The cicadas (Hemiptera: Cicadidae) of India, Bangladesh, Bhutan, Myanmar, Nepal and Sri Lanka: an annotated provisional catalogue, regional checklist and bibliography

Abstract Background The cicadas of the Indian subcontinent, like many other insects in the region, have remained understudied since the early part of the 20th Century, and await modern taxonomic, systematic and phylogenetic treatment. This paper presents an updated systematic catalogue of cicadas (Hemiptera: Cicadidae) from India, Bangladesh, Bhutan, Myanmar, Nepal and Sri Lanka, the first in over a century. New information This paper treats 281 species, including: India and Bangladesh (189 species), Bhutan (19 species), Myanmar (81 species), Nepal (46 species) and Sri Lanka (22 species). For each species all recognized junior synonyms are included with information on the type material and additional specimens where relevant. The global distributional range and notes on the taxonomy of each species are included where appropriate. Two lists are provided: (1) species known to occur in India and Bangladesh (treated as a geographic unit), Bhutan, Myanmar, Nepal and Sri Lanka; and (2) species previously listed from these countries in error. A bibliography of species descriptions is provided, with the papers containing the original descriptions provided where copyright allows.


Introduction
We share our planet with an estimated 8.7 million species (Mora et al. 2011), of which approximately 80% await scientific description. Insects, although understudied, are known to account for over half the world's described biodiversity (Foottit and Adler 2009), and likely constitute a much greater proportion of the undescribed biodiversity. The cicadas (Hemiptera: Cicadidae) of the Indian subcontinent, like many other insects in the region, have remained understudied since the early part of the 20th Century and await, for the most part, modern taxonomic, systematic and phylogenetic treatment.
The first illustrated publication dealing with the insect fauna of India was "An epitome of the natural history of the insects of India and the islands in the Indian Seas" by Donovan (1800), which included many superb illustrations made by the author. One cicada species (Cicada indica Donovan 1800) was described by Donovan in his epitome, although the illustration is dated as 1st of February 1804 (Fig. 1). The original description states that the specimen was collected in Bengal, a claim doubted by Distant: "According to Donovan, a single specimen of this species was found in Bengal by Mr. Fichtel, and deposited in the Imperial Cabinet at Vienna, but that habitat I consider liable to the greatest doubt." (Distant 1889a). Donovan may have confused Fabricius' label "Indiis" as India, when it could refer to the West Indies, East Indies or occasionally to Africa. In the present study Cicada indica is listed as unlikely to be from the Indian subcontinent, being synonymised with Tacua speciosa speciosa (Illiger, 1800) which is found in Indonesia and Malaysia.
By far the most influential figure in the history of cicadas in India is William Lucas Distant (12 November 1845-4 February 1922. Distant worked at the Natural History Museum, then the British Museum (Natural History) from 1889 to 1920, during which time he described a tremendous number of insect species. Over 100 cicada species which occur on the Indian subcontinent and were described by Distant are currently recognised as valid. His most significant works relevant to this fauna were: "A Monograph of Oriental Cicadidae" (Distant 1889b, Distant 1891, Distant 1892c; "The fauna of British India, including Ceylon and Burma" (Distant 1906c); and "A Synonymic Catalogue of Homoptera, Part I -Cicadidae" (Distant 1906b). In particular "The fauna of British India, including Ceylon and Burma" (Distant 1906c) provided the first treatment of the region as a whole, with 148 cicada species recognised in 44 genera. This was followed by an appendix (Distant 1916) which listed an additional 24 species and 3 new genera for the region, bringing the total to 172 cicada species in 47 genera.
Apart from a few isolated species descriptions, the cicada fauna of the Indian subcontinent has not been subjected to a detailed taxonomic and systematic review since Distant (1906c), Distant (1916), hence, it is highly likely that many unrecognized and undescribed cicada species exist in this region.
This study serves as a modern reference point for the cicada fauna of India, Bangladesh, Bhutan, Myanmar, Nepal and Sri Lanka, and hopefully, provides a backbone to recording efforts and studies of the underlying taxonomy. Pakistan, formerly a part of "British India" has been excluded from full review in this catalogue as it has recently been updated by Ahmed and Sanborn (2010), however the fauna are briefly compared. Donovan's illustration of Cicada indica Donovan 1800. This stunning species is a junior synonym of Tacua speciosa speciosa (Illiger, 1800) and is not found on the Indian subcontinent, an example of the historical misattribution of species from the "East Indies" with India.
