Skip to main content Skip to main navigation menu Skip to site footer
Type: Monograph
Published: 2018-08-06
Page range: 1–71
Abstract views: 414
PDF downloaded: 6

The firefly genus Pteroptyx Olivier revisited (Coleoptera: Lampyridae: Luciolinae)

Department of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore 117543, Singapore. Department of Biology, University of Turku 20014 Turku, Finland
School of Agricultural and Wine Sciences, Charles Sturt University, Wagga Wagga 2678, Australia.
Queensland Museum, PO Box 3300 South Brisbane, 4101 Australia.
Zasve @ UM Innovations Sdn. Bhd., University of Malaya 50603 Kuala Lumpur, Malaysia
Section of Biodiversity, Department of Biology, Lund University 223 62 Lund Sweden
Luciolinae taxonomy phylogeny morphology molecular habitat Southeast Asia ecotourism Luciola testacea Motsch. Poluninius Ballantyne Coleoptera

Abstract

The synchronous firefly genus Pteroptyx Olivier is reassessed from morphological, molecular, and habitat perspectives in Malaysia, and includes some reliably associated females described from morphological features and internal female reproductive anatomy. Phylogenetic analyses using combined morphological and molecular data (where available) for 158 taxa supported all the major features of the existing taxonomic categories within the Indopacific Luciolinae. They revealed a distinct Pteroptyx clade as a morphologically variable genus with Poluninius selangoriensis Ballantyne being newly synonymised with Luciola testacea Motschulsky, the type species, which is redescribed from the type series. Pteroptyx gelasina Ballantyne was shown to be distinct and three of the four morphological subdivisions within Pteroptyx malaccae (Gorham) considered useful. A new species Pt. balingiana Jusoh sp. nov. is described from Sarawak. A second specimen of Pt. gombakia Ballantyne is described and figured.

        Some females were reliably associated with identified males by molecular data, but investigation of their morphology showed consistent features that were for the most part not useful for species delineation, which still relies on association with the males and colour patterns. All females investigated had bursa plates.

Habitat details for most Pteroptyx revealed an association with a riparian environment likely to support mangroves but not necessarily an obligatory association with mangroves or any particular species. Pteroptyx galbina Jusoh was found up to 30 km from the sea, and Pt. bearni Olivier displays in a variety of flowering plants alongside rivers, including mangroves.

Keys to species and diagnoses of all species with coloured plates are given.

References

  1. Adobe Systems (2012) Adobe Illustrator CS6 Version 16.0.3. [software]

    Baker, R.H. & DeSalle, R. (1997) Multiple sources of character information and the phylogeny of Hawaiian Drosophilids. Systematic Biology, 46, 654–673.
    https://doi.org/10.1093/sysbio/46.4.654

    Ballantyne, L.A. (1987a) Further revisional studies on the firefly genus Pteroptyx Olivier (Coleoptera: Lampyridae: Luciolinae). Transactions of the American Entomological Society, 113, 117–170.

    Ballantyne, L.A. (1987b) Lucioline morphology, taxonomy and behaviour: a reappraisal. (Coleoptera, Lampyridae). Transactions of the American Entomological Society, 113, 171–188.

    Ballantyne, L.A. (2001) The bent winged fireflies of Cambodia, Indonesia, Malaysia, Philippines and Thailand (Coleoptera: Lampyridae: Luciolinae: Luciolini). Pteroptyx spp. of the Polunin collection. Serangga, 6 (1), 51–95.

    Ballantyne, L., Fu, X.H., Shih, C.H., Cheng, C.Y. & Yiu, V. (2011) Pteroptyx maipo Ballantyne, a new species of bent-winged firefly (Coleoptera: Lampyridae) from Hong Kong, and its relevance to firefly biology and conservation. Zootaxa, 2931, 8–34.

    Ballantyne, L.A. & Lambkin, C.L. (2000) Lampyridae of Australia (Coleoptera: Lampyridae: Luciolinae: Luciolini). Memoirs of the Queensland Museum, 46 (1), 15–93.

