u003e\"},{\"header\":\"Result\",\"content\":\"\\u003cp\\u003e \\u003cb\\u003eSurvey on H. itama nesting distribution on lamp posts.\\u003c/b\\u003e \\u003c/p\\u003e \\u003cp\\u003eThis study found that stingless bees occupied 82 lamp posts. \\u003cem\\u003eH. itama\\u003c/em\\u003e inhabited approximately 64 lamp posts, making up 78% of all lamp post nests (Supplementary Table\\u0026nbsp;1). The remaining nests belonged to other species: \\u003cem\\u003eTetragonula fuscobalteata\\u003c/em\\u003e (10 nests), \\u003cem\\u003eTetragonula laeviceps\\u003c/em\\u003e (7 nests), and \\u003cem\\u003eTetragonula geissleri\\u003c/em\\u003e (1 nest) (Supplementary Fig.\\u0026nbsp;1).\\u003c/p\\u003e \\u003cdiv id=\\\"Sec7\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eAre occupied lamp posts positioned in an ideal temperature location?\\u003c/h2\\u003e \\u003cp\\u003eThe ambient temperature at occupied (shaded) and unoccupied (exposed) locations show a similar pattern in temperature change throughout the day, \\u003cem\\u003ei.e.\\u003c/em\\u003e, a gradual rise in temperature from the early morning, peaking around midday, and then a gradual decline as the day progresses into the evening (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003e). The unoccupied-exposed location reaches a higher peak temperature (33.38\\u0026deg;C) than the occupied-shaded location (30.96\\u0026deg;C). The occupied-shaded location remains consistently lower than the unoccupied-exposed lamp posts throughout the day (mean occupied-shaded\\u0026thinsp;=\\u0026thinsp;25.89\\u0026deg;C, mean unoccupied-exposed\\u0026thinsp;=\\u0026thinsp;26.73\\u0026deg;C). The temperatures in both locations are similar in the early morning and late evening. In this study, the temperature of the unoccupied-exposed lamp posts was outside the optimal range (27\\u0026deg;C to 32\\u0026deg;C) for 78.12% of the day, which translates to 18.75 hours. In contrast, occupied-shaded lamp posts experienced temperatures outside the optimal range for 68.4% of the day or 16.42 hours. Consequently, unoccupied-exposed lamp posts were within the optimal temperature range for only 5.25 hours, while occupied-shaded lamp posts enjoyed a longer duration of 7.58 hours within this range. This notable difference was particularly pronounced during the peak foraging activity period of 09:30 to 13:00 hours.\\u003c/p\\u003e \\u003cp\\u003e \\u003c/p\\u003e \\u003cp\\u003e \\u003cem\\u003ePlease insert\\u003c/em\\u003e Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003e \\u003cem\\u003ehere\\u003c/em\\u003e\\u003c/p\\u003e \\u003c/div\\u003e \\u003cdiv id=\\\"Sec8\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eDoes temperature regulation occur in lamp post nests?\\u003c/h2\\u003e \\u003cp\\u003eThere is no significant difference in temperatures between occupied and unoccupied lamp posts (\\u003cem\\u003eF\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;2.709, \\u003cem\\u003ep\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.074). There is also no significant effect between the occupied and unoccupied lamp posts over the four-time blocks (\\u003cem\\u003eF\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.829, \\u003cem\\u003ep\\u003c/em\\u003e\\u0026thinsp;=\\u0026thinsp;0.450) (Supplementary Table\\u0026nbsp;2).