ZooKeys 319: 293–301, doi: 10.3897/zookeys.319.4148
Madagascar Flatidae (Hemiptera, Fulgoromorpha): state-of-the-art and research challenges
Dariusz Świerczewski 1, Adam Stroiński 2
1 Department of Zoology and Animal Ecology, Jan Długosz University, Al. Armii Krajowej 13/15, 42-201 Częstochowa, Poland
2 Museum and Institute of Zoology PAS, Wilcza 64, 00-679 Warszawa, Poland

Corresponding author: Dariusz Świerczewski (dswier@ajd.czest.pl)

Academic editor: N. Simov

received 20 October 2012 | accepted 4 February 2013 | Published 30 July 2013


(C) 2013 Dariusz Świerczewski. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.


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Citation: Świerczewski D, Stroiński A (2013) Madagascar Flatidae (Hemiptera, Fulgoromorpha): state-of-the-art and research challenges. In: Popov A, Grozeva S, Simov N, Tasheva E (Eds) Advances in Hemipterology. ZooKeys 319: 293–301. doi: 10.3897/zookeys.319.4148

Abstract

The paper provides a historical review of the research on Flatidae in Madagascar and indicates future prospects. While the first two species of Madagascar Flatidae were described by Guérin-Méneville (1844), it was Signoret (1860) who made the first real attempt to enhance our knowledge of the Hemiptera fauna of Madagascar by describing several additional species. Over the following century and a half, several investigators have turned their attention to this group of insects, with the final number of species recorded for the island reaching 79. Despite this long history of research, it is evident that much still remains to be done. Detailed taxonomic research will allow the natural history of Madagascar and changes in the biological diversity of its endemic ecosystems to be better understood. This paper should be considered as an introduction to a complex study on the systematics and phylogeny of worldwide Flatidae planthoppers.

Keywords

Fulgoromorpha, Flatidae, taxonomy, research, history, Madagascar

Introduction

Flatidae constitute one of the largest families within planthoppers (Fulgoromorpha, Hemiptera) with 1446 species described in 299 genera and 12 tribes distributed worldwide (Bourgoin 2013). These phytophagous insects are highly diverse in terms of their colour and size (from 4.5 up to 32 mm), and are found on all continents, but are especially common and abundant in the tropics (O’Brien 2002). They are divided into two subfamilies – Flatinae and Flatoidinae, which, in most cases, can be easily distinguished from each other by the shape of the body. A greater number of Flatinae are flattened laterally, in contrast to Flatoidinae which hold their wings horizontally (O’Brien and Wilson 1985). About 20 species of Flatidae are regarded as serious pests of economically important crops such as coffee, tea, cacao, mango, citrus, apple and cherry (Wilson and O’Brien 1986). In Europe the only recognized pest is Metcalfa pruinosa (Say, 1830), a flatid species introduced into Italy from USA in 1973 (Arzone et al. 1987).

Data resources

The data underpinning the analyses reported in this paper are deposited in the Dryad Data Repository at doi: 10.5061/dryad.d43j8

Madagascar as a global biodiversity hotspot

Madagascar is one of eight important global biodiversity hotspots owing to its unique biota and the high level of threat to its natural habitats (McNeely et al. 1990, Myers et al. 2000, Ganzhorn et al. 2001). Being a part of the southern supercontinent Gondwana, it started to separate from Africa as a Madgascar-India block ca. 130 million years ago. At ca. 90 million years, India started breaking off from Madagascar and drifting northeastwards (Ali and Aitchison 2008). Despite Madagascar’s extreme isolation from India and its geographical proximity to Africa, a high proportion of the biota of the Madagascar region reveals some Asian affinities (Yoder and Nowak 2006). The suggested explanation of this might be the results of analyses obtained by Warren et al. (2010), which support the repeated existence of sizeable islands across the western Indian Ocean, reducing the distance of open ocean to be crossed.