    Ballantyne, L.A. & Lambkin, C.L. (2001) A new firefly, Luciola (Pygoluciola) kinabalua, new species (Coleoptera: Lampyridae), from Malaysia, with observations on a possible copulation clamp. The Raffles Bulletin of Zoology, 49, 363–377.

    Ballantyne, L.A. & Lambkin, C.L. (2006) A phylogenetic reassessment of the rare South East Asian firefly genus Pygoluciola Wittmer (Coleoptera: Lampyridae: Luciolinae). The Raffles Bulletin of Zoology, 54, 21–48.

    Ballantyne, L.A. & Lambkin, C.L. (2009) Systematics of Indo-Pacific Fireflies with a redefinition of Australasian Atyphella Olliff, Madagascan Photuroluciola (Pic), and description of seven new genera from the Luciolinae (Coleoptera: Lampyridae). Zootaxa, 1997, 1–188.

    Ballantyne, L.A. & Lambkin, C.L. (2013) Systematics and phylogenetics of Indo-Pacific Luciolinae fireflies (Coleoptera: Lampyridae) and the description of new genera. Zootaxa, 3653 (1), 1–162.
    https://doi.org/10.11646/zootaxa.3653.1.1

    Ballantyne, L.A., Lambkin, C.L., Boontop, Y. & Jusoh, W.F.A. (2015) Revisional studies on the Luciolinae fireflies of Asia (Coleoptera: Lampyridae): 1. The genus Pyrophanes Olivier with two new species. 2. Four new species of Pteroptyx Olivier and 3. A new genus Inflata Boontop, with redescription of Luciola indica (Motsch.) as Inflata indica comb. nov. Zootaxa, 3959 (1), 1–84.

    Ballantyne, L.A. & McLean, M.R. (1970) Revisional studies on the firefly genus Pteroptyx Olivier (Coleoptera: Lampyridae: Luciolinae: Luciolini). Transactions of the American Entomological Society, 96, 223–305.

    Ballantyne, L.A. & Menayah, R. (2000) Redescription of the synchronous firefly, Pteroptyx tener Olivier (Coleoptera: Lampyridae), of Kampung Kuantan, Selangor. Malayan Nature Journal, 54 (4), 323–328.

    Ballantyne, L.A. & Menayah, R. (2002) A description of larvae and redescription of adults of the firefly Pteroptyx valida Olivier in Selangor, Malaysia (Coleoptera: Lampyridae: Luciolinae), with notes on Luciolinae larvae. Raffles Bulletin of Zoology, 50 (1), 101–109.

    Ballantyne, L.A., Wong, T.-W. & Goh, C.A.L. (2012) Pteroptyx macdermotti McLean-the unusual firefly. Poster presentation. Lampyrid, 2, 197.

    Bremer, K. (1988) The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution, 42, 795–803.
    https://doi.org/10.1111/j.1558-5646.1988.tb02497.x

    Bremer, K. (1994) Branch support and tree stability. Cladistics, 10, 295–304.
    https://doi.org/10.1006/clad.1994.1019

    Buck, J.B. (1938) Synchronous rhythmic flashing of fireflies. The Quarterly Review of Biology, 13 (3), 301–314.
    https://doi.org/10.1086/394562

    Buck, J. (1988) Synchronous rhythmic flashing of fireflies. II. The Quarterly Review of Biology, 265–289.
    https://doi.org/10.1086/415929

    Buck, J. & Buck, E. (1966) Biology of synchronous flashing of fireflies. Nature, 211, 562–564.
    https://doi.org/10.1038/211562a0

    Buck, J., Buck, E., Hanson, F.E., Case, J., Mets, L. & Atta, G.J. (1981a) Control of flashing in fireflies IV. Free run pacemaking in a synchronic Pteroptyx. Journal of Comparative Physiology, 144, 277–286.
    https://doi.org/10.1007/BF00612559

    Buck, J., Buck, E., Case, J.F. & Hanson, F.E. (1981b) Control of flashing in fireflies V. Pacemaker Synchronization in Pteroptyx cribellata. Journal of Comparative Physiology, 144, 287–298.
    https://doi.org/10.1007/BF00612560

    Calder, A.A. (1998) Coleoptera: Elateroidea. In: Wells, A. (Ed.), Zoological Catalogue of Australia 29.6. CSIRO Publishing, Melbourne, pp. 172–180.

