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It is with admiration and fondness that we dedicate this special issue of ZooKeys to Professor Kumar Krishna, dean of Isoptera research. This collection of papers is a humble testament of appreciation by the various biologists who are dedicated to studying termites, insects that are popularly maligned but actually of profound behavioral and ecological importance. Kumar’s influence extends beyond the scope of isopterological studies and so several of the papers included herein are from contributors on other lineages of insects who have been similarly inspired by his indomitable spirit.
Kumar grew up in Dehra Dun in northern India, in the foothills of the Himalayas. His father was a physician, and was also one of the first Indians to be commissioned in the British army, in fact serving as a major in World War I. Kumar went to Agra University, earning a Bachelor of Science in 1950, and shortly thereafter a Master of Science degree from Lucknow University in 1952. He then served as a Research Assistant (1952–1954) to Mittan Lal Roonwal (1908–1990) at the Forest Research Institute in Dehra Dun, where he developed his interest in Isoptera. Roonwal is well known among Isoptera workers for his comprehensive papers on the systematics and general biology of termites from India and surrounding areas. Immediately thereafter, he moved to the U.S. where he was a graduate student and employed as a research assistant at the University of Minnesota from 1954–55. It was during this time that he wrote to Prof. Alfred E. Emerson (1896–1976) about graduate studies, and was soon thereafter accepted into Emerson’s lab. Emerson was a professor at the University of Chicago from 1929–1962 and well known as the leading authority on the systematics and general evolution of termites. He was also a coauthor of the classic Principles of Animal Ecology (
Figure 1. Prof. Dr. Kumar Krishna.
At the University of Chicago, Kumar began a comprehensive revision of the world genera of drywood termites, still the major reference on the family Kalotermitidae (
Despite Kumar’s administrative and teaching loads, perhaps the most impressive achievements are that he continued to produce influential research, funded by National Science Foundation research grants. It was during this time, for example, that Kumar Krishna and Frances M. Weesner organized and contributed to the seminal, two-volume work, Biology of the Termites (1969–70), synthesizing all major topics on termite biology and systematics. Interestingly, publication of Biology of the Termites coincided with the first major book by E.O. Wilson, The Insect Societies (1971). It was the heyday of social insects. Although two recent volumes on termite biology have been produced (
Figure 2. Valerie Krishna, Kumar Krishna, Renato Araujo (1912–1978), and Alexander Sokoloff (1920–2011) at the XII International Congress of Entomology, London, 1964.
It is safe to say that no one today has such comprehensive knowledge of termites globally, particularly their systematics, taxonomy, morphology, biogeography, and the fossil record. When Emerson was working, for example, there was only one Cretaceous termite known, Cretatermes carpenteri Emerson. Now, there are 36 species and 32 genera, nearly a quarter of which we have had the pleasure of working with Kumar to describe, but more importantly he has united a critical study of fossil termites with that of their modern counterparts. Emerson would have been immensely pleased to see how much more we now know about termite diversity, relationships, and evolution, largely as a result of Kumar’s efforts. Kumar’s encyclopedic knowledge makes him the ideal person to have been the principal author of the upcoming and highly anticipated magnum opus, Treatise on the Isoptera of the World (2012). At 2400 single-spaced manuscript pages, the work is immense, 85% of which is a taxonomic compendium of the 3138 living and fossil termite species of the world (as of 26 March 2011) – incorporating a plethora of nomenclatural corrections made along the way, all based on direct study of over 4000 original taxonomic references and the more significant biological ones. Whereas many catalogues simply add to previous ones, propagating errors, this work was created de novo. Valerie applied her editorial acumen, spending months fastidiously proofing the manuscript for style and punctuation, a testament to the regard she and Kumar hold for each other.
