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Characterization of Nest Architecture of an Indian Ant Diacamma Indicum (Hymenoptera: Formicidae) Kushankur Bhattacharyya1 and Sumana Annagiri1,2

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Characterization of Nest Architecture of an Indian Ant Diacamma Indicum (Hymenoptera: Formicidae) Kushankur Bhattacharyya1 and Sumana Annagiri1,2 Journal of Insect Science, (2019) 19(6): 9; 1–8 doi: 10.1093/jisesa/iez083 Research Characterization of Nest Architecture of an Indian Ant Diacamma indicum (Hymenoptera: Formicidae) Kushankur Bhattacharyya1 and Sumana Annagiri1,2, 1Behaviour and Ecology Lab, Department of Biological Sciences, Indian Institute of Science Education and Research, Kolkata, Mohanpur, West Bengal-741246, India and 2Corresponding author, e-mail: [email protected] Subject Editor: Yael Lubin Received 24 April 2019; Editorial decision 6 August 2019 Abstract Nests are physical entities that give shelter to the inhabitants from natural adversities, predators and act as a platform for organization of tasks particularly in social insects. Social insect nests can range from simple structures consisting of a single entrance leading to a chamber to complex nests containing hundreds of connected shafts and chambers. This study characterizes nest architecture of a tropical ponerine ant Diacamma indicum (Hymenoptera: Formicidae), which has small colony sizes and is known to be a scavenger. We also examined if these nests vary seasonally. By examining the microhabitat in the vicinity of the nest, the nest entrance characteristics and casting 77 natural nests of D. indicum across a year, we found that this species occupies relatively simple nests consisting of a single entrance that leads to a single chamber. This chamber progressed to a secondary tunnel that terminated at a greater depth than the chamber. The nest volume was not correlated to the number of adult members in the colony. Even though the microhabitat around the nest and the entrance itself change across seasons, principle component analysis showed that the nest architecture remained similar. Only one parameter, the entrance tunnel showed significant difference and was longer during postmonsoon. Nests of colonies living in the immediate vicinity of human habitation were comparable to other nests. We conclude that D. indicum found in the Gangetic plains live in relatively simple nests that do not vary across seasons. Key words: Ponerinae, seasonal variation, nest microhabitat, nest chirality, nest volume A nest is a place or a specially modified structure serving as an abode but to the best of our knowledge, its impact on nest architecture in for animals and especially during their immature young. Animal tropics is relatively unexplored (Tschinkel 1999a,b; Cristiano et al. nests have fascinated us over the centuries with their monumental 2019). What we do know is that as colony size increases the number building efforts as well as their enigmatic architectural features of chambers as well as the volume of these chambers increase in case (Hansell and Ruxton 2008). Nests are important to organisms that of Pogonomyrmex badius, Phedole morrisi, Solenopsis invicta, and occupy them as they give protection from predators, environmental some other species (Tschinkel 2015). Sometimes, the chamber edges adversities, and provide a storage space for resources (Hansell 1993). become more lobed as the chamber grows (Tschinkel 2015). In the Nests are also important for the establishment of territories and pro- temperate ant Odontomachus brunneus, winter nests are twice as viding group identity and for the organization of work, particularly deep (up to 170 cm) as summer nests (up to 60 cm), even though in the case of social insects (Robson et al. 1995). Nesting sites of ants colonies consisted of similar number of adults (Hart and Tschinkel are diverse and can range from subterranean soil nests to cavities in 2012). When Temnothorax rugatulus, colonies were subjected to living trees or leaves stitched together forming arboreal chambers higher humidity and lower airflow, they built nests with thicker and occasionally even consist of living workers orienting themselves walls, indicating that colonies are potentially capable of adapting to form a chamber (Tschinkel 2015). The generalized architecture their nests to differences in their natural habitat (DiRienzo and of subterranean ant nests consists of a vertical shaft connecting the Dornhaus 2017). entrance to a horizontal chamber. This unit can be repeated many There are several methods to study the architecture of subter- times over to form complex structures such as those found in leaf ranean ant nests. Careful excavation of the nest, layer by layer, ex- cutter ant Atta leviagata nests, containing up to 7,800 chambers that posing one chamber at a time can give an idea about chamber size extend as deep as 7 m underground (Moreira et al. 2004). and depth, but fine structural details are lost. Casting of nests pro- The impact of seasonal variation has been examined on life his- vides detailed information regarding