© Prof. T.C. Narenderan Trust for Biosystematica http://www.tcntrust.org/journal.php ISSN: 0973-7871(online) ISSN: 0973-9955 (print)

A Review of Extant , Tribes and Genera in India

THRESIAMMA VARGHESE

Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560 012, India.

ABSTRACT. More than a century old monograph” Fauna of British India, including Ceylon and Burma – – 2. and Cuckoo-Wasps” by Bingham is still the guiding source for anybody who initiates taxonomic studies of Indian ants. However, over a century period, there are lots of changes in the status of many names, at the , , and levels, largely owing to several revision studies in many groups of the Formicidae. In the recent past, across the world there are several remarkable endeavours in identifying the major of ants and to understand the relationships among them. In this review, the author has looked at some of those changes, which are pertaining to Indian fauna and the results are summarised below. Out of 5 subfamilies of the family Formicidae, mentioned in Bingham, 4 of them have valid status, while one of them is now known under a different name. Of the 79 genera mentioned, 53 of them are currently known to be present in India and retains the same name and valid status. Further, as per a recent approximation of ant species in India estimates 633 species of ants coming under 82 genera distributed across 13 subfamilies.

KEYWORDS. Ants, Formicidae, subfamily, tribe, genera, and species.

Introduction More than a century old, the precious book “Fauna of provided an extensive review on the current status of British India” on ant taxonomy by C.T. Bingham (1903) ant systematics. Recently, one more subfamily, is surely the beginning point for each and every person, Martialinae is added to the family Formicidae and who gets interested in taxonomy of the Indian ants. More proposed to be the sister lineage to all extant ants ever, it is still the guiding light for the beginners in ant (Rabeling et al., 2008). taxonomy. More over, revisions of majority of subfamilies, But over the years, the developments occurred in genera and species groups have been carried out by ant systematics across the globe, with the aid of various various people across the globe. The main revisions are tools and techniques in biology, ecology, behaviour and by Brown (1952, 1954, 1958, 1959, 1960 & 1975), Bolton phylogeny are excitingly remarkable. With the help of (1974, 1976, 1977, 1986, 1994, 1995 & 2003), Shattuck refined morphological characters, molecular, ecological (1992), Ward (1990 & 2001), Wilson (2003), and LaPolla and behavioural data, along with information from new (2004). The book “The Ants” by Holldobler and Wilson , the ant phylogeny is better understood now than (1990) has compiled all known information on the biology ever before in the history of ant taxonomy and of ants till then known. systematics. Some of the recent molecular phylogenetic The best taxonomic achievement among all studies carried out by Ward & Brady (2003), Saux et developments is that identification keys for all subfamilies al., (2004), Brady et al., (2006), Moreau et al., (2006), and genera are available in English (Bolton,1994). In Ouellette et al., (2006), and other studies claim that addition, world class catalogue on ants of the world is “Formicoid ” is the largest clade comprising 14 of also made available by Bolton (1995). An outstanding, the 20 extant ant subfamilies (Ward, 2007). He has single piece of publication, which provides information