Putting his considerable academic accomplishments aside, Kumar is one of the most positive, jovial, and generous individuals we know. Now in his eighties, his enthusiasm and energy for termite research seem tireless. When he worked on the Termitidae in Miocene Dominican amber, for example, Kumar spent hours each day hunched over his microscope, examining, sorting, comparing, and measuring hundreds of specimens and describing dozens of species. He worked like a graduate student. Indeed, there is perhaps no more enjoyable way for us to spend an afternoon than taking turns peering into a microscope alongside Kumar, discussing the details of some challenging fossil or exotic living specimen. We look forward to his residence on the fifth floor of the museum for many years to come, where we can peer through the microscope together in excited discussion.
1. Roonwal ML, Krishna K (1955) Systematics of Oriental termites. II. A new species, Coptotermes guarii, from Ceylon. Indian Journal of Agricultural Science 25(2): 143–152.
2. Krishna K (1956) Two new species of Coptotermes Wasmann from Malaya (Isoptera, Rhinotermitidae, Coptotermitinae). American Museum Novitates 1809: 1–5.
3. Krishna K (1960) Social insects. In: McGraw-Hill Encyclopedia of Science and Technology [Volume 14]. McGraw-Hill, New York, NY, 405–407.
4. Krishna K (1961) A generic revision and phylogenetic study of the family Kalotermitidae (Isoptera). Bulletin of the American Museum of Natural History 122(4): 303–408.
5. Krishna K, Emerson AE (1962) New species of the genus Glyptotermes Froggatt from the Papuan, Oriental, Ethiopian, and Neotropical Regions (Isoptera, Kalotermitidae). American Museum Novitates 2089: 1–65.
6. Krishna K (1962) New species and a hitherto undescribed imago caste of the genus Calcaritermes Snyder (Isoptera, Kalotermitidae). American Museum Novitates 2098: 1–13.
7. Krishna K (1962) New species of the genera Allotermes Wasmann, Bicornitermes Krishna, Epicalotermes Silvestri, and Procryptotermes Holmgren (Isoptera, Kalotermitidae). American Museum Novitates 2119: 1–25.
8. Krishna K (1963) The evolution of the family Kalotermitidae, Isoptera. Symposia Genetica et Biologica Italica [Atti IV Congresso U.I.E.I.S., Pavia, 9–14 September 1961] 11: 202–209.
9. Krishna K (1963) The African genus Foraminitermes Holmgren (Isoptera, Termitidae, Termitinae). American Museum Novitates 2161: 1–23.
10. Krishna K (1963) A systematic catalogue of the main identified entomological collection at the Forest Research Institute, Dehradun, Part 32, Order Lepidoptera, Family Papilionidae. Indian Forest Leaflet 121(4): 396–408.
11. Krishna K (1965) A new species of termite from the Nicobar Islands (Isoptera, Termitidae, Nasutitermitinae). Entomologiske Meddelelser 34: 107–109.
12. Krishna K (1965) Termites (Isoptera) of Burma. American Museum Novitates 2210: 1–34.
13. Krishna K (1966) Key to eight termite genera: Kalotermes, Incisitermes, Cryptotermes, Zootermopsis, Heterotermes, Reticulitermes, Coptotermes, and Amitermes. Cooperative Economic Insect Report, Plant Pest Control Division, Agricultural Research Service, United States Department of Agriculture 16(47): 1091–1098.
14. Krishna K, Araujo RL (1968) A revision of the neotropical termite genus Neocapritermes (Isoptera, Termitidae, Termitinae). Bulletin of the American Museum of Natural History 138(3): 83–130.
15. Krishna K (1968) Phylogeny and generic reclassification of the Capritermes complex (Isoptera, Termitidae, Termitinae). Bulletin of the American Museum of Natural History 138(5): 261–324.
16. Krishna K, Weesner FM (1969) Biology of Termites [Volume I]. Academic Press, New York, NY, xiii+598 pp.
17. Krishna K (1969) Introduction. In: Krishna K, Weesner FM (Eds) Biology of Termites [Volume I]. Academic Press, New York, NY, 1–17 [total pages xiii+598 pp.].
18. Krishna K, Weesner FM (1970) Biology of Termites [Volume II]. Academic Press, New York, NY, xiv+[i]+643 pp.
19. Krishna K (1970) Taxonomy, phylogeny, and distribution of termites. In: Krishna K, Weesner FM (Eds) Biology of Termites [Volume II]. Academic Press, New York, NY, 127–152 [total pages xiv+[i]+643 pp.].