    Case, J.F. (1980) Courting behavior in a synchronously flashing, aggregative firefly, Pteroptyx tener. Biologists Bulletin, 159, 613–625.

    Case, J.F. (1984) Vision in Mating Behaviour of Fireflies. In: Lewis, T. (Ed.), Insect Communication. Royal Entomological Society symposium. Academic Press, Orlando, pp. 195–222.

    Chey, V.K. (2004) Fireflies of Sungai Klias and their display trees. Sepilok Bulletin, 1, 67–69.

    Chey, V.K. (2008) Fireflies of Sepilok. Sepilok Bulletin, 9, 3–11.

    Chey, V.K. (2009) Fireflies of Tuaran. Sepilok Bulletin, 10, 25–33.

    Chey, V.K. (2010) Fireflies of Beaufort with special reference to Sungai Garama and Sungai Klias. Sepilok Bulletin, 12, 13–19.

    Chey, V.K. (2011) Fireflies of Pulau Sakar. Sepilok Bulletin, 13 & 14, 27–32.

    Cunningham, C.W. (1997) Can three incongruence tests predict when data should be combined. Molecular Biology and Evolution, 14, 733–740.
    https://doi.org/10.1093/oxfordjournals.molbev.a025813

    de Pinna, M. (1991) Concepts and tests of homology in the cladistic paradigm. Cladistics, 7, 367–394.
    https://doi.org/10.1111/j.1096-0031.1991.tb00045.x

    Farris, J.S. (1990) Phenetics in camouflage. Cladistics, 6, 91–100.
    https://doi.org/10.1111/j.1096-0031.1990.tb00528.x

    Farris, J.S., Källersjö, M., Kluge, A.G. & Bult, C. (1995) Constructing a significance test for incongruence. Systematic Biology, 44, 570–572.
    https://doi.org/10.1093/sysbio/44.4.570

    Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3 (5), 294.

    Foo, K. & Mahadimenakbar, M.D. (2015) Diversity of fireflies (Coleoptera: Lampyridae) of Sungai Teratak, Sabah, Malaysia. Journal of Tropical Biology and Conservation, 12, 1–11.

    Foo, K. & Mahadimenakbar, M.D. (2016) Short notes on fireflies of Sungai Kawang, Sabah. Journal of Tropical Biology and Conservation, 13, 125–128.

    Fu, X., Ballantyne, L.A. & Lambkin, C.L. (2012a) Emeia gen. nov., a new genus of Luciolinae fireflies from China (Coleoptera: Lampyridae) with an unusual trilobite-like larva, and a redescription of the genus Curtos Motsch. Zootaxa, 3403, 1–53.

    Fu, X., Ballantyne, L.A. & Lambkin, C.L. (2012b) The external larval morphology of aquatic and terrestrial Luciolinae fireflies (Coleoptera: Lampyridae). Zootaxa, 3405, 1–34.

    Gabito, O.A. (2013) Growing rural employment through community-based tourism. 12th National Convention on Statistics (NCS), EDSA Shangri-La Hotel, Mandaluyong City, October 1–2, 2013. Available from: http://nap.psa.gov.ph/ncs/12thncs/papers/INVITED/IPS-16%20Tourism%20Statistics/IPS-16_2_Growing%20Rural%20Employment%20Through%20Community-Based%20Tourism.pdf (accessed 24 May 2015)

    Gorham, H.S. (1880) Materials for a revision of the Lampyridae. Transactions of the Entomological Society of London, 1880, 83–112, 1 pl.

    Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98.

    Hanson, F.E. (1978) Comparative studies of firefly pacemakers. Federation Proceedings, 37, 2158–2164.