\\u003c/p\\u003e \\u003cp\\u003eIn general, the inside temperature of the lamp posts was higher than the ambient temperature during the warmer parts of the day (Block 2 and 3: 0930\\u0026ndash;1630 hr) and closer to ambient temperature in the early morning (Block 1: 0700\\u0026ndash;0930 hr) and late afternoon to evening (Block 4: 1630\\u0026ndash;2000 hr), regardless of whether it is occupied or unoccupied. Temperature regulation (albeit not statistically significant) was detected over three of the four-time blocks. Temperature regulation was detected at Block 2 and 3: 0930\\u0026ndash;1630 hr, with \\u0026minus;\\u0026thinsp;0.017\\u0026deg;C, indicating inside \\u0026gt;\\u0026thinsp;ambient temperature between occupied and unoccupied lamp posts. Temperature regulation was detected again at Block 4: 1630\\u0026ndash;2000 hr, with \\u0026minus;\\u0026thinsp;0.889\\u0026deg;C, indicating inside \\u0026gt;\\u0026thinsp;ambient temperature between occupied and unoccupied lamp posts.\\u003c/p\\u003e \\u003cp\\u003e \\u003cem\\u003ePlease insert\\u003c/em\\u003e Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig3\\\" class=\\\"InternalRef\\\"\\u003e3\\u003c/span\\u003e \\u003cem\\u003ehere\\u003c/em\\u003e\\u003c/p\\u003e \\u003cp\\u003e \\u003c/p\\u003e \\u003c/div\\u003e\"},{\"header\":\"Discussion\",\"content\":\"\\u003cp\\u003eMost stingless bees would prefer pre-existing cavities such as tree hollows, brick walls, and termite mounds, regardless of whether these are naturally occurring or man-made structures (Aidar et al., 2013; Choudharya et al., 2021; Hora et al., 2023). In our study, \\u003cem\\u003eH. itama\\u003c/em\\u003e strongly preferred lamp posts as nesting materials at the Universiti Malaysia Sabah campus (Supplementary Fig.\\u0026nbsp;1; Lardizabal \\u0026amp; Juris, 2022). This preference likely stems from decreased suitable natural nesting sites, such as tree cavities in an increasingly urbanized campus environment. Urbanization and habitat alteration often reduce natural nesting sites, driving bees to adapt to artificial alternatives resembling preferred sites. For example, a re-evaluation of stingless bee species and abundance over eight years at the Federal University of Juiz de Fora campus in Minas Gerais, southeastern Brazil, recorded increased species richness and abundance in stingless bees to man-made nest sites. This is especially pronounced for \\u003cem\\u003eTrigona spinipes\\u003c/em\\u003e, which recorded a switch to 100% occupancy in man-made nests (Vieira et al. \\u003cspan citationid=\\\"CR59\\\" class=\\\"CitationRef\\\"\\u003e2016\\u003c/span\\u003e).\\u003c/p\\u003e \\u003cp\\u003eNot all lamp posts at UMS were occupied with stingless bee nests. The lamp posts that were not occupied generally were situated in an exposed area (\\u003cem\\u003epersonal observations\\u003c/em\\u003e). Unoccupied-exposed lamp posts have only 5.25 hours per day that falls within the optimal foraging range of \\u003cem\\u003eH. itama\\u003c/em\\u003e (27\\u0026ndash;32\\u0026deg;C) (Fahimee et al., \\u003cspan citationid=\\\"CR16\\\" class=\\\"CitationRef\\\"\\u003e2018\\u003c/span\\u003e). In contrast, occupied-shaded lamp posts have an additional 2.33 hours that falls within \\u003cem\\u003eH. itama\\u003c/em\\u003e optimal foraging range. The periods where the temperature falls outside the optimal range occur during peak foraging hours (1000\\u0026ndash;1300 hr), supporting the significance of shaded areas for nest site choice (Hoffmann et al., \\u003cspan citationid=\\\"CR24\\\" class=\\\"CitationRef\\\"\\u003e2016\\u003c/span\\u003e, Ramos et al., \\u003cspan citationid=\\\"CR44\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e). Stingless bees, when overheated, can affect foraging performance and colony survivability (Lopez et al. 2011; Santos et al., 