20. Krishna K (1972) A revision of the Indomalayan termite genus Homallotermes (Isoptera, Termitidae, Termitinae). American Museum Novitates 2489: 1–23.
21. Krishna K (1974) Isoptera. In: Preece WE (Ed) Encyclopaedia Britannica [15th Edition, Volume 9]. Encyclopaedia Britannica, Inc., London, UK, 1049–1054.
22. Emerson AE, Krishna K (1975) The termite family Serritermitidae (Isoptera). American Museum Novitates 2570: 1–31.
23. Krishna K, Adams C (1982) The Oriental termite genus Labritermes Holmgren (Isoptera, Termitidae, Termitinae). American Museum Novitates 2735: 1–14.
24. Krishna K, Emerson AE (1983) A new fossil species of termite from Mexican amber, Mastotermes electromexicus (Isoptera, Mastotermitidae). American Museum Novitates 2767: 1–8.
25. Krishna K, Bacchus S (1987) A new fossil species of termite from Dominican amber, Cryptotermes yamini (Isoptera, Kalotermitidae). American Museum Novitates 2884: 1–5.
26. Krishna K (1989) Order Isóptera: Termites. In: Borror DJ, Triplehorn CA, Johnson NF (Eds) An Introduction to the Study of Insects [6th Edition]. Saunders College Publishing, Philadelphia, PA, 234–241 [xiv+875 pp.].
27. Krishna K (1990) Isoptera. Bulletin of the American Museum of Natural History 195: 76–81. [Grimaldi DA (Ed) Insects from the Santana Formation, Lower Cretaceous, of Brazil. Bulletin of the American Museum of Natural History 195: 1–191].
28. Krishna K, Grimaldi D (1991) A new fossil species from Dominican amber of the living Australian termite genus Mastotermes (Isoptera: Mastotermitidae). American Museum Novitates 3021: 1–10.
29. Krishna K (1992) Isoptera. In: McHenry R (Ed) The New Encyclopaedia Britannica [15th Edition, Volume 21, Macropaedia]. Encyclopaedia Britannica, Inc., Chicago, IL, 675–679 [total volume pages [iii]+1016 pp.].
30. Krishna K (1996) New fossil species of termites of the subfamily Nasutitermitinae from Dominican and Mexican amber (Isoptera, Termitidae). American Museum Novitates 3176: 1–13.
31. Thorne BL, Grimaldi DA, Krishna K (2000) Early fossil history of the termites. In: Abe T, Bignell DE, Higashi M (Eds) Termites: Evolution, Sociality, Symbioses, Ecology. Kluwer Academic Publishers, Dordrecht, The Netherlands, 77–93 [total volume pages xxii+466 pp.].
32. Krishna K, Grimaldi D (2000) A new subfamily, genus, and species of termite (Isoptera) from New Jersey Cretaceous amber. In: Grimaldi D (Ed) Studies on Fossils in Amber, with Particular Reference to the Cretaceous of New Jersey. Backhuys Publishers, Leiden, The Netherlands, 133–140 [total volume pages viii+498 pp.].
33. Engel MS, Krishna K (2001) Kalotermes Hagen, 1853 (Insecta, Isoptera): Proposed designation of Termes flavicollis Fabricius, 1793 as the type species. Bulletin of Zoological Nomenclature 58(2): 100–104.
34. Engel MS, Krishna K (2001) Macrotermitinae Kemner, 1934 (Insecta, Isoptera): Proposed precedence over Acanthotermitinae Sjöstedt, 1926. Bulletin of Zoological Nomenclature 58(3): 206–209.
35. Krishna K (2001) Southeast Asian species of the genus Dicuspiditermes (Isoptera: Termitidae: Termitinae). Sociobiology 37(3A): 399–488.