Corresponding author: [email protected] Biosystematica, 2009, 3(2): 81-89 81 82 THRESIAMMA VARGHESE about the diagnosis and status of all known extinct and The web page developed and maintained by the extant subfamilies, tribes, and genera of ants across the California Academy of Sciences is a marvellous world is the synopsis and classification of the family reference place for ant genera of the world. The antbase Formicidae by Bolton (2003). also provides links to many useful databases, like primary As a result of many such studies, there are quite taxonomic publications, Formis bibliography, distribution a lot of changes in the classification and scientific names database, Integrated Taxonomic Information System, of many ant species across the globe. Subsequently, the Hymenoptera Name Server etc. In addition, Global status of many subfamilies, tribes, genera and species Information Facility (GBIF-http:// names of ants in India also have changed; there are www.gbif.org, Date of access- 05/11/09) and Global many synonyms, new combinations,new revivals etc. Taxonomic Initiative (GTI-http://www.cbd.int/gti, Date of access-05/11/09) coordinate the efforts of various Since these changes have happened and have national museums and universities, and provide data on been happening at an exhilarating rate, all over the world species all over the world in an effort to trim down the over many years and carried out by various people in taxonomic impediment. In Asian countries, the database different parts of the world, all information is naturally on ant taxonomy and ecology is the antbase (http:// scattered and is not easily accessible to a beginner; though www.antbase.net/, Date of access-05/11/09). In addition most of the information is available online and become to this, another international network for Asian more or less transparent. myrmecologists for the improvement of ant taxonomy However, through modern approaches of and ecology is ANeT (http://homepage.mac.com/dorylus/ enormously passionate groups of people, and institutions, jorney.html, Date of access-05/11/09). This excellent the whole field of taxonomy and systematics is gaining initiative publishes a scientific journal “Asian significant momentum. More over, they are trying hard Myrmecology”. to bring most of the available information on biology, However, the main problem one faces, when one ecology, behaviour, taxonomy and systematics of ants look at ant genera and species in India is that the status to everyone through internet and other means. One of of many names given in Bingham (1903) have such ultimate attempts is the “ E-type initiative” recently undergone some changes. In this review, the author has began at the Museum of Comparative Zoology at looked at some of those changes and have summarised Harvard University, with the aim of putting detailed them below with the hope that this information is useful pictures of many ant species on Earth on the internet, to the beginners, especially for those who do not have which will be accessible to ant researchers all over the easy access to internet facility. world (http://insects.oeb.harvard.edu/etypes/about.htm, Date of access-05/11/09). As a result, many of the The following is an abstract of the major changes, problems existing in ant taxonomy, because of the which took place at the subfamily and genus level inaccessibility of many type specimens and the data taxonomy of ants in India, over a century. Since the associated with them, will eventually fade away. author aims this publication, on the whole, at the amateur ant taxonomists, it isn’t represented here in a strict As a result of many such unfailing efforts, at taxonomic pattern, but rather in a general form. A present, there are many useful websites, which are of synonymic synopsis of the Bingham’s species list is extreme use to any ant taxonomist. Some of the excellent available at http://ces.iisc.ernet.in/thresi/ web based identification resources freely available are AntWeb (www.antweb.org, Date of access-05/11/09), Results and Discussions Antbase (www.antbase.org, Date of access-05/11/09), As a result of various studies based on Australian Ants online (www.ento.csiro.au/science/ants, morphology, molecular data, along with information from Date of access-05/11/09). The Antweb and another fossils, the evolution of ants and their characters, database on Japanese ants are the first image databases relationships between various subfamilies, tribes and on ants. They provide good quality detailed photographs genera and species are better understood now and of ants showing taxonomic features, which enables subsequently they are reclassified and reorganized in species identification, without physically accessing the the present form (Bolton, 2003, Bolton et al., 2006). type specimens. Since most of the images are taken An extensive review of the work relevant to with high resolution microscopes and processed using the Indian fauna is presented in the manuscript auto montage software, the images are spectacular and (Varghese, in press). When one looks at the scenario in the details are shown to finer details. India, there isn’t much development in ant taxonomy