The checklists presented here are full of compromises but our guiding intention has been to present all the relevant information in a manner that is as compact and informative as possible. No doubt there are errors which have crept in, or which have not been discovered in the previous literature, and we welcome all suggestions to rectify any errors or omissions. To aide future studies the next incarnation will be digital, and the list of species is already in the process of migrating to a website, where updates will be maintained as and when changes occur. The website will also allow the community to share images and information on each species, providing an up-to-date resource, hopefully for many years to come.
Historically the regions making up the Indian subcontinent have had a complex nomenclature (Figs 2, 3), further compounded by different authors interpreting locality labels to mean different regions. In particular the confusion between East India, East Indies, India, Indo-China and Indonesia, will mean that some species included in this list are unlikely to occur in the focal region, but are included until further study proves otherwise.

Checklists
The checklist was based on the "The fauna of British India, including Ceylon and Burma" (Distant 1906c) and its supplement (Distant 1916), the four catalogues which list the global cicada literature (Distant 1906b, Metcalf 1963, Duffels and van der Laan 1985, Sanborn 2014) and the material housed at the Natural History Museum, London. In addition species which have been described after 2010 were incorporated by searching Google Scholar (htt ps://scholar.google.co.uk/) and ScienceDirect (http://www.sciencedirect.com/) using a combination of "India", "Bangladesh", "Bhutan", "Myanmar", "Nepal", "Sri Lanka", and "cicada" and "Cicadidae". The checklist is based on either the locality of the type specimen, the locality of specimens whose identity has been verified or substantial literature evidence. This checklist follows the higher taxonomy of Sanborn (2014) with updated generic combinations including current information as of September 2015. Institutional abbreviations are summarized in Table 1. Map of the Indian subcontinent, including countries as politically delineated at present that were at one time considered part of "British India" and countries surrounding "British India" with which the fauna of Bangladesh, Bhutan, India, Myanmar, Nepal, Pakistan and to a lesser extent Sri Lanka likely overlaps.
The cicadas (Hemiptera: Cicadidae) of India, Bangladesh, Bhutan, Myanmar, ... Due to the significant gap since the last major reviews by Distant, and the dramatic changes in the geographic boundaries of the countries involved (Figs 2, 3), two lists have been developed: (1) Species present in India and Bangladesh (treated here as a single unit), Bhutan, Myanmar, Nepal and Sri Lanka.
(2) Species historically included in the countries listed above, but on analysis are very unlikely to be present in these countries, with reasons for their exclusion listed in the notes.
We have not considered the cicada fauna of Pakistan as these species have recently been studied (Ahmed and Sanborn 2010).

Specimen Images
All type specimens examined were photographed (dorsal and ventral) and are accessible at: http://www.indiancicadas.org/.

Bibliography
For each species recorded from the Indian subcontinent the original description was sourced. In addition, any publication listing the focal region (as summarized by the four major catalogues) was sourced and confirmed. The citation of each authority is linked in this manuscript through the Notes field and in addition is publically accessible on Mendeley as the "Cicadas of the Indian subcontinent": http://mnd.ly/1OFGZc1. Original species descriptions without copyright restrictions can be downloaded from the NHM data portal: ht tp://dx.doi.org/10.5519/0028307.

Data resources
The bibliography is publically accessible on Mendeley: http://mnd.ly/1OFGZc1 The species descriptions are available on the NHM data portal: http:// dx.doi.org/10.5519/0028307 The checklist files for each individual country are available on the NHM data portal: http:// dx.doi.org/10.5519/0028307 The above files and information, along with images of specimens examined are also available on the Cicadas of India website: http://www.indiancicadas.org/ under "Balinta sp" which may be the other syntype, however this requires additional study.
Notes: Authority: Distant 1892c; Lectotype was designated by Beuk (1996). There are an additional 3 male (2 MSNG; 1 BMNH) and 1 female (MSNG) paralectotypes, however only the examined material is included in this paper. Not from India: Sanborn (2014) states India in reference to Hua (2000) but no other records from India exist, it is likely the result of a mistranslation of Indo-China. The type locality (Carin Asciuii Ghecu, Tennasserim) is not near the India/Myanmar border.