Madagascar has evolved an incredible wealth of biodiversity, with thousands of species that can be found nowhere else on earth. On the one hand, this is due to its long isolation from all other landmasses (Storey et al. 1995), on the other, several alternative mechanisms have generated local endemism such as isolation within watersheds (Wilmé et al. 2006), adaptation along environmental gradients (Smith et al. 1997) and ecologically mediated postspeciation range shifts (Losos and Glor 2003). The relative importance of these mechanisms, both for Madagascar and globally, is still under discussion (Pearson and Raxworthy 2009, Vences et al. 2009). For example, of its estimated 12 000 plant species, nearly 10 000 are unique to Madagascar (Gautier and Goodman 2007). Among the fauna, vertebrates have been the most thoroughly studied Madagascan animals (Raxworthy and Nussbaum 1996, Raxworthy et al. 2002, Goodman and Ganzorn 2004). With regard to invertebrates, and insects in particular, the best known groups are butterflies (Torres et al. 2001, Zakharov et al. 2004), dung beetles (Wirta et al. 2008), minnow mayflies (Monaghan et al. 2005) and ants (Fisher 1997).

However, unsustainable use of natural resources such as wood for charcoal, bushmeat for protein supply and land for crop cultivations and cattle farming are having a profound impact on the Malagasy environment (Brown 2007). According to Kull (2012), an aerial photograph-based analysis of land-cover change of Madagascar’s high plateau in the latter half of the 20th century, based on a stratified rand sample of twenty eight sites, reveals a dramatic expansion of the cultural landscape of villages and agro-ecosystems into wetland and grassland environments. In response, significant environmental conservation efforts have been undertaken by the Malagasy government with international funding and technical support (Mittermeier et al. 2005). These environmental efforts in Madagascar have evolved over the past two decades from a Yellowstone model – transformation of large tracts of land into uninhabited, strictly protected areas, through the Integrated Conservation and Development Project (ICDP), which combines conservation of biological diversity with the social and economic needs of local people, to more recent initiatives that are the cutting edge of environmental innovation (Marcus and Kull 1999).

Flatidae research in Madagascar: a historical account

The Flatidae fauna of Madagascar has intrigued natural historians for centuries and continues to attract more interest with time. The first description of flatid species dates back to the first half of the 19th century when French entomologist Félix Édouard Guérin-Méneville (1799–1874) gave a drawing and a short note on Flata malgacha (presently in genus Flatida White, 1846) and Flatoides tortrix in the work Iconographie du Règne Animal (Guérin-Méneville 1844). However, it was Victor Signoret (1816–1889) who made the first real attempt to enhance our knowledge of the Hemiptera fauna of Madagascar (Signoret 1860) by describing several additional species, in particular: Nephesa antica, Nephesa suturalis, Phyllyphanta nivea, Elidiptera madagascariensis, Flatoides cicatricosus, Flatoides hyalinipennis, Flatoides sinuatus, Flatoides vicinus (presently in the genera Latois Stål, 1866, Flatopsis Melichar, 1902, Ulundia Distant, 1910 and Flatoidessa Melichar, 1923). Moreover, the second half of 19th century was the realase of further papers by such eminent naturalists-hemipterologists as Carl Stål (1833–1878) and Ferdinand Karsch (1853–1936), which provided descriptions of additional representatives of Madagascan Flatidae (Stål 1866, Karsch 1890). Leopold Melichar (1856–1924), the foremost worker on Hemiptera at that time, made a substantial contribution with his two-part monograph of the world fauna of Flatidae by publishing re-descriptions of a number of species and describing a large number of new taxa, including several restricted just to Madagascar (Melichar 1901, 1902). This work was followed by a more concise volume of Genera Insectorum (Melichar 1923) also dedicated in part to Flatidae. Further species were described by Victor Lallemand (1880–1965) and Jacques Auber (1916–1995) (Lallemand 1933, 1950, Auber 1954, 1955). In 1956, Belgian entomologist Henri Synave (1921–1980) gave a short overview of Madagascan Flatinae, providing a key to all known species and simplified illustrations of some genital characters (Synave 1956). He also published a faunistic paper based on material collected in Madagascar during an expedition organized by The Natural History Museum in Basel, which was in fact the last one dedicated to Madagascan flatids (Synave 1966). Although John T. Medler (1914–2006), an outstanding researcher on the world fauna of Flatidae, later published two papers on west and southern African flatids (Medler 1988, 2001), neither of them contained any data referring to Madagascar. Summarizing, the Madagascan flatid fauna presently consists of 18 genera with 40 species of Flatinae and 12 genera with 39 species of Flatoidinae (Metcalf 1957, Stroiński and Świerczewski 2011, Stroiński and Świerczewski 2012a).

