Research Article |
Corresponding author: Ming Kai Tan ( orthoptera.mingkai@gmail.com ) Academic editor: Corinna S. Bazelet
© 2017 Ming Kai Tan, Taksin Artchwakom, Rodzay Abdul Wahab, Chow-Yang Lee, Daicus M. Belabut, Hugh Tiang Wah Tan.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Tan MK, Artchwakom T, Abdul Wahab RH, Lee C-Y, Belabut DM, Wah Tan HT (2017) Overlooked flower-visiting Orthoptera in Southeast Asia. Journal of Orthoptera Research 26(2): 143-153. https://doi.org/10.3897/jor.26.15021
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The study of insect–plant interactions such as flower visitors, pollinators, and florivores, are important for understanding the natural world. However, not all flower-visiting insects are equally well known, especially in the biodiverse Southeast Asian region. One group is the orthopterans, comprising of grasshoppers, crickets, and katydids. Natural history observations were made around Southeast Asia to document flower-visiting orthopterans. Owing to the limited studies on the ecology of orthopterans in Southeast Asia, we provide here the first documentation of flower-visiting orthopterans from Southeast Asia and the most extensive one for the Tropics. Based on 140 incidences of orthopteran visiting flowers, 41 orthopteran species have so far been recorded to visit 35 different plant species, in mainly Singapore, Malaysia, part of Thailand, and Brunei Darussalam. We conclude that orthopterans are indeed overlooked flower-visitors in this region and warrant further investigation.
florivory, insect-plant, interaction, natural history, pollination
Insects and plants make up a large proportion of the organismal diversity on Earth. Interactions between plants and insects are complex and can be intriguing (
Many insects visit flowers, and some primarily feed on flowers (
Understanding the diversity of flower-visiting orthopterans can have potential applications. As many Southeast Asian countries still rely on agriculture as a main source of economic growth (
Here, we surveyed seven localities around Southeast Asia and made natural history observations of orthopterans visiting flowers. We identified the orthopteran and plant species, whenever possible, and provided notes and remarks on the behaviour and ecology. We aim to provide the first report of flower-visiting orthopterans in this region and provide baseline information for further investigation into flower-visiting, florivory and pollination by orthopterans.
Field observations.—
Natural history observations were carried out between 2015–2017 mainly in seven surveyed sites around Southeast Asia (Fig.
No. | Locality | Habitat(s) Surveyed | Sampling effort (in days) |
1 | Singapore | Scrublands, gardens, herbaceous plots, lowland secondary forests | >100 |
2 | Bukit Larut, Perak, Peninsular Malaysia | Gardens, lowland secondary forests, herbaceous plots, lower montane forests | 12 |
3 | Bukit Fraser, Pahang, Peninsular Malaysia | Gardens, lower montane forests | 27 |
4 | Ulu Gombak Field Studies Centre, Selangor, Peninsular Malaysia | Lowland secondary forests | 4 |
5 | Pulau Tioman, Pahang, Peninsular Malaysia | Gardens, lowland secondary forests | 3 |
6 | Sakaerat Biosphere Reserve, Nakhon Ratchasima, Thailand | Dry dipterocarp forest, dry evergreen forest, herbaceous plots | 36 |
7 | Kuala Belalong Field Studies Centre, Temburong, Brunei Darussalam | Gardens, primary hill dipterocarp forests | 21 |
Analyses
.—To visualise and summarise the respective orthopteran species that visited a specific flower species, an interaction network was constructed using the ‘plotweb’ function in bipartite package (
We recorded 140 incidences of orthopterans visiting flowers in five countries around Southeast Asia: Singapore (82), Peninsular Malaysia (23), Thailand (27), Brunei Darussalam (7), and Indonesia (1). While the sampling was distinctly higher in Singapore, the species that were recorded are mostly Southeast Asian species and can be found in most parts of Southeast Asia. Habitats with the largest number of observations include scrublands (59), and evergreen dipterocarp forest and gardens (both 13). In total, 99 records were of Ensifera (crickets and katydids) in contrast to 41 of Caelifera (grasshoppers). Fifty of the orthopteran records were adults whereas 90 were nymphs.
Forty-one orthopteran species from six families were recorded to visit flowers of 35 plant species from 15 families (Fig.
The interaction web between flower-visiting orthopterans (right row) and flower species (left row) in Southeast Asia. The width of the linkage represents the number of observations. Legends for orthopteran families: Acri = Acrididae; Grylla = Gryllacrididae; Grylli = Gryllidae; Mogo = Mogoplistidae; Pyrg =Pyrgomorphidae; Tett = Tettigoniidae. Legends for flower families: Aca = Acanthaceae; Ama = Amaryllidaceae; Apo = Apocynaceae; Ast = Asteraceae; Con = Convolvulaceae; Cos = Costaceae; Dil = Dilleniaceae; Fab = Fabaceae; Mal = Malvaceae; Mel = Melastomataceae; Orc = Orchidaceae; Poa = Poaceae; Rub = Rubiaceae; Ver = Verbenaceae; Vit = Vitaceae.
