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UC Riverside UC Riverside Previously Published Works UC Riverside UC Riverside Previously Published Works Title Rattan litter-collecting structures attract nest-building and defending ants. Permalink https://escholarship.org/uc/item/8s3837x7 Journal Plant signaling & behavior, 14(8) ISSN 1559-2316 Authors Liu, Kunpeng Mansor, Asyraf Ruppert, Nadine et al. Publication Date 2019 DOI 10.1080/15592324.2019.1621245 Peer reviewed eScholarship.org Powered by the California Digital Library University of California Plant Signaling & Behavior ISSN: (Print) 1559-2324 (Online) Journal homepage: https://www.tandfonline.com/loi/kpsb20 Rattan litter-collecting structures attract nest- building and defending ants Kunpeng Liu, Asyraf Mansor, Nadine Ruppert, Chow Yang Lee, Nur Munira Azman & Nik Fadzly To cite this article: Kunpeng Liu, Asyraf Mansor, Nadine Ruppert, Chow Yang Lee, Nur Munira Azman & Nik Fadzly (2019) Rattan litter-collecting structures attract nest-building and defending ants, Plant Signaling & Behavior, 14:8, 1621245, DOI: 10.1080/15592324.2019.1621245 To link to this article: https://doi.org/10.1080/15592324.2019.1621245 Published online: 27 May 2019. Submit your article to this journal Article views: 43 View Crossmark data Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=kpsb20 PLANT SIGNALING & BEHAVIOR 2019, VOL. 14, NO. 8, e1621245 (8 pages) https://doi.org/10.1080/15592324.2019.1621245 SHORT COMMUNICATION Rattan litter-collecting structures attract nest-building and defending ants Kunpeng Liu, Asyraf Mansor , Nadine Ruppert , Chow Yang Lee, Nur Munira Azman , and Nik Fadzly School of Biological Sciences, Universiti Sains Malaysia, Minden, Malaysia ABSTRACT ARTICLE HISTORY Rattan is an important climbing palm taxon in Malaysian tropical rain forests. Many rattan species have Received 23 March 2019 unique structures directly associated with certain ant species. In this study, four rattan species Revised 10 May 2019 (Daemonorops lewisiana, Calamus castaneus, Daemonorops geniculata and Korthalsia scortechinii) were Accepted 16 May 2019 inspected and documented in a field survey concerning their relationships with several ant species. We KEYWORDS noticed that two rattan species (D. lewisiana and C. castaneus) were more likely to be associated with Rattan; spine; leaf structure; ants compared to their neighbouring rattan (Plectomia griffithii). However, D. lewisiana and C. castaneus leaf litter; ants did not directly provide shelters for ant colonies, but possessed unique structures: upward-pointing spines and funnel-shaped leaves, which are equipped to collect more litter than P. griffithii. To test our litter collecting hypothesis, we measured the inclination of spines from the stem. Our results showed the presence of ant colonies in the litter-collecting rattans (D. lewisiana and C. castaneus), which was significantly higher compared to a non-litter-collecting rattan (P. griffithii). We propose a complex and novel type of adaptation (litter-collection and provision of nesting materials) for rattans, which promotes interactions between the rattan and ants through the arrangements of leaves, leaflets, and spines. In return, the rattan may benefit from ants’ services, such as protection, nutrient enhancement, and pollination. Introduction Asia, Korthalsia is a common rattan genus associated with ants.2,15,18,19 Other rattan taxa with similar relationships Rattans are spiny palms belonging to the subfamily include Calamus sp. and Daemonorops sp.20 Calamoideae. About 600 species of rattans belonging to 13 2 Dransfield reported six rattan adaptations to cater for genera are recorded worldwide.1 Some rattans are climbers, associated ant colonies. Ocreas (inflated extension of leaf which possess climbing structures, such as cirrus or sheath) are hollow chambers that form desirable shelters for a flagellum, but some are acaulescent or non-climbing ant colonies in some Korthalsia sp. and a few Calamus sp. species.2 The flexible stems of certain rattan species are used Calamus longipinna’s sac-like ocreas are domatia for ant by humans for matting, binding, and furniture production.3 nesting.21 Another type of elongated ocreas (only known in In Peninsular Malaysia, rattans can be found from sea level to Korthalsia hispida) provides tube-like chambers for ants to mountains over 1000-m high. They are an important compo- 2 colonize. Furthermore, ocreas are armed with spines which nent of the tropical primary rainforest and secondary forests may provide protection for ant colonies. Korthalsia furta- but remain poorly understood.2 doana has an obligate symbiont ant species, Camponotus Ants are often regarded as the most successful eusocial sp.18 K. furtadoana enjoys healthier leaves if colonized by animals. Their