36. Engel MS, Krishna K (2002) Cryptotermes dudleyi Banks, 1918 (Insecta, Isoptera): Proposed precedence over Calotermes (Cryptotermes) jacobsoni Holmgren, 1913. Bulletin of Zoological Nomenclature 59(2): 90–92.
37. Krishna K, Grimaldi DA (2003) The first Cretaceous Rhinotermitidae (Isoptera): A new species, genus, and subfamily in Burmese amber. American Museum Novitates 3390: 1–10.
38. Krishna K (2003) A new species, Cavitermes rozeni (Isoptera: Termitidae: Termitinae), from Brazil. Journal of the Kansas Entomological Society 76(2): 92–95.
39. Engel MS, Krishna K, Boyko C (2003) Termopsidae Holmgren, 1911, Termopsis Heer, 1849 and Miotermes Rosen, 1913 (Insecta, Isoptera): Proposed conservation of prevailing usage by the designation of Termes bremii Heer, 1849 as the type species of Termopsis. Bulletin of Zoological Nomenclature 60(2): 119–123.
40. Engel MS, Krishna K (2004) Family-group names for termites (Isoptera). American Museum Novitates 3432: 1–9.
41. Engel MS, Krishna K (2004) Comment on the proposed conservation of prevailing usage of Termopsidae Holmgren, 1911, Termopsis Heer, 1849 and Miotermes Rosen, 1913 (Insecta, Isoptera). Bulletin of Zoological Nomenclature 61(3): 169–170.
42. Engel MS, Krishna K (2004) An overlooked family-group name for termites (Isoptera). Entomological News 115(3): 168.
43. Engel MS, Krishna K (2005) Nasutitermes Dudley, 1890, Microcerotermes Silvestri, 1901 and Nasutitermitinae Hare, 1937 (Insecta, Isoptera): Proposed conservation. Bulletin of Zoological Nomenclature 62(1): 8–13.
44. Engel MS, Krishna K (2005) Comment on the proposed suppression of Eutermes Heer, 1849 to conserve the generic names Nasutitermes Dudley, 1890 and Microcerotermes Silvestri, 1901, and on the proposed designation of Eutermes costalis Holmgren, 1910 as type species of Nasutitermes Dudley, 1890 (Insecta, Isoptera). Bulletin of Zoological Nomenclature 62(4): 240.
45. Krishna K (2005) Order Isoptera: Termites. In: Triplehorn CA, Johnson NF (Ed) Borror and DeLong’s Introduction to the Study of Insects [7th Edition]. Brooks/Cole, Belmont, CA, 252–259 [total volume pages x+864 pp.].
46. Engel MS, Krishna K (2007) Two overlooked family-group names for fossil termites (Isoptera: Mastotermitidae). Entomological News 118(1): 105–106.
47. Lo N, Engel MS, Cameron S, Nalepa CA, Tokuda G, Grimaldi D, Kitade O, Krishna K, Klass K-D, Maekawa K, Miura T, Thompson GJ (2007) Spare Isoptera: A comment on Inward et al. Biology Letters 3(5): 562–563.
48. Engel MS, Krishna K (2007) New Dolichorhinotermes from Ecuador and in Mexican amber (Isoptera: Rhinotermitidae). American Museum Novitates 3592: 1–8.
49. Engel MS, Krishna K (2007) Drywood termites in Dominican amber (Isoptera: Kalotermitidae). Beiträge zur Entomologie 57(2): 263–275.
50. Engel MS, Krishna K (2007) Reticulitermes Holmgren, 1913 (Insecta, Isoptera): Proposed precedence over Maresa Giebel, 1856. Bulletin of Zoological Nomenclature 64(4): 230–234.
51. Engel MS, Grimaldi DA, Krishna K (2007) Primitive termites from the Early Cretaceous of Asia (Isoptera). Stuttgarter Beiträge zur Naturkunde, Serie B, Geologie und Paläontologie 371: 1–32.
52. Engel MS, Grimaldi DA, Krishna K (2007) A synopsis of Baltic amber termites (Isoptera). Stuttgarter Beiträge zur Naturkunde, Serie B, Geologie und Paläontologie 372: 1–20.