Notes: Authority: Distant 1888c; Distant (1906c) states that the type specimen is male.
Notes: Authority: Distant 1897; Metcalf (1963) listed India in reference to Mathur (1953) in which Burma was considerd a part of "British India". The type locality (Chin Hills) extends into India (Lushai Hills -Nagaland), thus this species may be recorded in Nagaland in future.
Notes: Authority: Distant 1905a; Not from India: Metcalf (1963) listed India in reference to Mathur (1953), in which it references a specimen "India.-Burma: Tavoy" with Burma being considerd part of "British India". The type locality (Thaungyang Valley, Tennasserim) and Mathur's locality Tavoy (now Dawei) are both in the far south of Myanmar thus this species has been removed from the Indian list. Subsequent "India" localities as stated in Sanborn (2014), in reference to Boulard (2001b), Sanborn et al. (2007) and Lee (2008), are likely to have resulted from this initial error. Hayashi (1993) excluded this species from India.
Notes: Authority: Distant 1902; Sanborn (2014) listed locality including Sikkim in reference to Lei (1994). This may be in error as the species has been recorded from Sri Lanka previously.
Notes: Authority: Distant 1913b; Female described by Boulard (2005). Sanborn (2014) listed India in reference to Boulard (2005). This species was described from Indo-China and there are currently no records of this species known from the Indian region, however it is present in the surrounding countries. Distribution: [Metcalf, 1963]  Notes: Authority: Distant 1888c; In the NHMUK there is a male specimen bearing a type label, however this male was described later by Distant (1913b). Holotype is a female from "Tenasserim: Houngdarau Valley" (MSNG). Not from India: Sanborn (2014) states India in reference to Sanborn et al. (2007) and Lee (2008), however their inclusion of India is in reference to previous distributions that may be incorrect, having resulted from Indo-China being included as India. Further evidence is required before this species can be included in the fauna of this region.
Notes: Authority: Distant 1879; Metcalf (1963) lists "India", however the type was collected from Upper Tenasserim (southern Myanmar), which was formerly a part of "British India" but is quite distant from the India/Myanmar border. No records have been found in modern India at present.
Notes: Authority: Walker 1858a; Species description states type specimen is male but the labelled type specimen in NHMUK is female. This observation agrees with Distant (1892a) which illustrates type specimen as a female. Notes: Authority: Walker 1858b; Metcalf (1963) states "Eastern India, India, Assam" in reference to India orientalis. Lee (2010b) states that the type locality is Myanmar and that records from China, Philippines, Indonesia, Malaysia, and Thailand may have been based on mididentifications of closely allied species. Subsequent references to India in Sanborn (2014) are likely to be due to previous inaccurate literature localities. Distribution: [Metcalf, 1963] India; Madras. [Sanborn, 2014] India.
Notes: Authority: Naruse in Naruse and Takagi 1977; An additional 1 male paratype (EIHU) was designated in the species description.
Notes: Authority : Ollenbach 1929;Metcalf (1963) listed India in reference Burma being part of "British India". The type locality (Thandaung) is within modern day Myanmar and the species has not yet been recorded from modern day India.
Notes: Authority: Walker 1850; Not from India: Metcalf (1963) incorrectly stated India in reference to Dohrn (1859) who listed it with an unknown locality. The species was described by Walker (1850) without a known locality. The type specimen is not in the NHMUK and this species has not been mentioned in literature since Distant (1906a). This species was not included in the Fauna of British India by Distant (1906c). Distribution: [Metcalf, 1963] Europe. [Sanborn, 2014] Metcalf (1963) stated India in reference to the type specimen location that was listed as "Indiis". Duffels (1991) states that this species is from Indonesia, and that the Indian listing by Metcalf is eronious. Thus subsequent references to India in Sanborn (2014) are likely to be due to previous inaccurate literature localities.
Notes: Authority: Matsumura 1917; Unlikely to be from India: Incorrectly listed in India and Burma by Schumacher (1915). According to Metcalf (1963), Schumacher referred to a specimen misidentified as Pycna repanda (actually Platypleura takasagona) and likely stated the locality information associated with P. repanda and not P. takasagona.