2018). Therefore, stingless bees will prioritize nest sites that ensure longest span of optimal temperatures for offspring development and colony survival (Silve et al., 2017). By selecting shaded lamp posts, \\u003cem\\u003eH. itama\\u003c/em\\u003e can reduce overheating and the need for active thermoregulation, thus potentially conserving energy for other essential tasks like foraging and brood care (Jones \\u0026amp; Oldroyd, \\u003cspan citationid=\\\"CR26\\\" class=\\\"CitationRef\\\"\\u003e2007\\u003c/span\\u003e; Stabentheiner, 2021).\\u003c/p\\u003e \\u003cp\\u003eWe expected thermoregulation to occur when the nest temperature was outside the optimal range for \\u003cem\\u003eH. itama\\u003c/em\\u003e. Our findings support this hypothesis with internal\\u0026thinsp;\\u0026gt;\\u0026thinsp;ambient temperature from 0930 to 1630 (time blocks 2 to 3; Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig3\\\" class=\\\"InternalRef\\\"\\u003e3\\u003c/span\\u003e). Putting aside the natural insulation of nesting materials, stingless bees are able to further cool their nests with certain behaviors. For example, some stingless bees increase water collection (Engels et al., \\u003cspan citationid=\\\"CR13\\\" class=\\\"CitationRef\\\"\\u003e1995\\u003c/span\\u003e), while other species practice water regurgitation, storing water internally and releasing it inside the nest for additional cooling (Roubik, \\u003cspan citationid=\\\"CR47\\\" class=\\\"CitationRef\\\"\\u003e2006\\u003c/span\\u003e). To further enhance the cooling process, the bees can use wing fanning to increase airflow and pushes hot air out of the nest (Jones and Oldroyd, \\u003cspan citationid=\\\"CR26\\\" class=\\\"CitationRef\\\"\\u003e2007\\u003c/span\\u003e). We do not know which of these behaviors applies in our model system, but we expected behaviors with the least energy expenditure (increase water collection and water regurgitation) would be prioritize over wing fanning.\\u003c/p\\u003e \\u003cp\\u003eThe widespread utilization of lamp posts as nest sites suggests that the metal material can retain some residual heat from the day, radiating warmth into the nest. Coupled with passive thermoregulatory behaviors, such as huddling in tight clusters, reducing their exposed surface area to minimize heat loss and share body warmth (Engels et al., \\u003cspan citationid=\\\"CR13\\\" class=\\\"CitationRef\\\"\\u003e1995\\u003c/span\\u003e; Jones and Oldroyd, \\u003cspan citationid=\\\"CR26\\\" class=\\\"CitationRef\\\"\\u003e2007\\u003c/span\\u003e), occupied lamp post nests have internal\\u0026thinsp;\\u0026gt;\\u0026thinsp;ambient temperature (time block 4: 1630\\u0026ndash;2000 hrs; Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig3\\\" class=\\\"InternalRef\\\"\\u003e3\\u003c/span\\u003e). The temperature can also be increased with enhanced activity levels that generate metabolic heat that contributes to warming the nest (Rold\\u0026atilde;o-Sbordoni et al., \\u003cspan citationid=\\\"CR46\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e). Moreover, stingless bees can reverse the direction of their fanning behavior. Instead of cooling the nest, this draws in warmer air from upper regions and circulates it around the brood (Jones and Oldroyd, \\u003cspan citationid=\\\"CR26\\\" class=\\\"CitationRef\\\"\\u003e2007\\u003c/span\\u003e). All these combined actions maintain a suitable temperature for the optimal functioning of the colony. Future investigations should include temperature readings and behavioral observations of \\u003cem\\u003eH. itama\\u003c/em\\u003e, especially during extreme temperatures. Understanding the adaptability of stingless bees in increasingly urbanized environment is crucial for pollinator conservation and meliponiculture industry improvement.