53. Grimaldi DA, Engel MS, Krishna K (2008) The species of Isoptera (Insecta) from the Early Cretaceous Crato Formation: A revision. American Museum Novitates 3626: 1–30.
54. Krishna K, Grimaldi DA (2009) Diverse Rhinotermitidae and Termitidae (Isoptera) in Dominican amber. American Museum Novitates 3640: 1–48.
55. Engel MS, Grimaldi DA, Krishna K (2009) Termites (Isoptera): Their phylogeny, classification, and rise to ecological dominance. American Museum Novitates 3650: 1–27.
56. Krishna K, Engel MS (2011) Cryptotermes dudleyi Banks, 1918 (Insecta, Isoptera): Proposed precedence over Calotermes havilandi parasita Wasmann, 1910 (currently Cryptotermes parasita). Bulletin of Zoological Nomenclature 68(2): 109–112.
57. Krishna K, Engel MS, Grimaldi DA, Krishna V (2012) Treatise on the Isoptera of the world. Bulletin of the American Museum of Natural History. In press.
Euisoptera Engel, Grimaldi, & Krishna
Neoisoptera Engel, Grimaldi, & Krishna
Family-group names (6):†Archeorhinotermitinae Krishna & Grimaldi
Archotermopsidae Engel, Grimaldi, & Krishna
†Carinatermitinae Krishna & Grimaldi
†Cratomastotermitidae Engel, Grimaldi, & Krishna
Sphaerotermitinae Engel & Krishna
Syntermitinae Engel & Krishna
Genus-group names (19):†Archeorhinotermes Krishna & Grimaldi
†Baissatermes Engel, Grimaldi, & Krishna
Bicornitermes Krishna
Bifiditermes Krishna
Ceratokalotermes Krishna
Comatermes Krishna
†Dharmatermes Engel, Grimaldi, & Krishna
Dicuspiditermes Krishna
†Garmitermes Engel, Grimaldi, & Krishna
Incisitermes Krishna
†Kachinitermes Engel, Grimaldi, & Krishna
†Khanitermes Engel, Grimaldi, & Krishna
Labiocapritermes Krishna
Marginitermes Krishna
†Melqartitermes Engel, Grimaldi, & Krishna
†Mylacrotermes Engel, Grimaldi, & Krishna
Postelectrotermes Krishna
†Tanytermes Engel, Grimaldi, & Krishna
Tauritermes Krishna
Species-group names (106):Aciculitermes maymyoensis Krishna
Allotermes denticulatus Krishna
Allotermes papillifer Krishna
†Amitermes lucidus Krishna & Grimaldi
Angulitermes paanensis Krishna
Angulitermes resimus Krishna
†Anoplotermes bohio Krishna & Grimaldi
†Anoplotermes cacique Krishna & Grimaldi
†Anoplotermes carib Krishna & Grimaldi
†Anoplotermes maboya Krishna & Grimaldi
†Anoplotermes naboria Krishna & Grimaldi
†Anoplotermes nitaino Krishna & Grimaldi
†Anoplotermes quisqueya Krishna & Grimaldi
†Anoplotermes taino Krishna & Grimaldi
†Archeorhinotermes rossi Krishna & Grimaldi
†Atlantitermes antillea Krishna & Grimaldi
†Atlantitermes caribea Krishna & Grimaldi
†Atlantitermes magnoculus Krishna & Grimaldi
†Baissatermes lapideus Engel, Grimaldi, & Krishna
Bicornitermes bicornis Krishna
Bicornitermes emersoni Krishna
Bulbitermes prabhae Krishna
Calcaritermes colei Krishna
Calcaritermes rioensis Krishna
Calcaritermes snyderi Krishna
†Caribitermes hispaniola Krishna & Grimaldi
Cavitermes rozeni Krishna
†Constrictotermes electroconstrictus Krishna
Coptotermes bentongensis Krishna
†Coptotermes hirsutus Krishna & Grimaldi
†Coptotermes paleodominicanus Krishna & Grimaldi