Flatidae as a tool for biodiversity conservation: preliminary results

The accurate and rapid measurement of patterns of species richness, species turnover, and endemism is fundamental to current conservation efforts in Madagascar (Fisher 2007). One approach is to sample taxa that are ecologically important, but at the same time relatively easily collected, diverse at site and contain a high level of information for conservation planning. Flatidae meet these criteria and so might serve as an appropriate environmental indicator. According to Prof. Thierry Bourgoin from The Natural History Museum in Paris, this is the most abundant and easily collected group of the hemipterans in Madagascar (pers. comm.). Yet, the knowledge of the group is still very limited.

The wildlife of Madagascar is considered to be one of the most important human heritage resources of our time. The lack of appropriate tools for the recognition of biodiversity, which would also be useful in the protection of environment and natural resources, is a great disadvantage. Studies of Flatidae as a model group of phytophagous insects could be the solution for these needs, especially in relation to rare and endangered ecosystems. One of these is tapia woodlands – a short, endemic, sclerophyllous forest formation in Madagascar, with dominant canopy tree species Uapaca bojeri Baillon, 1874 or tapia in Malagasy. It can be found in four zones located in the central and southern part of the island, covering approximately 2600 km2 (Kull 2002). Tapia woodlands reveal strong adaptations to fire and are specifically human-shaped through controlled burning and selective cutting as they serve as a source of non-timber forest products for local residents (Kull 2003). The characteristic species for this ecosystem was recently described Phleboterum tapiae Świerczewski & Stroiński, 2012 (Świerczewski and Stroiński 2012b). Moreover, we discovered two new species Flatopsis medleri Świerczewski & Stroiński, 2011 and Latois nigrofasciata Świerczewski & Stroiński, 2012 confined exclusively to littoral forests (Świerczewski and Stroiński 2011, Świerczewski and Stroiński 2012a). Littoral forests have once formed an uninterrupted, narrow band along the entire length of Madagascar’s eastern coastline but as a result of anthropogenic activity their cover has been dramatically reduced to isolated, remnant fragments. Littoral forests are presently one of the smallest yet most diverse habitats in Madagascar. However, they are relatively unexplored and poorly documented ecosystems (De Gouvenain and Silander 2003). Approximately 13% of Madagascar’s total native flora can been found in these ecosystems and over 25% of the 1535 plant species known from littoral forests are endemic to this community (Consiglio et al. 2006). With respect to fauna, littoral forest are one of four major areas of ant endemism in Madagascar (Fisher and Girman 2000) Additionally, mountain flatid fauna seems quite rich and diverse, with endemic species and genera restricted to particular mountain massifs (work in prep.). An example can be the representatives of the genus Urana Melichar, 1902 – Urana paradoxa Melichar, 1902 and Urana unica Stroiński & Świerczewski, 2012, which are exclusively related to high altitude mountain rainforests (Stroiński and Świerczewski 2012b). Finally, we discovered the first representatives of the tribe Phantiini in Madagascar – Soares testudinarius Stroiński & Świerczewski, 2012 and Madoxychara unicornis Stroiński & Świerczewski, 2013 (Stroiński and Świerczewski 2012a, Stroiński and Świerczewski 2013). Phantiini is a small flatid tribe established by Melichar (1923) distributed worldwide and numbering 12 genera with 35 species. In Afrotropic it is represented by 6 genera and 11 species.

