Kumar Krishna

Termites (Isoptera): Their Phylogeny, Classification, and Rise to Ecological Dominance

Abstract Like ants, termites are entirely eusocial and have profound ecological significance in the tropics. Following upon recent studies reporting more than a quarter of all known fossil termites, we present the first phylogeny of termite lineages using exemplar Cretaceous, Tertiary, and Recent taxa. Relationships among Recent families were largely unaffected by the addition of extinct taxa, but the analysis revealed extensive grades of stem-group taxa and the divergence of some modern families in the Cretaceous. Rhinotermitidae, Serritermitidae, and the “higher” termites (family Termitidae), which comprise 84% of the world termite species, diverged and radiated entirely in the Tertiary, corresponding to a significant increase in termite individuals in the fossil record. Radiation of the higher termites may have affected the formation of terrestrial carbon reserves like oil and coal. The higher classification of Isoptera is slightly revised based on the phylogenetic results. The following new taxa are proposed: Cratomastotermitidae, new family; Euisoptera, new clade; Archotermopsidae, new family; and Neoisoptera, new clade. In addition, the families Stolotermitidae, Stylotermitidae, and Archeorhinotermitidae are newly recognized or resurrected, and the families Termopsidae and Hodotermitidae are significantly restricted in composition.

Treatise on the Isoptera of the World

ABSTRACT A comprehensive compendium on the taxonomy and biology of the 3106 living and fossil species of the worlds termites is presented, along with reviews of Isoptera morphology and evolution, identification keys, the history of termite systematics, and summary of the worlds 363 significant pest species. A complete bibliography is provided of nearly 5000 references covering virtually all aspects of termite taxonomy and biology through December 2011. The morphology of Isoptera is thoroughly reviewed and illustrated with original scanning electron micrographs and photomicrographs, covering the cuticular anatomy and those internal organs that are taxonomically and phylogenetically significant, including several new character systems. Terminology is presented for the systems of tibial spines and spurs so as to establish homologs. Keys are presented to the nine living families of termites, and the world subfamilies and genera of Archotermopsidae, Hodotermitidae, Kalotermitidae, Mastotermitidae, Rhinotermiti...

Treatise on the Isoptera of the World: VOLUME 1 INTRODUCTION

ABSTRACT A comprehensive compendium on the taxonomy and biology of the 3106 living and fossil species of the worlds termites is presented, along with reviews of Isoptera morphology and evolution, identification keys, the history of termite systematics, and summary of the worlds 363 significant pest species. A complete bibliography is provided of nearly 5000 references covering virtually all aspects of termite taxonomy and biology through December 2011. The morphology of Isoptera is thoroughly reviewed and illustrated with original scanning electron micrographs and photomicrographs, covering the cuticular anatomy and those internal organs that are taxonomically and phylogenetically significant, including several new character systems. Terminology is presented for the systems of tibial spines and spurs so as to establish homologs. Keys are presented to the nine living families of termites, and the world subfamilies and genera of Archotermopsidae, Hodotermitidae, Kalotermitidae, Mastotermitidae, Rhinotermitidae, Serritermitidae, Stolotermitidae, and Stylotermitidae. A key to subfamilies of the Termitidae is included. A detailed morphological diagnosis for each family and subfamily is provided, along with images of exemplar species. The history of isopteran research in taxonomy, systematics, morphology, paleontology, and biology is reviewed from 1758 to the present, with emphasis on transformative workers such as Holmgren, Silvestri, Emerson, Roonwal, Noirot, and Sands. Evolution of the Isoptera is reviewed, including the diversity and natural history of genera and species in all Zoogeographic regions, major patterns in social biology, the phylogeny of Recent and fossil genera and families, and 135 million years of fossils preserved as compressions, mineralized replicas, and in amber. The definitive sister group to the Isoptera is the monogeneric family of wood roaches, Cryptocercidae (Cryptocercus), so the taxonomic ranks of the two groups are now Infraorder Isoptera and Infraorder Cryptocercoidea within Order Blattaria (roaches and termites). The compendium summarizes the taxonomic history, nomenclature, distribution, type locality, and repository, and all significant aspects of natural history and biology for each species of the world, exclusive of pest control and colony inquilines (termitophiles). The classification of Recent and fossil lower termites (all those exclusive of family Termitidae) used in the compendium is from Engel et al. (2009), which is based on morphology and largely congruent with molecular studies. Rhinotermitidae s.s. (exclusive of Stylotermitidae) may be paraphyletic with respect to Termitidae, although the six traditional subfamilies of the former are used here. A separate section summarizes the nomenclatural changes made in the compendium, including new synonymies, new combinations, status novus, lectotype selection etc. A detailed list is provided of museums and other institutional collections that house type specimens. An index is included. The Treatise is intended to provide an authoritative foundation for taxonomic work on the Isoptera, present and future.

Early Fossil History of the Termites

All Mesozoic records of termites are from the Cretaceous and are of the families Hodotermitidae, Termopsidae, and possibly the Mastotermitidae. All known fossil Isoptera from the Cretaceous, including described and undescribed specimens, have distinctly primitive wing venation and, to the extent that can be evaluated, primitive morphology of other body regions. Thus termites were primitive but reasonably diversified in the Cretaceous, suggesting an origin in the Upper Jurassic, but probably not earlier. The geographic range of known Mesozoic termite fossils includes Europe, Asia, and North and South America, indicating a broad Pangean distribution and both temperate and tropical habitats. We suggest that the driving forces behind modern termite distributions were not early evolution and continental drift, but rather an explosive Tertiary radiation followed by broad and rapid dispersal as termites became ecologically dominant consumers. Relict distributions of modem Mastotermitidae, Hodotermitidae, and Termopsidae, the origin and the radiation of social insects are discussed.