Biosystematica, 2009, 3(2) Review of Extant Subfamilies, Tribes and Ant Genera in India 83 over the last hundred years. Hence there is nothing much The major changes at the genus level are the to look beyond the fauna volume by Bingham (1903). following. Two genera, and Aenictus retain their This book provides descriptions and identification keys name. Under the subfamily , the Lioponera for ants up to species level, which still remains as the Mayr, Syscia Roger, and Ooceraea Roger, are major guiding source for the Indian ant taxonomists. synonymised under the genus Cerapachys Smith The fauna volume included the species of Sri (Brown, 1973 & 1975; Kempf, 1972). As mentioned Lanka and Burma also. Now the present author earlier, the genus Cerapachys is placed in the subfamily considers the fauna of political India only, mainly to make Cerapachyinae, not in the subfamily Ponerinae as it simple and to facilitate convenience for the Indian classified in the Bingham’s book (Wheeler, 1902, Bolton, audience. Bingham(1903) treated all ants in the single 2003). This is true for the genus Sphinctomyrmex Mayr family Formicidae, and classified them under 5 different also, which is being transferred to the subfamily subfamilies. Cerapachyinae from Ponerinae (Wheeler, 1902, 1922 & Bolton, 2003). Out of 5 subfamilies mentioned in Bingham (1903), 4 of them are still valid with the same name, Another change in the status has taken place while Camponotinae is now known as for the genus Stigmatomma Roger, as per the recent (Wheeler, 1920; Bolton, 1994, 2003). The two genera classifications the valid name for this is given under the subfamily , Aenictus Erichson (Emery and Forel, 1879 & Brown, 1960), and Shuckard, and Dorylus Fabricius, are now placed is placed under the subfamily (Forel, under 2 different subfamilies, Dorylinae and 1893). The other two genera, which are listed in the Aenictinae (Bolton, 1990). The subfamily Ponerinae fauna volume under the subfamily Ponerinae, Mystrium has undergone tremendous revisions (Emery, 1911; and Myopopone are also classified under this subfamily. Forel, 1917 & 1921; Wheeler, 1910, 1920 & 1922; Even though this subfamily name was in use for some Brown and Nutting, 1950; Brown, 1954, 1973, 1976, time it was given the status of a tribe only, until Bolton 1978a & b; Wilson & Taylor, 1967; Wheeler & (2003), revived its subfamily status. Wheeler, 1972, 1976 & 1985; Bolton, 1975; Snelling, Similarly, the name Drepanognathus Smith is 1981; Dlussky & Fedoseeva, 1988; Baroni Urbani et synonymised under Harpegnathos Jerdon (Roger, al., 1992; Bolton, 1994, 1995 and Grimaldi et al., 1997). 1861), and Lobopelta Mayr under Leptogenys Roger As per the recent classification, “Poneroids” ants are respectively (Emery, 1896). Another large and classified roughly into 5 subfamilies viz., complicated genus, where key changes have taken place , Amblyoponinae, Paraponerinae, is the Pachycondyla Smith. At present the following Ponerinae and (Bolton, 2003). As per five genera, Ectomomyrmex Mayr, Pseudoponera the recent classification, two genera, Cerapachys Emery, Bothroponera Mayr, Mesoponera Emery, and Smith, and Sphinctomyrmex Mayr, are placed in the Brachyponera Emery are all put under the genus subfamily Cerapachyinae. Other primitive genera, Pachycondyla (Brown, 1973). Though the genus name Amblyopone Erichson, Mystrium Roger, and Smith remains as a valid name, the mention Myopopone Roger are classified under the subfamily in this volume is referring to the genus Gnamptogenys Amblyoponinae (Bolton, 1994, 1995 & 2003). The Roger. At present, both the genera are grouped under subfamily is split into 2 subfamilies as the subfamily (Bolton, 2003). The genus and Myrmicinae (Wheeler, 1920; Latreille mentioned in this book is actually the Bolton, 1994). Similarly, the subfamily current Hypoponera Santschi, but the genus Ponera is is divided into 2 subfamilies as Dolichoderinae and absolutely a valid name under the subfamily Ponerinae. (Clark, 1951; Bolton, 1994 & 2003). Taking into consideration the enormous changes Of the 79 genera mentioned in the fauna volume, in the subfamily Ponerinae, there are only very few 53 of them are currently present in India, retains the changes in the subfamily Myrmicinae, as far as the valid status under the same name. Among them, 59 of names of Indian ants are concerned. One of the major them are present in India either in the same or different changes is the placing of the genus Sima under a name. While 59 of them retains the valid status, all valid separate subfamily Pseudomyrmecinae and given the genera, on the other hand, are not present in India. For name Tetraponera (Smith, 1852, 1877; & Wheeler, example, the genus Aneuretus Emery, mentioned in the 1922). The following are the major changes in genus book is endemic to . names in the subfamily Myrmicinae; the Sima is