We found that most flower-visiting orthopterans visit very few flower species (Fig.
Examples of common flower-visiting orthopterans from Southeast Asia: A. Phaneroptera brevis feeding on the anthers of Tithonia diversifolia in Bukit Larut, B. Nisitrus species feeding on the petals of Centrosema molle in Ulu Temburong, C. Valanga nigricornis feeding on a corolla lobe of Hibiscus rosa-sinensis in Pulau Tioman, D. Conocephalus species feeding on the florets of Ageratum conyzoides, E. Xenocatantops humilis feeding on a corolla lobe of Gardenia jasminoides in Singapore, and F. Atractomorpha species on the corolla of Ipomoea pes-caprae in Pulau Tioman.
Orthopterans are only some of the many invertebrates that visit flowers (
There are two main types of orthopterans that visit flowers. Firstly, some orthopterans are floriphilic, clearly preferring flowers over other plant parts as their diet (
Examples of unidentified Phaneropterinae nymphs visiting flowers of various plants: A. Dillenia suffruticosa in Singapore, B. Acacia auriculiformis in Singapore, C. Costus lucanusianus in Singapore, D. Youngia japonica in Bukit Larut, E. Praxelis clematidea in Sakaerat, and F. Lantana camara in Sakaerat.
The second group of flower-visiting orthopterans are opportunistic polyphagous species. These polyphagous species are usually folivores (feeding on the foliage) but can be facultative florivores when floral parts are available (
Nonetheless, there are also other interesting encounters of flower-visiting orthopterans. These include a predatory katydid from the subfamily Meconematinae feeding on flowers of Dillenia suffruticosa (Griff. ex Hook.f. and Thomson) (Fig.
Examples of flower-visiting orthopterans on native plant species: A. Alloteratura species on Dillenia suffruticosa in Singapore, B. Tremellia timah on Dillenia suffruticosa in Singapore, C. Elimaea carinata on Melastoma malabathricum in Singapore, and D. Ornebius rufonigrus on Ixora congesta in Singapore.
We observed that non-native weeds in Southeast Asia are frequently visited by many orthopterans. These weeds tend to flower frequently and abundantly. In scrublands in Singapore, B. pilosa is a prominent weed and is known to be an important food source for many flower-visitors including the pollinator bees (
Gardens, where different kinds of flowers (often of non-native species) are planted, are another habitat with numerous records of flower-visiting orthopterans. In the highlands of Peninsular Malaysia, gardens are common in hill resorts and the flowers tend to attract floriphilic orthopterans. Some of these flowering plants (e.g. Ageratum houstonianum Miller and Tithonia diversifolia (Hemsl.) (Fig.
Native plant species that flower regularly are also visited by orthopterans. These include D. suffruticosa (Fig.
While we provide baseline information of flower-visiting orthopterans in Southeast Asia, there is still a dearth of information on the ecology and behaviours of flower-visiting orthopterans from this region. A major area for further study is to monitor how flower-visiting orthopterans respond to synchronous flowering events in the dipterocarp forests (
We should also aim to better understand the distribution of flower-visiting orthopterans as well as how to predict their occurrences. Although presence-only data can be challenging to analyse using conventional modelling techniques, recent development of MaxEnt modelling can help to overcome the shortcomings of such data (
MKT thanks L. Roman Carrasco for providing constructive comments on the manuscript and Louise Neo for verification of plant and flower identification. The permission for the collection of material in Kuala Belalong Field Studies Centre, Brunei Darussalam was kindly granted by the Institute for Biodiversity and Environmental Research, Universiti Brunei Darussalam (UBD/AVC-RI/1.21.1 [a]). The authors are grateful to the Institute for Biodiversity and Environmental Research, Universiti Brunei Darussalam, for allowing us to work in Kuala Belalong Field Studies Centre, and to the Biodiversity and Research Innovation Centre (BioRIC), Ministry of Industry and Tourism, Brunei Darussalam for the issuance of our export permits (BioRIC/HoB/TAD/51-73 and 51-80). The permission for the collection of material in the Sakaerat Environmental Research Station was kindly granted by the National Research Council, Thailand (No. 0002/209, Registration no. 9/57). MKT is also grateful to the staff members of the Sakaerat Environmental Research Station for their support and hospitality. Permission for the collection of material in Peninsular Malaysia was kindly granted by the Research Promotion and Co-Ordination Committee, Economic Planning Unit, Prime Minister’s Department (UPE: 40/200/19/3103 for Bukit Larut, 40/200/19/2923 for Bukit Fraser and 40/200/19/3395 for Pulau Tioman) and supported by the Institute for Biodiversity, Department of Wildlife and National Parks (Perhilitan) and Universiti Malaya. Permission for the collection of material in Singapore was kindly granted by the National Parks Board (NP/RP10-073). The work of MKT was supported by the Lady Yuen Peng McNeice Graduate Fellowship of the National University of Singapore.