Future research perspectives

Summarizing, despite a long history of observation and investigation, the state of knowledge of Madagascar’s Flatidae is deeply unsatisfactory. After gathering material from the most important entomological collections and two years of study we realized how much still remains to be discerned. In the area of systematics our research efforts are focused on redescriptions of all Madagascan Flatidae based on type specimens, descriptions of new species, complete synonymy, identification keys for genera and species, documentation of research in the form of drawings, SEM images and maps and a distribution catalogue of the species. Reinterpretation of characters formerly used for the identification of species and distinguishing higher taxa (tegmen venation and male genital structures) will also be included. Additionally, as a novelty, we will provide analysis of Flatidae female genital structures.

With respect to biodiversity and endemism, the basic parameters of the distribution, dispersal and zoogeographic status have yet to be understood. Furthermore, distinguishing species characteristic for particular vegetational formations is key to understanding the dynamics of endangered ecosystems. The revision will make it possible to estimate the level of endemism of Madagascan flatid fauna and its connections with the fauna of Oriental and Afrotropical regions. The project presented above is the first step in a complex study on the systematics and phylogeny of worldwide Flatidae planthoppers. It aims to clear up views on the species variability and diversity, create a modern classification system, and know in detail all the world genera and species. The goal of the long-term project is to complete the work on cladistic analysis, phylogeny and the evolutionary scenario proposal for the family.