    Hanson, F.E., Case, J.F., Buck, E. & Buck, J. (1971) Synchrony and flash entrainment in a New Guinea Firefly. Science, 174, 161–164.
    https://doi.org/10.1126/science.174.4005.161

    Hashim, N.R. (2011) Handbook for Field Survey and SWOT Analysis of Congregating Fireflies in Malaysia. Universiti Putra Malaysia Press, Serdang, 23 pp.

    Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 17 (8), 754–755.
    https://doi.org/10.1093/bioinformatics/17.8.754

    Jusoh W.F.A. (2015) Integrative taxonomy and molecular phylogenetic analysis of the bent-winged fireflies (Coleoptera: Lampyridae: Pteroptyx) in Peninsular Malaysia and Sarawak. PhD Dissertation, Universiti Putra Malaysia, Serdang, 119 pp.

    Jusoh, W.F.A.W. & Hashim, N.R. (2012) The effect of habitat modification on firefly populations at the Rembau-Linggi estuary, Peninsular Malaysia. Lampyrid, 2, 149–155.

    Jusoh, W.F.A., Hashim, N.R. & Adam, N.A. (2013) Distribution of the synchronous flashing beetle Pteroptyx tener Olivier (Coleoptera: Lampyridae) in Malaysia. The Coleopterists Bulletin, 67 (4), 604–605.
    https://doi.org/10.1649/0010-065X-67.4.604

    Jusoh, W.F.A., Hashim, N.R. & Ibrahim, Z.Z. (2010a) Firefly distribution & abundance on mangrove vegetation assemblages in Sepetang estuary, Peninsular Malaysia. Wetlands Ecology & Management, 18 (3), 367–373.
    https://doi.org/10.1007/s11273-009-9172-4

    Jusoh, W.F.A.W., Hashim, N.R. & Ibrahim, Z.Z. (2010b) Distribution and Abundance of Pteroptyx Fireflies in Rembau-Linggi Estuary, Peninsular Malaysia. EnvironmentAsia, 3 (special issue), 56–60.

    Jusoh, W.F.A., Hashim, N.R. & Wong, C.-H. (2011) Zonation of firefly species and their display trees along Kerteh River, Terengganu. Serangga, 16 (2), 59–66.

    Jusoh, W.F.A., Hashim, N.R., Sääksjärvi, I., Adam, N.A. & Wahlberg, N. (2014) Species delineation of Malaysian mangrove fireflies (Coleoptera: Lampyridae) using DNA barcodes. The Coleopterists Bulletin, 68 (4), 703–711.

    https://doi.org/10.1649/0010-065X-68.4.703

    Kallersjö, M., Farris, J.S., Kluge, A.G. & Bult, C. (1992) Skewness and permutation. Cladistics, 8, 275–287.
    https://doi.org/10.1111/j.1096-0031.1992.tb00071.x

    Källersjö, M., Farris, J.S. & Albert, V.A. (1999) Homoplasy increases phylogenetic structure. Cladistics, 15, 91–93.
    https://doi.org/10.1111/j.1096-0031.1999.tb00400.x

    Kawashima, I. (2003) Two new species of the lampyrid genus Pteroptyx Olivier (Coleoptera, Lampyridae, Luciolinae) from Sulawesi, Central Indonesia, with a list of the congeneric species. Special Bulletin of the Japanese Society of Coleopterology, 6, 263–274.

