Systematics and species radiation of the sheoak lace bug genus Epimixia Kirkaldy (Insecta : Heteroptera : Tingidae) in Australia, New Caledonia and Papua New Guinea
Gerasimos Cassis A B D , Celia Symonds A and Lauren Branson CA Evolution & Ecology Research Centre, University of New South Wales, Sydney, NSW 2052, Australia.
B Research Associate, 1 College Street, Australian Museum, Sydney, NSW 2000, Australia.
C Independent researcher.
D Corresponding author. Email: gcassis@unsw.edu.au
Invertebrate Systematics 33(2) 277-366 https://doi.org/10.1071/IS18040
Submitted: 22 May 2018 Accepted: 26 July 2018 Published: 4 April 2019
Abstract
On the basis of new survey, knowledge of Australia’s heteropteran fauna has been shown to have a significant taxonomic impediment, requiring the description of many new lineages. In this work we provide a monographic treatment of the lace bug genus Epimixia, including a phylogenetic analysis validating its monophyly. We redescribe the genus and previously described species E. alitophrosyne Kirkaldy, E. nigriceps (Signoret), E. nigripes dysmica Drake & Ruhoff, E. nigripes nigripes (Horváth), E. veteris Drake, E. vittata Horváth and E. vulturna (Kirkaldy). Eighteen new species are described, including the first recorded species from Papua New Guinea (E. gagnei, sp. nov.), a second species from New Caledonia (E. fulva, sp. nov.) and 16 new Australian species (E. aboccidente, sp. nov., E. acclivis, sp. nov., E. albimons, sp. nov., E. allocasuarina, sp. nov., E. bicolor, sp. nov., E. caerulamons, sp. nov., E. christopherdarwini, sp. nov., E. eneabba, sp. nov., E. kimberley, sp. nov., E. leai, sp. nov., E. megacosta, sp. nov., E. minor, sp. nov., E. pilbara, sp. nov., E. schuhi, sp. nov., E. tropica, sp. nov. and E. verticordiaphila, sp. nov.). Epimixia dysmica Drake & Ruhoff is elevated to species rank, from its previous subspeciefic status within E. nigripes. This work also reports host plants for Epimixia, predominantly belonging to the Casuarinaceae, with a handful of species associated with the Myrtaceae, Proteaceae and Fabaceae. Host plants are mapped to the Epimixia phylogeny, indicating that the Casuarinaceae-affiliated Epimixia species form a subclade. Area relationships of the Epimixia species are analysed using subtree analysis, resulting in the south-west and south-east corners of Australia being sisters, relative to the Adelaide subregion, and in turn to Atherton + New Caledonia.
http://zoobank.org/urn:lsid:zoobank.org:pub:E7FC4E37-517D-4AB7-A918-B9104D63AB25
Additional keywords: biogeography, host plants, phylogeny, taxonomy.
References
Bagils, R. Z., Ung, V., Grand, A., Vignes-Lebbe, R., Cao, N., and Ducasse, J. (2012). LisBeth: new cladistics for phylogenetics and biogeography. Comptes Rendus. Palévol 11, 563–566.| LisBeth: new cladistics for phylogenetics and biogeography.Crossref | GoogleScholarGoogle Scholar |
Cassis, G. (2008). The Lattinova complex of Austromirine plant bugs (Hemiptera: Heteroptera: Miridae: Orthotylinae). Proceedings of the Entomological Society of Washington 110, 845–939.
| The Lattinova complex of Austromirine plant bugs (Hemiptera: Heteroptera: Miridae: Orthotylinae).Crossref | GoogleScholarGoogle Scholar |
Cassis, B., and Bulbert, M. (2016). Lacebugs of Australia. LUCID key at Lucidcentral. http://keys.lucidcentral.org/keys/v3/aus_museum/lace_bugs/
Cassis, G., and Gross, G. F. (1995). ‘Zoological Catalogue of Australia. Hemiptera: Heteroptera (Coleorrhyncha to Cimicomorpha). Volume 27.3A.’ (CSIRO Publising: Melbourne.)
Cassis, G., and Gross, G. (2002). ‘Zoological Catalogue of Australia. Hemiptera: Heteroptera (Pentatomomorpha). Volume 27.3B.’ (CSIRO Publishing: Melbourne.)