\\u003c/p\\u003e\"},{\"header\":\"Declarations\",\"content\":\"\\u003ch2\\u003eAcknowledgment\\u003c/h2\\u003e \\u003cp\\u003eWe would like to thank Alvinus Joseph, Ling Chi Wong, and a dedicated group of volunteers for their assistance with data collection. The present study complied with the Sabah Biodiversity Protection laws (Access License: JKM/MBS.1000-2/2 JLD.16(18)) and was financially supported by a Swiss National Science Foundation grant (SPIRIT: IZSTZ0_189496) to SHY (UMS Grant Code: LPA2201).\\u003c/p\\u003e\"},{\"header\":\"References\",\"content\":\"\\u003col\\u003e\\u003cli\\u003e\\u003cspan\\u003eAmano K, Nemoto T, Heard TA (2000) What are stingless bees, and why and how to use them as crop pollinators? \\u0026ndash; a review. JARQ 34:183\\u0026ndash;190\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eAntoine CM, Forrest JR (2021) Nesting habitat of ground-nesting bees: a review. Ecol Entomol 46(2):143\\u0026ndash;159\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eAra\\u0026uacute;jo ED, Costa M, Chaud-Netto J, Fowler HG (2004) Body size and flight distance in stingless bees (Hymenoptera: Meliponini): inference of flight range and possible ecological implications. Brazilian J Biology 64:563\\u0026ndash;568\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eBarbieri C, Pinheiro GL, Drago PM, Francoy TM (2019) A scientific note on a stingless beehive model for ecological and behavioral studies and for environmental education. Sociobiology 66(1):186\\u0026ndash;189\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eBasari N, Ramli SN, Mohd Khairi NAS (2018) Food reward and distance influence the foraging pattern of stingless bee, \\u003cem\\u003eHeterotrigona itama\\u003c/em\\u003e. Insects 9(4):138\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eBenedick S, Gansau JA, Ahmad AH (2021) Foraging Behaviour of Heterotrigona itama (Apidae: Meliponini) in Residential Areas. Pertanika J Trop Agric Sci, 44(2)\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eBrown JC, Albrecht C (2001) The effect of tropical deforestation on stingless bees of the genus Melipona (Insecta: Hymenoptera: Apidae: Meliponini) in central Rondonia, Brazil. J Biogeogr 28(5):623\\u0026ndash;634\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eCampbell AJ, Lichtenberg EM, Carvalheiro LG, Menezes C, Borges RC, Coelho BWT, Mau\\u0026eacute;s MM (2022) High bee functional diversity buffers crop pollination services against Amazon deforestation, vol 326. Agriculture, Ecosystems \\u0026amp; Environment, p 107777\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eCorregidor-Castro A, Jones OR (2021) The effect of nest temperature on growth and survival in juvenile Great Tits Parus major. Ecol Evol 11(12):7346\\u0026ndash;7353\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eda Silva MA, Ferreira NDS, Teixeira-Souza VHDS, Maia-Silva C, de Oliveira FDA, Hrncir M (2017) On the thermal limits for the use of stingless bees as pollinators in commercial greenhouses. J Apic Res 56(1):81\\u0026ndash;90\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eDantas MRT (2020) Thermogenesis in stingless bees: an approach with emphasis on brood's thermal contribution. J Anim Behav Biometeorol 4(4):101\\u0026ndash;108\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eEltz T, Br\\u0026uuml;hl CA, Imiyabir