    Kumar, S., Tamura, K. & Nei, M. (2004) MEGA3: integrated software for molecular evolutionary genetics analysis & sequence alignment. Briefings in Bioinformatics, 5 (2), 150–163.
    https://doi.org/10.1093/bib/5.2.150

    Lambkin, C.L., Lee, M.S.Y., Winterton, S.L. & Yeates, D.K. (2002) Partitioned Bremer support and multiple trees. Cladistics, 18, 436–444.
    https://doi.org/10.1111/j.1096-0031.2002.tb00159.x

    Larson, A. & Dimmick, W.W. (1993) Phylogenetic relationships of the salamander families: An analysis of congruence among morphological and molecular characters. Herpetological Monographs, 7, 77–93.
    https://doi.org/10.2307/1466953

    Lewis, P.O. (2001) A likelihood approach to estimating phylogeny from discrete morphological character data. Systematic Biology, 50, 913–925.
    https://doi.org/10.1080/106351501753462876

    Lloyd, J.E. (1979) Sexual Selection in luminescent beetles. In: Blum, M.S. & Blum, N.A. (Eds.), Sexual Selection and Reproductive Competition in Insects. Academic Press, New York, pp. 293–342.
    https://doi.org/10.1016/B978-0-12-108750-0.50015-X

    Lloyd, J.E. (1983) Bioluminescence and Communication in insects. Annual Review of Entomology, 28, 131–160.
    https://doi.org/10.1146/annurev.en.28.010183.001023

    Lloyd, J.E. (1984) On deception, a way of all flesh, and firefly signalling and systematics. In: Dawkins, R. & Ridley, M. (Eds.), Oxford Survey of Evolutionary Biology. Vol. I. Oxford University Press, Oxford, pp. 48–84.

    Lloyd, J.E. & Wing, S.R. (1981) Photostory (copulation clamp). Florida Entomologist, 64, 459.

    Lloyd, J.E., Wing, S.R. & Hongtrakul, T. (1989) Ecology, flashes and behavior of congregating Thai fireflies. Biotropica, 21 (4), 373–376.
    https://doi.org/10.2307/2388290

    Lucas, R. (1920) Catalogus alphabeticus generum et subgenerum Coleopterorum orbis terrarum totius, Pars 1. Archiv fűr Naturgeschichte, 84, 557.

    Lunt, D.H., Zhang, D.X., Szymura, J.M. & Hewitt, O.M. (1996) The insect cytochrome oxidase I gene: evolutionary patterns and conserved primers for phylogenetic studies. Insect Molecular Biology, 5 (3), 153–165.
    https://doi.org/10.1111/j.1365-2583.1996.tb00049.x

    Maddison, W.P., Donoghue, M.J. & Maddison, D.R. (1984) Outgroup analysis and parsimony. Systematic Zoology, 33, 83–108.
    https://doi.org/10.2307/2413134

    Maddison, W.P. & Maddison, D.R. (2011) Mesquite: a modular system for evolutionary analysis. Version 2.75. Available from: http://mesquiteproject.org (accessed 1 December 2011)

    Mahadimenakbar, M.D., Hairul, H.M. & Mazidi A.G. (2007).The distribution and phenology of Pteroptyx fireflies (Coleoptera: Lampyridae) along Garama River, Sabah, Malaysia. Journal of Tropical Biology and Conservation, 3, 1–9.

    Mahadimenakbar, M.D. & Saikim, F.H. (2016) Studies on congregating fireflies (Coleoptera: Lampyridae: Pteroptyx sp.) on Sabah, Malaysia: A review. Journal of Tropical Biology and Conservation, 13, 13–25.

    McDermott, F.A. (1959) Two new species of fireflies (Coleoptera: Lampyridae). Entomological News, LXX (1), 7–12.

    McDermott, F.A. (1964) The taxonomy of the Lampyridae. Transactions of the American Entomological Society, 90, 1–72.

    McDermott, F.A. (1966) Lampyridae. In: Steel, W.O. (Ed.), Coleopterorum Catalogus Supplementa. Pars 9. Editio Secunda. W. Junk, S’Gravenhage, 149 pp.

    Margush, T. & McMorris, F. (1981) Consensus n-trees. Bulletin of Mathematical Biology, 43, 239–244.

    Meier, R. (1994) On the inappropriateness of presence/absence recoding for non-additive multistate characters in computerised cladistic analysis. Zoologischer Anzeiger, 232, 201–212.

    Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop (GCE). New Orleans, Louisiana, pp. 1–8.
    https://doi.org/10.1109/gce.2010.5676129

    Morse, E.S. (1916) Fireflies flashing in unison. Science, 43, 169–170.
    https://doi.org/10.1126/science.43.1101.169

    Motschulsky, V. de (1854) Lampyrides. Etudes Entomologiques, III, 47–62.

    Nada, B., Ballantyne, L.A., Kirton, L. & Norma-Rashid, Y. (2010) Discovery of an additional species of firefly along the Selangor River based on morphologically distinct larvae. Poster presentation. Lampyrid, 2, 201.

    Nallakumar, K. (2003) The synchronously flashing aggregative fireflies of Peninsular Malaysia. Biodiversity, 4 (2), 11–16.
    https://doi.org/10.1080/14888386.2003.9712684

    Nixon, K.C. & Carpenter, J.M. (1993) On outgroups. Cladistics, 9, 413–426.
    https://doi.org/10.1111/j.1096-0031.1993.tb00234.x

    Nurancha, P., Inkapatanakul, W. & Chunkao, K. (2013) Guidelines to the management of firefly watching tour in Thailand. Modern Applied Science, 7 (3), 8–14.
    https://doi.org/10.5539/mas.v7n3p8

    Ohba, N. & Sim, S.H. (1994) The morphology, behaviour and life cycle of Pteroptyx valida (Coleoptera: Lampyridae) in Singapore. Science Report of the Yokosuka City Museum, 42, 1–11.

    Ohba, N. & Wong, C.H. (2004) External morphology and ecological study of the firefly Pteroptyx tener at Kampong Kuantan, Selangor, Malaysia. Science Report of the Yokosuka City Museum, 51, 1–33.

    Ohba, N. & Wong, C.H. (2012) Observations on the external morphology and behaviour of Pteroptyx tener and Pteroptyx bearni in Malaysia. Lampyrid, 2, 80–86.

    Olivier, E. (1885) Catalogue des Lampyrides faisant partie des collections du Musée Civique de Gênes. Annali del Museo Civico di Storia Naturale di Genova, 2a (2), 333–374.

    Olivier, E. (1886) Baer, G.-A., Catalogue des Coléoptères des Iles Philippines. Annales de la Société Entomologique de France, 1886, 183–187.

    Olivier, E. (1902) Catalogue des espèces de 'Luciola' et genres voisins décrits jusqu'à ce jour. Revue Scientifique du Bourbonnais et du centre de la France, 15, 69–88.

    Olivier, E. (1907) Coleoptera, Lampyridae. Genera Insectorum, 53, 1–74. [Luciolinae, 48–56].

    Olivier, E. (1909a) Déscriptions de Lampyrides nouveaux du Musée Civique de Gênes. Annali del Museo Civico di Storia Naturale di Genova, 3a (IV), 316–319.

    Olivier, E. (1909b) Sur les Luciola d'Australie. Transactions of the Entomological Society of London, 1909, lxxxi–lxxxii.

    Olivier, E. (1910) Pars 9. Lampyridae. Coleopterorum Catalogus, 1910 (Luciolinae), 40–48.

    Olivier, E. (1911) Révision des Lampyrides. Revue Scientifique du Bourbonnais et du centre de la France, 24, 24–27, 39–58, 63–85, 98–112.

    Olivier, E. (1913) The Lampyridae of Borneo. Sarawak Museum Journal, 3, 55–60.

    Olivier, E. & Pic, M. (1909) Lampyride et Alleculide Nouveaux. L'Echange, 1909, 139–140.

