Early Fossil History of the Termites

Proc. Natl. Acad. Sci. U.S.A.

Agosti²

2000

175

129

A worker ant preserved with microscopic detail has been discovered in Turonian-aged New Jersey amber [ca. 92 mega-annum (Ma)]. The apex of the gaster has an acidopore and, thus, allows definitive assignment of the fossil to the large extant subfamily Formicinae, members of which use a defensive spray of formic acid. This specimen is the only Cretaceous record of the subfamily, and only two other fossil ants are known from the Cretaceous that unequivocally belong to an extant subfamily (Brownimecia and Canapone of the Ponerinae, in New Jersey and Canadian amber, respectively). In lieu of a cladogram of formicine genera, generalized morphology of this fossil suggests a basal position in the subfamily. Formicinae and Ponerinae in the mid Cretaceous indicate divergence of basal lineages of ants near the Albian (ca. 105-110 Ma) when they presumably diverged from the Sphecomyrminae. Sphecomyrmines are the plesiomorphic sister group to all other ants, or they are a paraphyletic stem group ancestral to all other ants-they apparently became extinct in the Late Cretaceous. Ant abundance in major deposits of Cretaceous and Tertiary insects indicates that they did not become common and presumably dominant in terrestrial ecosystems until the Eocene (ca. 45 Ma). It is at this time that modern genera that form very large colonies (at least 10,000 individuals) first appear. During the Cretaceous, eusocial termites, bees, and vespid wasps also first appear-they show a similar pattern of diversification and proliferation in the Tertiary. The Cretaceous ants have further implications for interpreting distributions of modern ants.

Section: Discussionmentioning

confidence: 99%

A formicine in New Jersey Cretaceous amber (Hymenoptera: Formicidae) and early evolution of the ants

Proc. Natl. Acad. Sci. U.S.A.

Agosti²

2000

175

129

“…Mastotermes darwiniensis Froggatt, from northern Australia and southern New Guinea, is the sole survivor of the formerly global Mastotermitidae (Thorne et al, 2000;Wappler and Engel, 2006), and it retains striking plesiomorphic features with roaches, such as laying its eggs in a vestigial pod or ootheca. Interestingly, the Mastotermitidae as it has historically been classified was recovered as monophyletic, despite opinion to the contrary (e.g., Jarzembowski, 1981).…”

Section: Phylogenymentioning

confidence: 99%

“…100-120 Ma, with some modern subfamilies diverging in the Late Cretaceous. Though their evolutionary history has been studied less, it is now acknowledged that termites are highly modified, eusocial roaches (Cleveland et al, 1934;McKittrick, 1964;Lo et al, 2000;Deitz et al, 2003;Grimaldi and Engel, 2005;Klass and Meier, 2006), whose earliest fossils predate those of ants and bees by approximately 35 million years (Thorne et al, 2000;. Recent phylogenetic work on termites involves molecular and some morphologically based analyses of living species only (e.g., Kambhampati et al, 1996;Donovan et al, 2000;Thompson et al, 2000;Bitsch and Noirot, 2002;Klass and Meier, 2006;Inward et al, 2007aInward et al, , 2007bLegendre et al, 2008).…”

Section: Introductionmentioning

confidence: 99%

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

Engel

American Museum Novitates

Krishna

2009

198

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.

“…For example, Mastotermes is the only termite that, like cockroaches, oviposits an ootheca (egg case), although it is rudimentary (2). Numerous mastotermitid fossil specimens, corresponding to four extinct genera and about 20 species, occur from the Eocene [40 million years ago (mya)] to the Miocene (20-5 mya) of Australia, Brazil, the Caribbean, Central America, and especially Europe (3,4). The two finest preserved fossil species, extremely similar to M. darwiniensis, occur in Oligocene and Miocene amber from southern Mexico (Mastotermes electromexicus) (5) and the Dominican Republic (Mastotermes electrodominicus) (6).…”

mentioning

confidence: 99%

Spirochete and protist symbionts of a termite ( Mastotermes electrodominicus ) in Miocene amber

Wier

Dolan

Proc. Natl. Acad. Sci. U.S.A.

et al. 2002

Extraordinary preservation in amber of the Miocene termite Mastotermes electrodominicus has led to the discovery of fossil symbiotic microbes. Spirochete bacteria and wood-digesting protists were identified in the intestinal tissue of the insect. Fossil wood (xylem: developing vessel-element cells, fibers, pit connections), protists (most likely xylophagic amitochondriates), an endospore (probably of the filamentous intestinal bacterium Arthromitus ‫؍‬ Bacillus), and large spirochetes were seen in thin section by light and transmission electron microscopy. The intestinal microbiota of the living termite Mastotermes darwiniensis, a genus now restricted to northern Australia, markedly resembles that preserved in amber. This is a direct observation of a 20-million-year-old xylophagus termite fossil microbial community. Mastotermes, a genus of large tropical wood-ingesting termites, is of evolutionary importance for two reasons: (i) they harbor Mixotricha paradoxa and other amitochondriate many-genomed protists considered key to the early history of nucleated cells, and (ii) they are phylogenetically proximal to cockroaches (1). Both morphological and molecular features consistently place the single extant species (Mastotermes darwiniensis, family Mastotermitidae) as an early taxon ancestral to other termites. For example, Mastotermes is the only termite that, like cockroaches, oviposits an ootheca (egg case), although it is rudimentary (2). Numerous mastotermitid fossil specimens, corresponding to four extinct genera and about 20 species, occur from the Eocene [40 million years ago (mya)] to the Miocene (20-5 mya) of Australia, Brazil, the Caribbean, Central America, and especially Europe (3, 4). The two finest preserved fossil species, extremely similar to M. darwiniensis, occur in Oligocene and Miocene amber from southern Mexico (Mastotermes electromexicus) (5) and the Dominican Republic (Mastotermes electrodominicus) (6). The notoriously polyphagous ''living fossil'' M. darwiniensis is limited to northern Australia.Termites exhibit a complex and unique symbiosis with prokaryotic (eubacterial and archaebacterial) and eukaryotic microorganisms that live in their hindgut. The symbiotic microbiota digest cellulose to sugars and acetate and produce hydrogen, methane, and carbon dioxide (7). In addition to Mixotricha paradoxa, a giant trichomonad motile by means of a unique set of attached surface spirochetes (8, 9), M. darwiniensis harbors other Archaeprotista (class Parabasalia, those with distinctive parabasal bodies, a type of Golgi apparatus), including the two hypermastigote genera (Koruga and Deltotrichonympha), a devescovinid (Metadevescovina extranea), and a trichomonad (Pentatrichomonoides scroa) that harbors endosymbiotic methanogenic bacteria (10).Where and when did this symbiosis evolve? Here we use the remarkable preservative properties of termites in amber to seek direct paleontological evidence for the evolution of termite intestinal symbionts. Cellular and subcellular structures of amber-preserved tis...