References
Ali JR, Aitchison JC (2008) Gondwana to Asia: Plate tectonics, paleogeography and the biological connectivity of the Indian sub-continent from the Middle Jurassic through latest Eocene (166–35 Ma). Earth-Science Review 88: 145-166. doi: 10.1016/j.earscirev.2008.01.007
Auber J (1954) Un noveau Phromnia de Madagascar (Hom. Flatidae). Bulletin de la Société Entomologique de France 59: 29-30.
Auber J (1955) Deux nouveaux Flatides malgaches (Homopt.). Bulletin de la Société Entomologique de France 60: 77-80.
Arzone A, Vidano C, Alma A (1987) Auchenorrhyncha introduced into Europe from the Nearctic region: taxonomic and phytopathological problems. In: Wilson MR, Nault LR (Eds). Proc. 2nd Int. Workshop on Leafhoppers and Planthoppers of Economic Importance, Provo, Utah, USA, 28th July-1st Aug. 1986, CIE, London: 3-17.
Baillon H (1874) Stirpes exoticae novae. Adansonia 11: 175-182.
Bourgoin Th (2013) FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to Fulgoromorpha. Version 8, updated [2013–01–23]. http://hemiptera-databases.org/flow/
Brown M (2007) The wealth of trees and the pressures of population. Madagascar Conservation & Development 2: 3.
Consiglio T, Schatz GE, McPherson G, Lowry II PP, Rabenantoandro J, Rogers ZS, Rabevohitra R, Rabehevitra D (2006) Deforestation and plant diversity of Madagascar’s Littoral Forests. Conservation Biology 20: 1799-1803. doi: 10.1111/j.1523-1739.2006.00562.x
De Gouvenain RC, Silander Jr JA (2003) Littoral Forest. In: Goodman SM, Benstead JP (Eds). The Natural History of Madagascar. The University of Chicago Press, Chicago-London: 103-109.
Distant WL (1910) Cercopidae concluded, Jassidae with additions to the Fulgoridae and many new genera and species. Insecta Transvaaliensia. A contribution to a knowledge of the entomology of South Africa 10: 229-252.
Fisher BL (1997) Biogeography and ecology of the ant fauna of Madagascar (Hymenoptera: Formicidae). Journal of Natural History 31 (2): 269-302. doi: 10.1080/00222939700770141
Fisher BL (2007) Formicidae, Ants. Role in Conservation. In: Goodman SM, Benstead JP (Eds). The Natural History of Madagascar. The University of Chicago Press, Chicago-London: 811-819.
Fisher BL, Girman DJ (2000) Biogeography of ants in eastern Madagascar. In: Lourenco WR, Goodman SM (Eds). Diversité et Endémisme à Madagascar. Mémoires de la Société de Biogéographie, Paris: 331-344.
Ganzhorn JU, Lowry II PP, Schatz GE, Sommer S (2001) The biodiversity of Madagascar: one the world’s hottest hotspots on its way out. Oryx 35 (4): 346-348.
Gautier L, Goodman SM (2007) Introduction to the Flora of Madagascar. In: Goodman SM, Benstead JP (Eds). The Natural History of Madagascar. The University of Chicago Press, Chicago-London: 229-250.
Goodman SM, Ganzhorn JU (2004) Biogeography of lemurs in the humid forests of Madagascar: the role of elevational distribution and rivers. Journal of Biogeography 31 (1): 47-55. doi: 10.1111/j.1365-2699.2004.00953.x
Guérin-Méneville FE (1844) Insectes. In: Cuvier GLCFD. Iconographie du Règne Animal, Paris, 355–370.
Karsch FAF (1890) Afrikanische Fulgoriden. Berliner Entomologische Zeitschrift 35: 57-70. doi: 10.1002/mmnd.18900350105
Kull ChA (2002) The “Degraded” Tapia Woodlands of Highland Madagascar: Rural Economy, fire Ecology, and Forest Conservation. Journal of Cultural Geography 19 (2): 95-128. doi: 10.1080/08873630209478290
Kull ChA (2003) Uapaca Woodlands. In: Goodman SM, Benstead JP (Eds). The Natural History of Madagascar. The University of Chicago Press, Chicago-London: 393-398.
Kull ChA (2012) Air photo evidence of historical land cover change in the highlands: Wetlands and grasslands give way to crops and woodlots. Madagascar Conservation & Development 7 (3): 144-152.