Z, Linsenmair KE (2003) Nesting and nest trees of stingless bees (Apidae: Meliponini) in lowland dipterocarp forests in Sabah, Malaysia, with implications for forest management. For Ecol Manag 172(2\\u0026ndash;3):301\\u0026ndash;313\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eEngels W, Rosenkranz P, Engels E (1995) Thermoregulation in the nest of the Neotropical stingless bee Scaptotrigona postica and a hypothesis on the evolution of temperature homeostasis in highly eusocial bees. Stud Neotropical fauna Environ 30(4):193\\u0026ndash;205\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eFaheem M, Aslam M, Razaq M (2004) Pollination ecology with special reference to insect a review. JRes (Sci) 15:395\\u0026ndash;409\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eFahimee J, Halim M, Mispan MR, Jajuli R, Saranum MM, Zainuddin MY, Ghani A, I (2016) The diversity and abundance of stingless bee (Hymenoptera: Meliponini) in Peninsular Malaysia. Adv Environ Biology 10(9):1\\u0026ndash;8\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eFahimee J, Jajuli R, Mispan MR, Ghani IA (2018) Foraging behavior of stingless bee Heterotrigona itama (Cockerell, 1918) (Hymenoptera: Apidae: Meliponini). In AIP Conference proceedings (Vol. 1940, No. 1). AIP Publishing\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eFahimee J, Zulidzham MS, Reward NF, Muzammil N, Jajuli R, Abdullah M, Yaakop S (2024) Development of Heterotrigona itama (Cockerell, 1918) queens by in vitro culture for conservation purposes. J Apic Res 63(1):153\\u0026ndash;161\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eFierro MM, Cruz-Lopez L, Sanchez D, Villanueva-Gutierrez R, Vandame R (2012) Effect of biotic factors on the spatial distribution of stingless bees (Hymenoptera: Apidae, Meliponini) in fragmented neotropical habitats. Neotrop Entomol 41(2):95\\u0026ndash;104\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eGhazi R, Zulqurnain NS, Azmi WA (2018) Melittopalynological studies of stingless bees from the east coast of peninsular Malaysia. Pot-pollen stingless bee melittology, 77\\u0026ndash;88\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eGr\\u0026uuml;ter C, Gr\\u0026uuml;ter C (2020) Stingless bees: an overview. Stingless Bees: Their Behaviour, Ecology and Evolution, 1\\u0026ndash;42\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eHamid SA, Salleh MS, Thevan K, Hashim NA (2016) Distribution and morphometrical variations of stingless bees (Apidae: Meliponini) in urban and forest areas of Penang Island, Malaysia. J Trop Resour Sustain Sci 4:1\\u0026ndash;5\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eHeard TA (1999) The role of stingless bees in crop pollination. Ann Rev Entomol 44(1):183\\u0026ndash;206\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eHe M, Ran N, Jiang H, Han Z, Dian Y, Li X, Wang H (2022) Effects of landscape and local factors on the diversity of flower-visitor groups under an urbanization gradient, a case study in Wuhan, China. Diversity 14(3):208\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eHoffmann S, Lee ES, McNeil A, Fernandes L, Vidanovic D, Thanachareonkit A (2016) Balancing daylight, glare, and energy-efficiency goals: An evaluation of exterior coplanar shading systems using complex fenestration modeling tools. Energy Build 112:279\\u0026ndash;298\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eHubbell SP, Johnson LK (1977) Competition and nest spacing in a tropical stingless bee community. Ecology 58(5):949\\u0026ndash;963\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eJones JC, Oldroyd BP (2007) Nest thermoregulation in social insects. Adv insect Physiol 33:153\\u0026ndash;191\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eKelly N, Farisya MSN, Kumara TK, Marcela P (2014) Species diversity and external nest characteristics of stingless bees in meliponiculture. 