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Tetraponera Smith, Triglyphothrix Forel is were, involved in ant diversity and taxonomic studies in Tetramorium Mayr (Bolton, 1985), Trichomyrmex Mayr India. (Ettershank, 1966), and Holcomyrmex Mayr are under 1. Prof. R. Gadagkar, Centre for Ecological Monomorium Mayr (Emery, 1908), and Trigonogaster Sciences, Indian Institute of Science, Bangalore, Forel is Recurvidris Bolton (Bolton, 1992). The genus 560012, Karnataka, India. Oligomyrmex is a synonym of Carebara Westwood 2. Dr. Himender Bharathi, Department of Zoology, (Fernandez, 2004). Punjabi University, Patiala , Punjab, India , The subfamily Camponotinae is now the subfamily 147002. Formicinae. There are only very few changes in this 3. Sr. Karmaly K.A., Dept of Zoology, St. Xavier’s subfamily as far as the Indian fauna is concerned. The college for Women, Aluva, 683101, Kerala, India. genus Westwood mentioned in this book refers the genus Cataglyphis Forster (Brown, 1973), 4. Merry Zacharias (Earlier at Ashoka Trust for where as Acantholepis Mayr is synonymised under the Research in Ecology and the Environment, genus Santschi (Bolton, 1994). Likewise, the Royal Enclave, Srirampura, Jakkur P.O., genus Colobopsis Mayr is synonymised with the genus Bangalore, 560064, Karnataka). Camponotus Mayr and Hemioptica Roger is placed 5. Ali T. M. University of Agricultural Sciences, under the genus Polyrhachis Smith. Other few explicit, Gandhi Krishi Vignyan Kendra, Bangalore, but not so apparent changes are also there. The genus 560065, Karnataka, India. Mayr in this book includes the current 6. Dr. Narendra, A. Currently working as research Santschi also. Additionally, the current associate at Australian National University, genus Motschoulsky is given under the Canberra, Australia. genus Mayr. However, the status of all the above mentioned genera remains valid (Bolton, 2003). 7. Prof. Narendran T.C., Department of Zoology, University of Calicut, 673635, Kerala, India. The subfamily Dolichoderinae has only very few changes to supplement. One of the major changes is 8. Dr. Neelkamal Rastogi, Department of Zoology, that the genus Aneuretus Emery is now placed in a Centre of Advanced Study, Banaras Hindu separate subfamily Aneuretinae (Clark, 1951; Bolton, University, Varanasi, U.P. -221 005, India. 2003). 9. Dr. Parthiba Basu, Reader, Ecology Research According to a recent world statistics, there are Unit, Department of Zoology, Kolkatta approximately 12,592 described species of ants University, 35, Ballygunge Circular Road, (www.antbase.org, Date of access-05/11/09; as per Kolkata, 700019, India. Antweb’s (www.antweb.org) world statistics, there are 10. Dr. Priyadharsan Dharmarajan, Ashoka Trust 14,062 species of described ant species distributed over for Research in Ecology and the Environment, 294 genera across 22 subfamilies-date of access-5/11/ Royal Enclave, Srirampura, Jakkur P.O. 09). As per a current estimate, there may be as many as Bangalore, 560 064, Karnataka. 633 species of described ant species in India, distributed 11. Sabu K. Thomas, Litter Entomology Research over 82 valid genera under 13 subfamilies. A detailed Unit, P.G. & Research Department of Zoology, list of all valid genera and extant species currently known St. Joseph’s College, Devagiri, Calicut, 673 008, to exist in India is freely available at the following Kerala, India. address, http://ces.iisc.ernet.in/thresi/AntsOfIndia.htm. 12. Dr. Sheela S., Zoological Survey of India, Prani The following table summarises the number of current Vigyan Bhawan, ‘M’ Block, New Alipore, subfamilies, tribes and genera, which are present in India, Kolkatta - 700 053 (WB), India. in alphabetical (Table 1). (Based on Bingham, 13. Sreenivasa, Y.B., Institute of Wood Science and 1903; Bolton, 1995 & 2003) (Compiled by Thresiamma Technology, 18th Cross, Malleswaram, Varghese). Bangalore- 560 006, Karnataka, India. For the last two decades, ants are noticeably 14. Sunil Kumar M. (Earlier at attaining enormous concern in India also. As a result, Deccan Herald, many more people are getting interested in ants and Bangalore, Karnataka, India) consequently, many more publications are out in this line 15. Tiwari, R.N. (Earlier Director of ZSI, Kolkatta) in India. The following is a list of few people, who are/ Few important readings (other than the ones given