Lallemand V (1933) Homoptères Africains. Revue de Zoologie et de Botanique Africaines 24 (2): 198-199.
Lallemand V (1950) Contribution à l’étude des Homoptères de Madagascar. Mémoires de l’Institut Scientifique de Madagascar Serie A 4: 83-95.
Losos JB, Glor RE (2003) Phylogenetic comparative methods and the geography of speciation. Trends in Ecology and Evolution 18 (5): 220-227. doi: 10.1016/S0169-5347(03)00037-5
Marcus RR, Kull ChA (1999) Setting the Stage: The Politics of Madagascar’s Environmental Efforts. African Studies Quaterly 3 (2): 1-8.
McNeely JA, Miller KR, Reid WV, Mittermeier RA, Werner TB (1990) Conserving the World’s Biological Diversity. International Union for Conservation of Nature and Natural Resources, 193 pp.
Medler JT (1988) Flatidae from the Tai Forest, Cote d’Ivore, and taxonomic notes on the family in West Africa [Homoptera, Auchenorrhyncha, Fulgoroidea]. Revue Francaise d’Entomologie (N. S. ) 10: 117-148.
Medler JT (2001) Review of Flatidae in Southern Africa, with keys and descriptions of new species (Homoptera: Fulgoroidea). Contributions on Entomology-International 4: 323-375.
Melichar L (1901) Monographie der Acanaloniiden und Flatiden (Homoptera). Annalen des k.k Naturhistorischen Hofmuseums. Wien 16: 178-258.
Melichar L (1902) Monographie der Acanaloniiden und Flatiden (Homoptera) (Fortsetzung). Annalen des k.k Naturhistorischen Hofmuseums. Wien 17: 1-253.
Melichar L (1923) Homoptera, fam. Acanaloniidae, Flatidae et Ricaniidae. Genera Insectorum. Bruxelles 182: 1-185.
Metcalf ZP (1957) Flatidae and Hypochthonellidae. Part 13. General Catalogue of the Homoptera. Fascicule IV. North Carolina State College, Raleigh (USA), 565 pp.
Mittermeier RA, Hawkins F, Rajaobelina S, Langrand O (2005) Wilderness conservation in a biodiversity hotspot. International Journal of Wilderness 11 (3): 42-46.
Monaghan MT, Gattolliat J-L, Sartori M, Elouard J-M, James H, Derleth P, Glaizot O, de Moor F, Vogler AP (2005) Trans-oceanic and endemic origins of the small minnow mayflies (Ephemeroptera, Baetidae) of Madagascar. Proceedings of the Royal Society. Biological Sciences, Series B 272: 1829-1836.
Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GAB, Kent J (2000) Biodiversity hotspots for conservation priorities. Nature 403: 853-858. doi: 10.1038/35002501
O’Brien L (2002) The Wild Wonderful World of Fulgoromorpha. Denisia (NF) 4: 83-102.
O’Brien L, Wilson SW (1985) Planthopper Systematics and External Morphology. In: Nault LR, Rodriguez JG (Eds). The Leafhoppers and Planthoppers. John Wiley & Sons, New York: 61-102.
Pearson RG, Raxworthy ChJ (2009) The evolution of local endemism in Madagascar: watershed versus climatic gradient hypotheses evaluated by null biogeographic models. Evolution 63 (4): 959-967. doi: 10.1111/j.1558-5646.2008.00596.x
Raxworthy CJ, Nussbaum RA (1996) Montane amphibian and reptile communities in Madagascar. Conservation Biology 10 (3): 750-756. doi: 10.1046/j.1523-1739.1996.10030750.x
Raxworthy CJ, Forstner MRJ, Nussbaum RA (2002) Chameleon radiation by oceanic dispersal. Nature 415: 784-787. doi: 10.1038/415784a
Say T (1830) Descriptions of new North American Hemipterous Insects, belonging to the first family of the section Homoptera of Latreille. Journal of the Academy of Natural Sciences of Philadelphia 6: 235-244.
Signoret V (1860) Faune des hémiptères de Madagascar. 1ère partie. Homoptères. Annales de la Société Entomologique de France. Paris (Ser. 3) 8: 177–206.
Smith TB, Wayne RK, Girman DJ, Bruford MW (1997) A role for ecotones in generating rainforest biodiversity. Science 276: 1855-1857. doi: 10.1126/science.276.5320.1855
Stål C (1866) Hemiptera Homoptera Latr. Hemiptera Africana 4: 1-276.
Storey M, Mahoney JJ, Sounders AD, Duncan RA, Kelley SP, Coffin MF (1995) Timing of hot spot-related volcanism and the breakup of Madagascar and India. Science 267: 852-855. doi: 10.1126/science.276.5320.1855