the nest structure and is con- tory traits like alate formation, brood production, and colony size, sidered to be the best and most convenient option. Materials like © The Author(s) 2019. Published by Oxford University Press on behalf of Entomological Society of America. This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs licence (http://creativecommons.org/licenses/by-nc-nd/4.0/), which permits non-commercial 1 reproduction and distribution of the work, in any medium, provided the original work is not altered or transformed in any way, and that the work is properly cited. For commercial re-use, please contact [email protected] 2 Journal of Insect Science, 2019, Vol. 19, No. 6 paraffin wax, dental plaster, or metals with low melting point like foragers that were returning to their nest. A total of 77 nests were Al or Zn are poured into the nest. The molten metal occupies the sampled across the study period: monsoon (28), postmonsoon (25), hollow structures within the nest and solidifies to produce a three- and premonsoon (24). Upon discovery of a nest several entrance dimensional cast of the nest. While wax and dental plaster allow for features like diameter, decoration, and presence of mound were the recovery of the dead ants from the nest cast, metal casts provide recorded and building index (BI) scores were assigned. BI values greater detail of the three-dimensional nest structure and is less prone ranged from 0 to 5, where 0 was allocated for a bare open nest and 5 to fragmentation during recovery and storage (Tschinkel 2010). Both for a nest entrance which had elaborate decorations and an elevated of these methods lead to the destruction of the nest and its inmates. entrance with a consolidated mound and many decorations around Two noninvasive methods namely X-ray and computed tomography the entrance (Kolay and Annagiri 2015). For further examination, a can also give information on the three-dimensional structure of the photograph of each nest entrance was taken with a scale in the back- nest without destroying the nest and its inmates(Halley et al. 2005). ground using a still camera (Canon Powershot SX15X). Additional Diacamma indicum (Ponerinae), is a black, 1-cm long ant, found analyses of entrance diameter and area were carried out using these in India, Sri Lanka, and perhaps Japan. Diacamma indicum colonies pictures with ImageJ software. In order to evaluate the area sur- consist of adult monomorphic adult females, that is, all the individ- rounding the nest entrance a circle with a radius of three D. indicum uals (reproductive caste and worker caste) are similar in size and body length (3 cm) from the center of the nest entrance was drawn. morphology and they show solitary foraging to bring back parts of Within this focal microhabitat, the area covered by the four most termites and other dead insects (personal observation) (Kushankur common materials, that is, soil, vegetative material, caterpillar cu- Bhattacharyya). The colony size ranges from 12 to 260 adults. They ticle, and bird feathers were measured using ImageJ software. For live mostly in subterranean nests and occasionally occupy other lo- each of these materials, scores from 0 to 4 were assigned. Scores cations like crevices below rocks, bricks, walls, and tree branches were given according to the percentage area covered by each of these (Viginier et al. 2004, Kaur and Sumana 2014, Kolay and Annagiri materials except for soil. When any given material was absent, it was 2015). This species is known to be monodomous and occupy scored as 0. When a given material covered 1 to 25% area within shallow nests or move to higher elevations during monsoon (Kaur the focal area, a score of 1 was assigned; similarly coverage between and Sumana 2014, Kolay and Annagiri 2015). Across several relo- 25–50%, 51–75%, and 76–100% were scored 2, 3, and 4, respect- cations conducted in nature, D. indicum occupied pre-existing holes ively. However, with soil, the calculation of percentage soil would or cavities and they were not seen excavating a nest of their own underestimate the height of soil present and also did not account for (Kaur et al. 2012). In this study, we did field surveys across a year the labor associated with consolidation of soil as compared to loose to characterize the living quarters or nests of D. indicum in its nat- soil around the entrance. Thus, for soil, we scored small amount of ural habitat, in eastern India along the Gangetic plains. The annual loose soil as 1; medium amount of loose soil as 2; small but consoli- temperature ranges from 8 to 48°C in this region and rainfall occurs dated soil mound as 3, and consolidated mound with large amount mostly in the months of June to September while the remainder of of soil as 4. Sum of the scores from all these four materials was the year it is mostly dry. We hypothesized that the seasonal variation termed as ‘nest entrance microhabitat or NEMH’ and was used to will impact nest architecture of this species. We characterized the get a comprehensive view of the nest entrance surroundings.
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