293\\u0026ndash;298\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eKhan DH, Ali M, Khan FZ, Mehmood MA, Saeed S (2024) Effect of landscape complexity, nesting substrate, and nest orientation on cavity-nesting solitary bees in southern Punjab, Pakistan. Int J Trop Insect Sci, 1\\u0026ndash;11\\u003c/span\\u003e\\u003c/li\\u003e \\u003cli\\u003e\\u003cspan\\u003eKiatoko N, Van Langevelde F, Raina SK (2018) Forest degradation influences nesting site selection of Afro-tropical stingless bee species in a tropical rain forest, Kenya. 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Apidologie 52:388\\u0026ndash;399\\u003c/span\\u003e\\u003c/li\\u003e\\u003c/ol\\u003e\"}],\"fulltextSource\":\"\",\"fullText\":\"\",\"funders\":[],\"hasAdminPriorityOnWorkflow\":false,\"hasManuscriptDocX\":true,\"hasOptedInToPreprint\":true,\"hasPassedJournalQc\":\"\",\"hasAnyPriority\":false,\"hideJournal\":false,\"highlight\":\"\",\"institution\":\"\",\"isAcceptedByJournal\":true,\"isAuthorSuppliedPdf\":false,\"isDeskRejected\":\"\",\"isHiddenFromSearch\":false,\"isInQc\":false,\"isInWorkflow\":false,\"isPdf\":false,\"isPdfUpToDate\":true,\"isWithdrawnOrRetracted\":false,\"journal\":{\"display\":true,\"email\":\"info@researchsquare.com\",\"identity\":\"insectes-sociaux\",\"isNatureJournal\":false,\"hasQc\":true,\"allowDirectSubmit\":false,\"externalIdentity\":\"inso\",\"sideBox\":\"Learn more about [Insectes Sociaux](http://link.springer.com/journal/40)\",\"snPcode\":\"40\",\"submissionUrl\":\"https://www.editorialmanager.com/inso/default2.aspx\",\"title\":\"Insectes Sociaux\",\"twitterHandle\":\"\",\"acdcEnabled\":true,\"dfaEnabled\":true,\"editorialSystem\":\"em\",\"reportingPortfolio\":\"Springer Hybrid\",\"inReviewEnabled\":true,\"inReviewRevisionsEnabled\":false},\"keywords\":\"thermoregulation, meliponiculture, nest sites, Universiti Malaysia Sabah main campus\",\"lastPublishedDoi\":\"10.21203/rs.3.rs-4845014/v1\",\"lastPublishedDoiUrl\":\"https://doi.org/10.21203/rs.3.rs-4845014/v1\",\"license\":{\"name\":\"CC BY 4.0\",\"url\":\"https://creativecommons.org/licenses/by/4.0/\"},\"manuscriptAbstract\":\"\\u003cp\\u003eThe commercial stingless bee \\u003cem\\u003eHeterotrigona itama\\u003c/em\\u003e naturally nests in tree cavities but was kept in wooden boxes in meliponiculture farms. However, at Universiti Malaysia Sabah (UMS), these bees primarily nest in lamp posts. We conducted a temperature survey to assess lamp posts as potential nesting sites for \\u003cem\\u003eH. itama\\u003c/em\\u003e, aiming to determine if the preference for occupied lamp posts was related to their distribution. We measured ambient temperatures in occupied lamp posts in shaded areas and unoccupied lamp posts in exposed areas on the UMS campus, calculating the percentage of time these temperatures fell outside the optimal range for \\u003cem\\u003eH. itama\\u003c/em\\u003e. Additionally, we analyzed the occurrence and timing of temperature regulation in lamp post nests by comparing temperature differences between occupied and unoccupied lamp posts across four-time blocks. Temperature measurements of occupied (shaded) and unoccupied (exposed) lamp posts revealed that shaded lamp posts experienced temperatures outside the bees' optimal range (27\\u0026deg;C-32\\u0026deg;C) less often than exposed lamp posts (68.4% vs. 78.12%). This suggests that \\u003cem\\u003eH. itama\\u003c/em\\u003e may prefer shaded lamp posts due to their more stable temperature profile. Additionally, the internal temperature of lamp posts, whether occupied or not, was consistently 1.54\\u0026ndash;1.76\\u0026deg;C warmer than ambient during hotter periods and closer to ambient during cooler periods, indicating inherent insulation properties of the metal lamp posts. However, a notable difference in temperature between occupied and unoccupied posts was observed in the late afternoon and evening, suggesting active thermoregulation by bees to maintain optimal nest temperature.\\u003c/p\\u003e\",\"manuscriptTitle\":\"Temperature regulation of Heterotrigona itama (Cockerell, 1918) in lamp posts nests\",\"msid\":\"\",\"msnumber\":\"\",\"nonDraftVersions\":[{\"code\":1,\"date\":\"2024-08-30 13:25:13\",\"doi\":\"10.21203/rs.3.rs-4845014/v1\",\"editorialEvents\":[{\"type\":\"communityComments\",\"content\":0},{\"type\":\"decision\",\"content\":\"Major Revisions Needed\",\"date\":\"2024-08-29T02:00:55+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"reviewerAgreed\",\"content\":\"\",\"date\":\"2024-08-09T00:36:02+00:00\",\"index\":0,\"fulltext\":\"\"},{\"type\":\"reviewersInvited\",\"content\":\"\",\"date\":\"2024-08-05T03:49:00+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"editorAssigned\",\"content\":\"\",\"date\":\"2024-08-04T14:34:54+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"submitted\",\"content\":\"Insectes Sociaux\",\"date\":\"2024-08-01T21:56:29+00:00\",\"index\":\"\",\"fulltext\":\"\"}],\"status\":\"published\",\"journal\":{\"display\":true,\"email\":\"info@researchsquare.com\",\"identity\":\"insectes-sociaux\",\"isNatureJournal\":false,\"hasQc\":true,\"allowDirectSubmit\":false,\"externalIdentity\":\"inso\",\"sideBox\":\"Learn more about [Insectes Sociaux](http://link.springer.com/journal/40)\",\"snPcode\":\"40\",\"submissionUrl\":\"https://www.editorialmanager.com/inso/default2.aspx\",\"title\":\"Insectes Sociaux\",\"twitterHandle\":\"\",\"acdcEnabled\":true,\"dfaEnabled\":true,\"editorialSystem\":\"em\",\"reportingPortfolio\":\"Springer Hybrid\",\"inReviewEnabled\":true,\"inReviewRevisionsEnabled\":false}}],\"origin\":\"\",\"ownerIdentity\":\"315baf10-a93a-4867-9fa9-3deac1d43331\",\"owner\":[],\"postedDate\":\"August 30th, 2024\",\"published\":true,\"recentEditorialEvents\":[],\"rejectedJournal\":[],\"revision\":\"\",\"amendment\":\"\",\"status\":\"published-in-journal\",\"subjectAreas\":[],\"tags\":[],\"updatedAt\":\"2024-11-11T16:07:05+00:00\",\"versionOfRecord\":{\"articleIdentity\":\"rs-4845014\",\"link\":\"https://doi.org/10.1007/s00040-024-01006-w\",\"journal\":{\"identity\":\"insectes-sociaux\",\"isVorOnly\":false,\"title\":\"Insectes Sociaux\"},\"publishedOn\":\"2024-11-08 15:56:59\",\"publishedOnDateReadable\":\"November 8th, 2024\"},\"versionCreatedAt\":\"2024-08-30 13:25:13\",\"video\":\"\",\"vorDoi\":\"10.1007/s00040-024-01006-w\",\"vorDoiUrl\":\"https://doi.org/10.1007/s00040-024-01006-w\",\"workflowStages\":[]},\"version\":\"v1\",\"identity\":\"rs-4845014\",\"journalConfig\":\"researchsquare\"},\"__N_SSP\":true},\"page\":\"/article/[identity]/[[...version]]\",\"query\":{\"redirect\":\"/article/rs-4845014\",\"identity\":\"rs-4845014\",\"version\":[\"v1\"]},\"buildId\":\"qtupq5eGEP_6zYnWcrvyt\",\"isFallback\":false,\"isExperimentalCompile\":false,\"dynamicIds\":[84888],\"gssp\":true,\"scriptLoader\":[]}","source_license":"CC-BY-4.0","license_restricted":false}