Stroiński A, Świerczewski D (2011) A new flatid species of the genus Panormenis Melichar, 1923 from Madagascar (Hemiptera: Flatidae). Genus. Wrocław 22 (2): 191-203.
Stroiński A, Świerczewski D (2012a) Soares testudinarius gen. et sp. nov. (Hemiptera: Fulgoromorpha: Flatidae), a new representative of the tribe Phantiini from Madagascar. Zootaxa 3256: 38-50.
Stroiński A, Świerczewski D (2012b) Revision of an extraordinary Selizini genus Urana Melichar, 1902 from Madagascar (Hemiptera: Fulgoromorpha: Flatidae). Journal of Natural History 46(41–42): 2577-2593. doi: 10.1080/00222933.2012.70845
Stroiński A, Świerczewski D (2013) Madoxychara gen. nov. (Hemiptera: Fulgoromorpha: Flatidae), a new genus of the tribe Phantiini Melichar from Madagascar. Zootaxa 3599 (4): 377-389. doi: 10.11646/zootaxa.3599.4.6
Synave H (1956) Les Flatidae de Madagascar (Hemiptera-Homoptera). Mémoires de l’Institut Scientifique de Madagascar (Ser. E) 7: 197-217.
Synave H (1966) Homoptères de Madagascar. Verhandlungen der Naturforschenden Gesellschaft in Basel 77: 55-75.
Świerczewski D, Stroiński A (2011) Flatopsis medleri sp. n. – a new flatid species from Madagascar (Hemiptera: Fulgoromorpha: Flatidae). Acta zoologica cracoviensia 54B (1–2): 23–30.doi: 10.3409/azc.54b_1-2.23-30
Świerczewski D, Stroiński A (2012a) A new species of the genus Latois Stål, 1866 from Madagascar (Hemiptera: Fulgoromorpha: Flatidae). Acta zoologica cracoviensia 55 (1): 65-77. doi: 10.3409/azc.55_1.65
Świerczewski D, Stroiński A (2012b) A new species of Phlebopterum Stål, 1854 (Hemiptera: Fulgoromorpha: Flatidae) from the tapia woodlands of Madagascar. Annales Zoologici 62 (4): 577–592. doi: 10.3161/000345412X659641
Torres E, Lees DC, Vane-Wright RI, Kremen C, Leonard JA, Wayne RK (2001) Examining Monophyly in a Large Radiation of Madagascan Butterflies (Lepidoptera: Satyrinae: Mycalesina) Based on Mitochondrial DNA Data. Molecular Phylogenetics and Evolution 20 (3): 460-473. doi: 10.1006/mpev.2001.0984
Vences M, Wollenberg KC, Vieites DR, Lees DC (2009) Madagascar as a model region of species diversification. Trends in Ecology and Evolution 24 (8): 456-465. doi: 10.1006/mpev.2001.0984
Warren BH, Strasberg D, Bruggemann JH, Prys-Jones RP, Thébaud Ch (2010) Why does the biota of the Madagascar region have such a strong Asiatic flavour? Cladistics 26 (5): 526–538. doi: 10.1111/j.1096-0031.2009.00300.x
White A (1846) Descriptions of some apparently new species of Orthopterous and Homopterous insects. Annals and Magazine of Natural History. London 18: 23-26.
Wilmé L, Goodman SM, Ganzhorn JU (2006) Biogeographic Evolution of Madagascar’s Microendemic Biota. Science 312: 1063-1065. doi: 10.1126/science.1122806
Wilson SW, O’Brien LB (1986) A Survey of planthopper pests of economically important plants (Homoptera: Fulgoroidea). In: Wilson MR, Nault LR (Eds). Proceedings of 2nd International Workshop on Leafhoppers and Planthoppers of Economic Importance, Provo, Utah, USA, 28th July-1st August 1986. CIE, London: 343-360.
Wirta H, Orsini L, Hanskii I (2008) An old adaptive radiation of forest dung beetles in Madagascar. Molecular Phylogenetics and Evolution 47 (3): 1076-1089. doi: 10.1016/j.ympev.2008.03.010
Yoder AD, Nowak MD (2006) Has vicariance or dispersal been the predominant biogeographic force in Madagascar? Only time will tell. Annual Review of Ecology, Evolution and Systematics 37: 405-431. doi: 10.1146/annurev.ecolsys.37.091305.110239
Zakharov EV, Smith CR, Lees DC, Cameron A, Vane-Wright RI, Sperling FAH (2004) Independent gene phylogenies and morphology demonstrate a Malagasy origin for a wide-ranging group of swallowtail butterflies. Evolution 58 (12): 2763-2782.