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Table 1. List of current subfamilies, tribes and genera in India Sl.No. Subfamilies Tribes Genera 1 Aenictinae Aenictini 1. Aenictus, Shuckard 2 Amblyoponinae Amblyoponini 1. Amblyopone, Erichson Amblyoponini 2. Myopopone, Roger 3 Cerapachyinae Cerapachyini 1. Cerapachys, Smith Cerapachyini 2. Sphinctomyrmex, Mayr 4 Dolichoderinae Dolichoderini 1. Bothriomyrmex, Emery Dolichoderini 2. , Lund Dolichoderini 3. Dorymyrmex, Mayr Dolichoderini 4. Liometopum, Mayr Dolichoderini 5. Philidris, Shattuck Dolichoderini 6. Tapinoma, Forster Dolichoderini 7. Technomyrmex, Mayr 5 Dorylinae Dorylini 1. Dorylus, Fabricius 6 Ectatomminae Ectatommini 1. Gnamptogenys, Roger 7 Ecitoninae Ecitonini 1. Eciton, Latreille 8 Formicinae 1. , Roger Lasiini 2. Anoplolepis, Santschi Camponotini 3. Camponotus, Mayr Formicini 4. Cataglyphis, Forster Formicini 5. , Linnaeus Gesomyrmecini 6. Gesomyrmex, Mayr Lasiini 7. , Fabricius 8. Lepisiota, Santschi Myrmoteratini 9. Myrmoteras, Forel Oecophyllini 10. Oecophylla, Smith Plagiolepidini 11. Paratrechina, Motschoulsky . Plagiolepidini 12. Plagiolepis, Mayr Camponotini 13. Polyrhachis, Smith Plagiolepidini 14. Prenolepis, Mayr Plagiolepidini 15. , Emery Camponotini 16. Echinopla, Smith 9 Leptanillini 1. , Emery Leptanillini 2. Yavnella, Kugler 10 Myrmicinae Myrmecinini 1. Acanthomyrmex, Emery Pheidolini 2. Aphaenogaster, Mayr Stenammini 3. Calyptomyrmex, Emery Formicoxenini 4. Cardiocondyla, Emery Solenopsidini 5. Carebara, Westwood Cataulacini 6. Cataulacus, Smith 7. Crematogaster, Lund Formicoxenini 8. Dilobocondyla, Santschi Stenammini 9. Indomyrma, Brown Pheidolini 10. Kartidris, Bolton Liomyrmecini 11. Liomyrmex, Mayr

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Formicoxenini 12. Leptothorax, Mayr Pheidolini 13. Lophomyrmex, Emery Meranoplini 14. Meranoplus, Smith Pheidolini 15. Messor, Forel Metaponini 16. Metapone, Forel Solenopsidini 17. Monomorium, Mayr Myrmecinini 18. Myrmecina, Curtis Myrmicini 19. , Latreille Myrmicariini 20. Myrmicaria, Saunders Paratopulini 21. Paratopula, Wheeler Pheidolini 22. Pheidole, Westwood Solenopsidini 23. Pheidologeton, Mayr Crematogastrini 24. Recurvidris, Bolton Melisotarsini 25. Rhopalomastix, Forel Tetramorini 26. Rhoptromyrmex, Mayr Solenopsidini 27. Solenopsis, Westwood Dacetini 28. Strumigenys, Smith Tetramorini 29. Tetramorium, Mayr Stenammini 30. Vollenhovia, Mayr Formicoxenini 31. Vombisidris, Bolton 11 Ponerinae 1. Anochetus, Mayr Ponerini 2. Centromyrmex, Mayr Ponerini 3. Cryptopone, Emery Ponerini 4. Diacamma, Mayr Ponerini 5. Emeryopone, Forel Ponerini 6. Harpegnathos, Jerdon Ponerini 7. Hypoponera, Santschi Ponerini 8. Leptogenys, Roger Ponerini 9. , Latreille Ponerini 10. Odontoponera, Mayr Ponerini 11. Pachycondyla, Smith Platythyreini 12. Platythyrea, Roger Ponerini 13. Ponera, Latreille 12 Proceratiinae Proceratiini 1. Discothyrea, Roger Probolomyrmecini 2. Probolomyrmex, Mayr Proceratiini 3. , Roger 13 Pseudomyrmecinae Pseudomyrmecini 1. Tetraponera, Smith Subfamilies - 13 Tribes - 36 Genera - 82 (Compiled by Thresiamma Varghese, based mainly on Bingham, 1903, Bolton, 1995 & 2003).

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Received: 12-Nov-2009 Accepted: 03-Dec-2009