relationships with plants have long been estab- certain ant species.19 In addition to Korthalsia, Daemonorops lished. These interactions include leaf-harvesting for fungal verticillaris, has overlapping spine structures that provide 4 5,6 gardens; seed-harvesting and seed-dispersal; Plants bearing cavities for ants. In return, the ants help the rattan absorb domatia for ant colonies are called myrmecophytes, benefiting nutrients and water at higher efficiency.22 Simple adaptations ’ 7-11 from symbiont ants protection against herbivores; and involving leaflet or inflorescences as well as adaptation with protection from Crematogaster, which are the obligate ant leaf sheath auricles are known but rarely studied.2 partner of Macaranga, regulated by the food supply (food In this study, we approached the fundamental function 12,13 bodies) of their host plants. Several cases showed that of a rattan spine from a different perspective. We ask the 14 ants can also serve as pollinators. question of whether the spines could function as a leaf litter Rattan–ant relationship across various species has been collecting structure. We hypothesize that there will be ant reported in the past.2,3,15-18 Sunderland17 reported on colonies in litter-collecting rattans (Daemonorops lewisiana a variety of ant species colonizing various parts of several and Calamus castaneus)incomparisontonon-littercollect- rattan species in Africa, with some ant species showing ing rattan (Plectomia griffithii). This study provides about a specific preference towards certain rattan species, both in a novel adaptation of certain rattan species which facilitates terms to their protective behavior and relationships with the a relationship with ants, which has yet to be reported to resident scale insects. In the tropical rain forests of Southeast date. CONTACT Nik Fadzly [email protected] School of Biological Sciences, Universiti Sains Malaysia, 11800 Minden, Penang, Malaysia © 2019 Taylor & Francis Group, LLC e1621245-2 K. LIU ET AL. Figure 1. Study site locations at (a) Bukit Genting Hill, (b) Penang National Park, (c) Taman rimba, Teluk Bahang and (d) Cherok Tokun, Penang. Materials and methods Table 1. Ant genera found on various rattan species. – Individual Rattan species Location Ant species Surveys on rattan ant relationship were conducted in north- Daemonorops ern Peninsular Malaysia, Bukit Genting Hill (5°18ʹ31.9”N, 1 D. lewisiana PNP Philidris sp. 100°13ʹ14.0”E), Penang National Park (5°28ʹ01.4”N, 100° 2 D. lewisiana PNP Dolichoderus thoracicus ʹ ” ʹ ” 3 D. lewiisana PNP Crematogaster sp.1 11 56.0 E) and Taman Rimba, Teluk Bahang (5°26 52.51 N, 4 D. lewiisana PNP Dolichoderus thoracicus 100°13ʹ4.89”E) in Penang island, and in Cherok Tokun, 5 D. lewisiana PNP Philidris sp. Penang mainland (05°21ʹ54.6”N, 100°28ʹ58.7”E; Figure 1). 6 D. lewisiana PNP Philidris sp. 7 D. lewisiana PNP Philidris sp. Rattans were identified based on keys developed by 8 D. lewisiana PNP Crematogaster sp.2 Dransfield.2 The associated ants were collected using forceps, 9 D. lewisiana PNP Philidris sp. 10 D. lewiisana PNP Philidris sp. kept in 70% ethanol, and transported to the laboratory for 11 D. lewisiana PNP Tapinoma melanocephalum identification using keys developed by Hashimoto.23 12 D. lewisiana PNP Philidris sp. 13 D. lewisiana PNP Philidris sp. The three most abundant rattan species in Penang National 14 D. lewisiana BGH Technomyrmex sp.2 Park and Taman Rimba, Teluk Bahang are Plectomia griffithii, 15 D. lewisiana BGH Technomyrmex sp.2 Daemonorops lewisiana and Calamus castaneus. We recorded 16 D. geniculata BGH Camponotus sp. 17 D. geniculata CT Crematogaster sp.5 for each individual rattan clump (stem length > 50 cm) whether 18 D. geniculata CT Crematogaster sp.6 an ant colony was present, absent or abandoned. Any rattan with 19 D. geniculata CT Dolichoderus thoracicus Calamus a sign of tunnels on the surface, but an absence of ants, was 20 C. castaneus TRTB Crematogaster sp.3 considered abandoned. Rattan seedlings (< 50 cm) were ignored 21 C. castaneus TRTB Crematogaster sp.3 as they were too small to collect leaf litter. 22 C. castaneus TRTB Crematogaster sp.3 23 C. castaneus CT Tapinoma melanocephalum To examine the leaf-collecting structures, we measured the 24 C. castaneus CT Dolichoderus thoracicus inclination of spines from the stem. Ten spines were randomly 25 C. castaneus CT Pheidole sp. 26 C. castaneus CT Technomyrmex sp.1 chosen from each species. We compared the spine angles among 27 C. castaneus CT Crematogaster sp.4 the three species. The paired leaflet angles were also measured in 28 C. castaneus CT Pheidole sp. Korthalsia D. lewisiana and C. castaneus. Using a protractor, we measured 29 K. scortechinii CT Camponotus beccarii the angles
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