Cassis, G., and Laffan, S. (2014). ‘Final Report on Heteroptera Integration and Spatial Analysis – Quantifying Bush Blitz: Heteroptera.’ (University of New South Wales: Sydney.)
Cassis, G., and Schuh, R. T. (2012). Systematics, biodiversity, biogeography, and host associations of the Miridae (Insecta: Hemiptera: Heteroptera: Cimicomorpha). Annual Review of Entomology 57, 377–404.
| Systematics, biodiversity, biogeography, and host associations of the Miridae (Insecta: Hemiptera: Heteroptera: Cimicomorpha).Crossref | GoogleScholarGoogle Scholar | 22149267PubMed |
Cassis, G., and Symonds, C. (2008). Systematics, biogeography and host associations of the lace bug genus Inoma (Hemiptera: Heteroptera: Tingidae). Acta Entomologica Musei Nationalis Pragae 48, 433–484.
Cassis, G., and Symonds, C. (2011). Systematics, biogeography and host plant associations of the lace bug genus Lasiacantha Stål in Australia (Insecta: Hemiptera: Heteroptera: Tingidae). Zootaxa 2818, 1–63.
Cassis, G., and Symonds, C. (2014). Systematics and host plant associations of a new genus of Acacia-inhabiting plant bugs from arid Australia (Insecta: Hemiptera: Heteroptera: Miridae: Orthotylinae). Invertebrate Systematics 28, 522–554.
| Systematics and host plant associations of a new genus of Acacia-inhabiting plant bugs from arid Australia (Insecta: Hemiptera: Heteroptera: Miridae: Orthotylinae).Crossref | GoogleScholarGoogle Scholar |
Cassis, G., and Symonds, C. (2016). Plant bugs, plant interactions and the radiation of a species rich clade in south-western Australia: Naranjakotta, gen. nov. and eighteen new species (Insecta: Heteroptera: Miridae: Orthotylinae). Invertebrate Systematics 30, 95–186.
| Plant bugs, plant interactions and the radiation of a species rich clade in south-western Australia: Naranjakotta, gen. nov. and eighteen new species (Insecta: Heteroptera: Miridae: Orthotylinae).Crossref | GoogleScholarGoogle Scholar |
Cassis, G., Wall, M. E., and Schuh, R. T. (2007). Biodiversity and industrializing the taxonomic process: the plant bug case study (Insecta: Heteroptera: Miridae). In ‘Reconstructing the Tree of Life: Taxonomy and Systematics of Species Rich Taxa’. (Eds T. R. Hodkinson, J. Parnell and S. Waldren.) pp. 193–212. (CRC Press: Boca Raton, FL.)
Cassis, G., Koenig, P., Symonds, C., and Shofner, R. (2017). Systematics and host plant associations of the Australian lace bug genus Nethersia (Insecta: Heteroptera: Tingidae), including the description of eighteen new species. Insect Systematics & Evolution 48, 1–95.
| Systematics and host plant associations of the Australian lace bug genus Nethersia (Insecta: Heteroptera: Tingidae), including the description of eighteen new species.Crossref | GoogleScholarGoogle Scholar |
Drake, C. J. (1944). A new genus and ten new species of serenthiines (Hemiptera: Tingitidae). Proceedings of the Entomological Society of Washington 46, 67–76.
Drake, C. J. (1961). Some Australian Tingidae (Hemiptera), including new genera and new species. Records of the Australian Museum 25, 107–113, pls III–IV.
Drake, C. J., and Ruhoff, F. A. (1960). Lace-bug genera of the world (Hemiptera: Tingidae). Proceedings of the United States National Museum 112(3431), 1–105, pls I–IX.
Drake, C. J., and Ruhoff, F. A. (1961). The genus Epimixia Kirkaldy (Hemiptera: Tingidae). Transactions of the Royal Society of New Zealand (Zoology) 1, 1–6, 2 pls.