Notes: Authority: Illiger 1800; Unlikely to be from India: Metcalf (1963), among others, states India in reference to the type locality of Cicada indica Donovan, 1800 which is a junior synonym of Tacua speciosa speciosa (Illiger, 1800). Distant (1892a), however noted that "According to Donovan, a single specimen of this species was found in Bengal by Mr. Fichtel, and deposited in the Imperial Cabinet at Vienna, but that habitat I consider liable to the greatest doubt." Cicada indica is illustrated in Fig. 1.
Notes: Authority: Walker 1850; Distant (1906a) refers to a male specimen, but lists two locations. Unlikely to be from India or Sri Lanka: Lee (2012b) states that the type specimen of Terpnosia psecas (Walker, 1850) is a female from Java. Male specimens identified by former authors were not true T. psecas but were T. elegans (Kirby, 1891) from Sri Lanka. The possible confusion with India has not been resolved due to specimen unavailability Kato (1932), however it is highly unlikely given the reported range (Indonesia and Malaysia). Subsequent references to India and Sri Lanka in Duffels and van der Laan (1985) and Sanborn (2014) are likely in reference to these previous misidentifications.
Unduncus connexa (Distant, 1910) Nomenclature: Lemuriana connexa Distant, 1910 Material Holotype: a. scientificName: Unduncus connexa (Distant, 1910); continent: Asia; country: Malaysia; locality: Lawas, Sarawak; sex: female; recordedBy: Moulton; basisOfRecord: PreservedSpecimen Cicadatra sankara (Distant, 1904) x Cicadatra walkeri Metcalf, 1963 x  Bliven, 1964 x Calcagninus nilgirensis (Distant, 1887) x Calcagninus picturatus (Distant, 1888) x Cicada conspurcata (Fabricius, 1777) x Cicada olivierana Metcalf, 1963 x Taungia abnormis Ollenbach, 1929 Neoterpnosia oberthuri (Distant, 1912) x Neoterpnosia versicolor (Distant, 1912) x x Platypleurini 21 2 6 5 5 Oxypleura atkinsoni (Distant, 1912) x Distantalna splendida splendida (Distant, 1878) x x Hyalessa expansa (Walker, 1858) x x Hyalessa mahoni (Distant, 1906) x Hyalessa melanoptera (Distant, 1904) x Hyalessa obnubila (Distant, 1888) x x x Hyalessa stratoria (Distant, 1905) x The cicada fauna inventoried above for the Indian region represents approximately 17% of the currently recognized global cicada genera (73 of 434 genera: data derived from Sanborn 2014). At a country level the generic diversity of India (including Bangladesh) ranks highest in the world (64 of 434 genera), followed by China (61 of 434 genera). The species diversity also compares favourably with that of other countries in the region which have recently been studied (Table 3). Of the 30 species currently recorded from Pakistan (Ahmed and , Ahmed et al. 2010) 21 are recorded from India, while 9 are either not present or yet to be recorded. Although India has larger landmass compared to most of its neighboring countries, the biogeographic placement of modern India and the topographic and climatic diversity of its bioregions may have contributed more to its rich cicada fauna than the size of the landmass. However, this needs to be explored further with phylogenetic / phylogeographic studies. India is rapidly developing and expanding its scientific infrastructure and personnel, and investing heavily in scientific research. There is also growing interest in biodiversity studies and conservation among professional biologists and citizen scientists, which will help in Table 3.
Regional comparison of species and generic diversity, ordered by the number of valid species recorded in each country. exploration and mapping of the current state of Indian biodiversity. As a result, we expect rapid scientific developments involving cicadas of India. To facilitate this development, we have launched a website on Indian cicadas (http://www.indiancicadas.org/), which will act as a central repository of reference images representing spot records, and various species attributes such as cicada songs and other phenotypic variation. Acoustic recordings of Indian cicadas may also be found at: http://bio.acousti.ca/). Following the publication of this paper, we aim to provide newly generated taxonomic and biological information on Indian cicadas through the Cicadas of India website, where updated information may be readily accessed in a centralised database.