Family-Group Names for Termites (Isoptera)

Abstract Thirty-nine available family-group names are identified within the insect order Isoptera (termites). For all names the correct author, date, type genus, and combining stem are provided for the first time. This nomenclatural compilation is done to stabilize the usage of family- group names in the Isoptera in advance of a world catalog. Several problems of priority are identified and discussed. The little understood subfamily Foraminitermitinae is diagnosed; while generally believed by many authors to be a new, unnamed subfamily, it was in fact established by Holmgren nearly a century ago. The subfamilies Syntermitinae and Sphaerotermitinae are newly proposed for the mandibulate genera of nasute termites and for Sphaerotermes, respectively. The classification of Isoptera is briefly outlined.

Save Isoptera: A comment on Inward et al.

A number of phylogenetic studies during the last decade have shown that termites—one of the main groups of eusocial insects—are a type of cockroach, whose closest living relative is the wood-feeding genus Cryptocercus (reviewed in [Klass & Meier (2006)][1] and [Inward et al. (2007)][2]). Inward

The Species of Isoptera (Insecta) from The Early Cretaceous Crato Formation: A Revision

Abstract The termite species from Brazils Early Cretaceous (Aptian-aged) Crato (Santana) Formation are evaluated on the basis of the degree of character variation seen in modern species, using a series of 56 specimens, scanning electron microscopy of minute structures, and a bivariate plot of the proportional sizes of sclerotized body structures. Of the previously described species only the following are considered valid: Mariconitermes talicei Fontes and Vulcano, Meiatermes araripena Krishna, Cratomastotermes wolfschwenningeri Bechly, Cratokalotermes santanensis Bechly, and Cretarhinotermes novaolindense Bechly. The combination M. araripena Krishna (once placed in Cretatermes) is restored, and Cretatermes pereirai Fontes and Vulcano is proposed as a junior synonym of M. araripena. The following new species is described: Meiatermes hariolus Grimaldi, new species. The following are considered nomina dubia based on superficial and even contradictory diagnoses: Caatingatermitinae Martins-Neto et al. (likely synonymous with Hodotermitinae); Araripetermes nativa Martins-Neto et al. (nomen incorrectum, recte: nativus), Caatingatermes megacephalus Martins-Neto et al., and Nordestinatermes obesa Martins-Neto et al. (nomen incorrectum, recte: obesus). These are probably all new synonyms of the Cretaceous genus Meiatermes Lacasa-Ruiz and Martínez-Delclòs and of M. araripena in particular, but this assertion cannot be verified without access to the type specimens. Of the six definitive species of Crato termites, reconstructions are provided for four. All species appear to be basal taxa, either a stem group to Isoptera, to Hodotermitidae sensu lato, or to Kalotermitidae. Despite very partial preservation, Cretarhinotermes appears to be within the hodotermitid grade and not a rhinotermitid. Some specimens have yielded detailed preservation of soft internal tissues, including the midgut, which has further phylogenetic implications.

The First Cretaceous Rhinotermitidae (Isoptera): A New Species, Genus, and Subfamily in Burmese Amber

Abstract A new subfamily, genus, and species, Archeorhinotermitinae, Archeorhinotermes rossi, from Burmese amber, dated as Turonian-Cenomanian (90–100 mya) of the Cretaceous period, are described and figured. Comparisons are made between the other subfamilies of the Rhinotermitidae and the new subfamily. This is the first fossil record of the family Rhinotermitidae from the Cretaceous.

Diverse Rhinotermitidae and Termitidae (Isoptera) in Dominican Amber

Abstract The most diverse and best-preserved paleofauna of the higher termites heretofore known, all found in Miocene amber of the Dominican Republic, is described. The imago of Coptotermes priscus Emerson is redescribed, and the soldier of C. priscus, the first known fossil soldier of this genus, is described. The fauna includes the following 29 new species, all in existing genera, with Krishna and Grimaldi as authors of each: in the Rhinotermitidae, two new species based on imagoes of each—Coptotermes hirsutus and C. paleodominicanus; in the Termitidae, 23 new species based on imagoes—Amitermes lucidus, Anoplotermes bohio, A. cacique, A. carib, A. maboya, A. naboria, A. nitaino, A. quisqueya, A. taino, Atlantitermes antillea, A. caribea, A. magnoculus, Microcerotermes insulanus, M. setosus, Nasutitermes amplioculatus, N. incisus, N. magnocellus, N. medioculatus, N. pilosus, N. seminudus, Subulitermes hispaniola, S. insularis, and Termes primitivus; in the Nasutitermitinae four new species based on nasute soldiers—Caribitermes hispaniola, Nasutitermes rotundicephalus, Parvitermes longinasus, and Velocitermes bulbus. This brings the total termite fauna in Dominican amber to four families, 17 genera, and 39 species, a number that exceeds that of the present-day fauna of Hispaniola. Biogeographical, paleoecological, and phylogenetic implications of the Dominican amber termites are discussed.

Treatise on the Isoptera of the World: VOLUME 4 TERMITIDAE (PART ONE)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 A Brief History of Termite Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Key to the Extant Families of Isoptera. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 Key to the Subfamilies of Termitidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74 Key to Genera of the Family Kalotermitidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 Diagnoses of Families and Subfamilies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79 Pest Species of Isoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133 Termite Evolution: Diversity, Distributions, Phylogeny, Fossil Record . . . . . . . . . . . . . . . . 147 Summary Classification of Isoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183 The Taxonomic Compendium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193 Nomenclatural Changes Made in This Work . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193 Museums and Repositories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196