Drake C. J. Ruhoff F. A. 1965
Ebach, M. C., Murphy, D. J., Gonzalez-Orozco, C. E., and Miller, J. T. (2015). A revised area taxonomy of phytogeographical regions within the Australian Bioregionalisation Atlas. Phytotaxa 208, 261–277.
| A revised area taxonomy of phytogeographical regions within the Australian Bioregionalisation Atlas.Crossref | GoogleScholarGoogle Scholar |
Goloboff, P. A., Farris, J. S., and Nixon, K. C. (2008). TNT, a free program for phylogenetic analysis. Cladistics 24, 774–786.
| TNT, a free program for phylogenetic analysis.Crossref | GoogleScholarGoogle Scholar |
González-Orozco, C. E., Ebach, M. C., Laffan, S., Thornhill, A. H., Knerr, N. J., Schmidt-Lebuhn, A. N., Cargill, C. C., Clements, M., Nagalingum, N. S., Mishler, B. D., and Miller, J. T. (2014). Quantifying phytogeographical regions of Australia using geospatial turnover in species composition. PlosOne 9, e92558.
| Quantifying phytogeographical regions of Australia using geospatial turnover in species composition.Crossref | GoogleScholarGoogle Scholar |
Guilbert, E. (2004). Immature stages of New Caledonian Tingidae (Heteroptera): description and development. European Journal of Entomology 101, 261–271.
| Immature stages of New Caledonian Tingidae (Heteroptera): description and development.Crossref | GoogleScholarGoogle Scholar |
Guilbert, E. (2008). New Caledonian Tingidae (Insecta: Heteroptera): new species and new records. In ‘Zoologia Neocaledonica 6. Biodiversity Sstudies in New Caledonia’. (Ed. P. Grandcolas.) Mémoires du Museum National de Histoire Naturelle 197, 63–78.
Guilbert, E., and Moir, M. L. (2010). A new species of Carldrakeana (Insecta: Heteroptera: Tingidae) from Western Australia. Records of the Western Australian Museum 25, 382–386.
| A new species of Carldrakeana (Insecta: Heteroptera: Tingidae) from Western Australia.Crossref | GoogleScholarGoogle Scholar |
Gustafsson, B. (2006). A checklist of the Heteroptera in the Swedish Museum of Natural History. In ‘Entomology Collections. Vol. 2010’. (Naturhistoriska Riksmuseet: Stockholm, Sweden.)
Hacker, H. (1927). New Tingitoidea (Hemiptera) in the Queensland Museum. Memoirs of the Queensland Museum 9, 19–32, pls VI–X.
Hacker, H. (1928). New species and records of Australian Tingitoidea (Hemiptera). Memoirs of the Queensland Museum 9, 174–188.
Hacker, H. (1929). New species of Australian Tingitidae (Hemiptera). Memoirs of the Queensland Museum 9, 324–334, pls XXXII–XXXV.
Horváth, G. (1925). Results of Dr E. Mjöberg’s Swedish Scientific Expeditions to Australia 1910–1913. 45. Tingitidae. Arkiv för Zoologi 17A, 1–17.
Kirkaldy, G. W. (1908). Memoir on a few heteropterous Hemiptera from eastern Australia. Proceedings of the Linnean Society of New South Wales 32, 768–788, pl. xliii.
Kozub, D., Khmelik, V., Shapoval, J., Chentsov, V., and Yatsenko, S. (2008). ‘Helicon Focus Pro.’ (Helicon Soft: Kharkov, Ukraine.)
Lethierry, L., and Severin, G. (1896). ‘Catalogue Général des Hémiptères. Tome III. Hétéroptères Tingidae, Phymatidae, Aradidae, Hebridae, Hydrometridae, Henicocephalidae, Reduvidae, Saldidae, Apophilidae, Ceratocombidae, Cimicidae, Anthocoridae.’ (R. Friedländer & Fils: Berlin, Germany.)
Lis, B. (1997). Australocader kerzhneri gen. n., sp. n. (Heteroptera: Tingidae: Cantacaderinae) from Australia. Polskie Pismo Entomologiczne 66, 3–4.
Lis, B. (2000). Carldrakeana pallida, a new species of Australian Cantacaderidae (Hemiptera: Heteroptera: Tingoidea). Polskie Pismo Entomologiczne 69, 405–410.