    O'Reilly, J.E., Puttick, M.N., Parry, L., Tanner, A.R., Tarver, J.E., Fleming, J., Pisani, D. & Donoghue, P.C.J. (2016) Bayesian methods outperform parsimony but at the expense of precision in the estimation of phylogeny from discrete morphological data. Biology Letters, 12.
    https://doi.org/10.1098/rsbl.2016.0081

    Rambaut, A. & Drummond, A.J. (2012) FigTree: Tree figure drawing tool. Version 1.4.0. Available from: http://tree.bio.ed.ac.uk/ (accessed 21 May 2012)

    Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2013) Tracer Version 1.6. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 22 December 2013)

    Ronquist, F., Huelsenbeck, J.P. & Teslenko, M. (2011) MrBayes Version 3.2 Manual: Tutorials and Model Summaries. Available from: http://mrbayes.sourceforge.net/mb3.2_manual.pdf (accessed 20 November 2014)

    Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542.
    https://doi.org/10.1093/sysbio/sys029

    Smith, H.M. (1935) Synchronous flashing of fireflies. Science, 82, (2120), 151–152.

    https://doi.org/10.1126/science.82.2120.151

    Sorenson, M.D. & Franzosa, E.A. (2007) TreeRot. Version 3. Boston University, Boston, MA. [software]

    Sriboonlert, A., Swatdipong, A., Wonnapinij, P., E-Kobon, T. & Thancharoen, A. (2015) New record of Pteroptyx tener (Coleoptera: LAMPYRIDAE: Luciolinae) in Thailand. The Coleopterists Bulletin, 69 (2), 332–336.
    https://doi.org/10.1649/0010-065X-69.2.332

    Swofford, D.L. & Begle, D.P. (1993) PAUP: Phylogenetic Analysis Using Parsimony Version 3.1.1. User's Manual. Illinois Natural History Survey, Champaign, 257 pp.

    Swofford, D.L. (2002) PAUP*: Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4.0b.10. Sinauer Associates, Sunderland, Massachusetts. [software]

    Villalba, S., Lobo, J.M., Martín-Piera, F. & Zardoya, R. (2002) Phylogenetic relationships of Iberian dung beetles (Coleoptera: Scarabaeinae): insights on the evolution of nesting behavior. Journal of Molecular Evolution, 55 (1), 116–126.
    https://doi.org/10.1007/s00239-002-2314-4

    Wahlberg, N. & Wheat, C.W. (2008) Genomic outposts serve the phylogenomic pioneers: designing novel nuclear markers for genomic DNA extractions of Lepidoptera. Systematic Biology, 57, 231–242.
    https://doi.org/10.1080/10635150802033006

    Watson, J.G. (1928) Mangrove forests of the Malay Peninsula. No. 6. Fraser & Neave, Singapore, 275 pp.

    Wild, A.L. & Maddison, D.R. (2008) Evaluating nuclear protein-coding genes for phylogenetic utility in beetles. Molecular Phylogenetics and Evolution, 48 (3), 877–891.
    https://doi.org/10.1016/j.ympev.2008.05.023

    Will, K.W., Mishler, B.D. & Wheeler, Q.D. (2005) The perils of DNA barcoding and the need for integrative taxonomy. Systematic Biology, 54 (5), 844–851.
    https://doi.org/10.1080/10635150500354878

    Wing, S.R., Lloyd, J.E. & Hontrakul, T. (1983) Male competition in Pteroptyx fireflies: Wing-cover clamps, female anatomy, and mating plugs. Florida Entomologist, 66 (1), 86–91.
    https://doi.org/10.2307/3494553

    Wong, C.H. & Yeap, C.A. (2012) Conservation of Congregating Firefly Zones (CFZs) in Peninsular Malaysia. Lampyrid, 2, 174–187.

    Wright, A.M. & Hillis, D.M. (2014) Bayesian analysis using a simple likelihood model outperforms parsimony for estimation of phylogeny from discrete morphological data. PLoS ONE, 9, e109210.
    https://doi.org/10.1371/journal.pone.0109210

    Zaidi, M.I., Azman, S. & Wong, C.Y. (2005) Synchronous fireflies on berembang trees along Sungai Sepetang, Taiping, Perak. In: Shaharuddin, M.I., Azahar, M., Razani, U., Kamaruzaman, A.B., Lim, K.L., Suhaili, R., Jalil, M.S. & Latiff, A. (Eds.), Sustainable Management of Matang Mangroves: 100 Years and Beyond. Forestry Department Peninsular Malaysia, Kuala Lumpur, pp. 283–288.