Coptotermes sepangensis Krishna
†Cryptotermes yamini Krishna & Bacchus
†Cryptotermes glaesarius Engel & Krishna
†Dharmatermes avernalis Engel, Grimaldi, & Krishna
Dicuspiditermes cacuminatus Krishna
Dicuspiditermes fissifex Krishna
Dicuspiditermes kistneri Krishna
Dicuspiditermes spinitibialis Krishna
†Dolichorhinotermes apopnus Engel & Krishna
Dolichorhinotermes lanciarius Engel & Krishna
Epicalotermes planifrons Krishna
Foraminitermes coatoni Krishna
Foraminitermes harrisi Krishna
†Garmitermes succineus Engel, Grimaldi, & Krishna
Glyptotermes adamsoni Krishna & Emerson
Glyptotermes concavifrons Krishna & Emerson
†Glyptotermes grimaldii Engel & Krishna
Glyptotermes guamensis Krishna & Emerson
Glyptotermes hendrickxi Krishna & Emerson
Glyptotermes jurioni Krishna & Emerson
Glyptotermes kirbyi Krishna & Emerson
Glyptotermes lighti Krishna & Emerson
Glyptotermes longipennis Krishna & Emerson
Glyptotermes longuisculus Krishna & Emerson
Glyptotermes nissanensis Krishna & Emerson
Glyptotermes palauensis Krishna & Emerson
†Glyptotermes paleoliberatus Engel & Krishna
Glyptotermes parki Krishna & Emerson
Glyptotermes parvoculatus Krishna & Emerson
Glyptotermes rotundifrons Krishna & Emerson
Glyptotermes schmidti Krishna & Emerson
Glyptotermes seeversi Krishna & Emerson
Glyptotermes sicki Krishna & Emerson
Glyptotermes sinomalatus Krishna & Emerson
Glyptotermes truncatus Krishna & Emerson
Glyptotermes tuberifer Krishna & Emerson
Homallotermes eleanorae Krishna
Homallotermes exiguus Krishna
†Incisitermes peritus Engel & Krishna
Labritermes emersoni Krishna & Adams
Labritermes kistneri Krishna & Adams
†Mastotermes electrodominicus Krishna & Grimaldi
†Mastotermes electromexicus Krishna & Emerson
†Meiatermes araripena Krishna
†Melqartitermes myrrheus Engel, Grimaldi, & Krishna
†Microcerotermes insulanus Krishna & Grimaldi
†Microcerotermes setosus Krishna & Grimaldi
Microcerotermes uncatus Krishna
Mirocapritermes valeriae Krishna
†Mylacrotermes cordatus Engel, Grimaldi, & Krishna
†Nasutitermes amplioculatus Krishna & Grimaldi
†Nasutitermes electrinus Krishna
†Nasutitermes electronasutus Krishna
Nasutitermes fabricii Krishna
†Nasutitermes incisus Krishna & Grimaldi
†Nasutitermes magnocellus Krishna & Grimaldi
†Nasutitermes medioculatus Krishna & Grimaldi
†Nasutitermes pilosus Krishna & Grimaldi
†Nasutitermes rotundicephalus Krishna & Grimaldi
†Nasutitermes seminudus Krishna & Grimaldi
Neocapritermes araguaia Krishna & Araujo
Neocapritermes guyana Krishna & Araujo
Neocapritermes talpoides Krishna & Araujo
Neocapritermes taracua Krishna & Araujo
Neocapritermes utiariti Krishna & Araujo
†Parvitermes longinasus Krishna & Grimaldi
Procapritermes parvulus Krishna
Procryptotermes falcifer Krishna
†Proelectrotermes holmgreni Engel, Grimaldi, & Krishna
†Subulitermes hispaniola Krishna & Grimaldi
†Subulitermes insularis Krishna & Grimaldi
†Tanytermes anawrahtai Engel, Grimaldi, & Krishna
†Termes primitivus Krishna & Grimaldi
†Termopsis ukapirmasi Engel, Grimaldi, & Krishna
†Velocitermes bulbus Krishna & Grimaldi