Lis, B. (2001). Australocader secundus, a new Australian species of Cantacaderidae (Hemiptera: Heteroptera: Tingoidea). Polskie Pismo Entomologiczne 70, 2.
Moir, M. L. (2009). Two new species of the lace bug genus Radinacantha (Hemiptera: Heteroptera: Tingidae) from Australia. Records of the Western Australian Museum 25, 269–275.
| Two new species of the lace bug genus Radinacantha (Hemiptera: Heteroptera: Tingidae) from Australia.Crossref | GoogleScholarGoogle Scholar |
Moir, M. L., and Guilbert, E. (2012). Swaustraltingis isobellae, a new genus and new species of Australian lacebug (Insecta: Heteroptera: Tingidae), with a redescription of Cysteochila cracentis Drake, and notes on the lacebug fauna of south-west Australia. Australian Journal of Entomology 51, 258–265.
| Swaustraltingis isobellae, a new genus and new species of Australian lacebug (Insecta: Heteroptera: Tingidae), with a redescription of Cysteochila cracentis Drake, and notes on the lacebug fauna of south-west Australia.Crossref | GoogleScholarGoogle Scholar |
Moir, M. L., and Lis, B. (2012). Description of three new species of Ceratocader (Hemiptera: Heteroptera: Tingidae) from Western Australia. Records of the Western Australian Museum 27, 148–155.
| Description of three new species of Ceratocader (Hemiptera: Heteroptera: Tingidae) from Western Australia.Crossref | GoogleScholarGoogle Scholar |
Nixon, K. C. (1999–2002). ‘WinClada Ver. 1.0000.’ (Published by the author: Ithaca, NY.)
Noack, A., Cassis, G., and Rose, H. A. (2011). Systematic revision of Thaumastocoris Kirkaldy (Hemiptera: Heteroptera: Thaumastocoridae). Zootaxa 3121, 1–60.
Preece, M., Harding, J., and West, J. G. (2015). Bush Blitz: journeys of discovery in the Australian outback. Australian Systematic Botany 27, 325–332.
| Bush Blitz: journeys of discovery in the Australian outback.Crossref | GoogleScholarGoogle Scholar |
Schuh, R. T. (2012). Integrating specimen databases and revisionary systematics. ZooKeys 209, 255–267.
| Integrating specimen databases and revisionary systematics.Crossref | GoogleScholarGoogle Scholar |
Schuh, R. T. (2002–2013). Plant Bugs Inventory website [Online]. Available at: http://research.amnh.org/pbi/ [Verified 1 May 2018].
Signoret, V. (1881). [no title]. Bulletin de Societe de Entomogique de. France 6, L.
Slater, J. A., Schuh, R. T., Cassis, G., Johnson, C. A., and Pedraza-Peñalosa, P. (2009). Revision of Laryngodus Herrich-Schaeffer, an Allocasuarina feeder, with comments on its biology and the classification of the family (Heteroptera: Lygaeoidea: Rhyparochromidae). Invertebrate Systematics 23, 111–133.
| Revision of Laryngodus Herrich-Schaeffer, an Allocasuarina feeder, with comments on its biology and the classification of the family (Heteroptera: Lygaeoidea: Rhyparochromidae).Crossref | GoogleScholarGoogle Scholar |
Symonds, C., and Cassis, G. (2013). New species of the lace bug genus Lasiacantha Stål (Insecta: Hemiptera: Heteroptera: Tingidae) from Western Australia, including a revised key to all Australian species. Australian Journal of Entomology 52, 53–66.
| New species of the lace bug genus Lasiacantha Stål (Insecta: Hemiptera: Heteroptera: Tingidae) from Western Australia, including a revised key to all Australian species.Crossref | GoogleScholarGoogle Scholar |
Symonds, C. L., and Cassis, G. (2014). A new genus Ittolemma (Heteroptera: Tingidae) gen. nov. and three included species of hirsute lace bugs from temperate woodlands of southern Australia. Austral Entomology 53, 380–390.
| A new genus Ittolemma (Heteroptera: Tingidae) gen. nov. and three included species of hirsute lace bugs from temperate woodlands of southern Australia.Crossref | GoogleScholarGoogle Scholar |