TY - JOUR
AU1 - Berenbaum,, May
AB - View largeDownload slide “Yes, I’m an alien extraterrestrial termite here to exploit your planet’s natural resources. Take me to your cedar ...” View largeDownload slide “Yes, I’m an alien extraterrestrial termite here to exploit your planet’s natural resources. Take me to your cedar ...” The heavenly body called Pluto was discovered by Clyde Tombaugh in 1930 and, amid much fanfare, was designated as the ninth planet in our solar system. As such, it was immortalized in mnemonic devices designed to help elementary school students remember the names of the planets in order of their distance from the sun. That’s how Pluto came to be the “Pizzas” in “My Very Educated Mother Just Served Us Nine Pizzas.” Always small relative to the other eight, Pluto’s status as a full-fledged planet came into question in 2005, when CalTech astronomers discovered Eris, a celestial object larger than Pluto but with an orbit at an even greater distance from the sun than Pluto’s orbit. This finding, among others, motivated the International Astronomical Union (IAU) to rethink its definition of a planet and to create a new category, “dwarf planet,” into which Pluto, Eris, and three other bodies were placed. Now, very educated mothers are apparently just serving unspecified quantities of noodles or nachos. Objectively, there’s not a lot to commend Pluto, other than rock and ice. At its greatest distance from the sun, temperatures can dip below -240 °C (-400 °F), so cold that methane gas freezes, forming snowcaps on mountains and methane ice dunes (Telfer et al. 2018). Notwithstanding, people around the world were distraught when Pluto’s status was downgraded to “dwarf planet.” The Illinois Senate, which in 2015–2016 couldn’t manage to approve a state budget, could nonetheless effortlessly pass a resolution in 2009 declaring 19 March “Pluto Planet Day” in defiance of IAU. According to the text of SR0046 of the 96th General Assembly, WHEREAS, Clyde Tombaugh, discoverer of the planet Pluto, was born on a farm near the Illinois community of Streator, and...is so far the only Illinoisan and only American to ever discover a planet...be it RESOLVED, BY THE SENATE OF THE NINETY-SIXTH GENERAL ASSEMBLY OF THE STATE OF ILLINOIS, that as Pluto passes overhead through Illinois’ night skies, that it be reestablished with full planetary status, and that March 13, 2009 be declared “Pluto Day” in the State of Illinois in honor of the date its discovery was announced in 1930 (https://tinyurl.com/y88hez5r). By contrast, the Illinois legislature remained unmoved when another iconic scientific entity was demoted, shattering a generation of entomologists who experienced the loss of an entire order of insects, not to extinction but to taxonomy. For centuries, biologists have known about termites but have struggled to find a taxonomic place for the more than 3,500 species that, at least superficially, aren’t quite like any other insects. A possible early attempt to classify termites appears in Etymologia, a 20-volume encyclopedia written by St. Isidore of Seville in 630 CE (Barney et al. 2006). In the fifth chapter de verminibus (vermin), St. Isidore defined “vermin” as “animals that are generated for the most part from flesh or wood or some earthy substance, without any sexual congress.” Of these, teredo were “wood vermin that eat by grinding.” Although today the genus Teredo houses shipworms (marine bivalves that do indeed grind their way through submerged wood), etymological history suggests that St. Isidore may have intended teredo to mean “termites.” The Oxford English Dictionary, for example, dates the origin of the English word “termite” in 1849 as a back-formation from the three-syllable plural word termites, derived in turn from the Late Latin term termes, “woodworm,” reflective of the Latin terere, “to rub, wear, or erode.” If it’s surprising that the English word “termite” is so new, it’s worth mentioning that, before the 19th century, they were best known as white ants (a name that persists even today). Brullé (1832) coined the name “Isoptera”; that’s the term used by Hagen (1858) in his Monographie der Termiten, in which he perceptively “insisted on the close affinity of the Termitidae with the Blattidae, or cockroaches” (Wheeler 1904). Since 1858, termites, as Isoptera, have variously been grouped as a suborder of long-gone grab-bag orders including Corrodentia, Pseudoneuroptera, and Platyptera. Borner (1904) may have been the first to upgrade Isoptera to ordinal status, where the group has remained until about a decade ago. Termite taxonomic troubles began to stir a half-century ago, when F. A. McKitterick (1964) pointed out morphological similarities between some termites and Cryptocercus wood roaches. Phylogenetic studies, mostly carried out with limited taxon sampling, failed to yield sufficient persuasive evidence for a taxonomic reshuffling, but, on 5 April 2007, Inward et al. (2007) sampled more than 100 in-group Dictyoptera species, along with 11 outgroups, using five gene loci to produce a phylogenetic tree showing termites nesting (as it were) within the cockroaches and with Cryptocercus as the termite sister group. These authors interpreted their findings as justification for the “death of an order,” Isoptera, and for combining all termites into a single family, Termitidae. This suggestion met with resistance in the form of a plea published on 14 August 2007 by Lo et al. to “Save Isoptera” based on multiple criteria, not least of which was that demoting all termites to a single family (and therefore demoting seven well-established families, including one already called Termitidae, to subfamily status) would be a hard sell and would create chaos for “future researchers who will in perpetuity need to reconcile past with future designations.” Moreover, in view of still-unresolved phylogenetic associations among cockroaches, these authors proposed instead that “Isoptera be retained as an unranked name within Blattaria,” with future taxonomic unit options including suborder, infraorder, superfamily, or epifamily status. This paper in turn elicited a much less provocatively titled response (i.e., “Response to Lo et al.”) by Eggleton et al. (2007), who, in the spirit of compromise, proposed “the new epifamily Termitoidae for the termites…and the new epifamily Cryptocercoidae…for the Cryptocercus lineage… and the new epifamily Blattoidae…for the lineage which includes the Blattidae.” These names, along with some suprafamilial and infraordinal names, and at least one nanordinal name (e.g., Engel 2011), all appear in post-2007 publications with varying degrees of frequency, but my approach to this problem is never to work on termites or to write an introductory entomology textbook. Truth be told, I’m still not exactly sure what an epifamily is, and I’m even less certain about nanorders, given that the only definition I could find, from Wiktionary.org, explained that a nanorder is “a taxon below order and hypoorder” (which begs the question of what a hypoorder is). However confusing entomologists have found termites, the general public has been remarkably consistent in its assessment of termites—no one likes them. Their remarkably sophisticated social behavior has inspired surprisingly few metaphors or allegories over the centuries. In fact, while “busy bees” and “industrious ants” are admired, termites are almost universally reviled, not unreasonably given that the one thing everyone seems to know about them is that they eat wood, a habit that even today costs humanity up to 40 billion dollars a year (Rust and Su 2012, Govorushko 2018). What I find sad is that the one thing people think they know about them isn’t entirely accurate; most termites do consume cellulose, but of the more than 3,000 species, only about 83 species attack and damage wooden structures. Moreover, some higher termites eat soil or grass, and more than 100 species feed on mammal dung (although in all likelihood, this dietary habit also isn’t going to endear them to most people any more than eating houses does). In recent years, there’s one more fact that people seem to have willingly embraced about termites—it’s that they are among the planet’s major contributors to producing greenhouse gases, particularly methane. Estimates put their contribution to global methane production at millions of tons (Berenbaum 2000). A Google search with the phrase “termite flatulence” turns up more than 100,000 hits (including a song by the science rock band Bungee Jumpin’ Cows: https://www.youtube.com/watch?v=ynKBnAn-mOs). But laying global climate change at the feet (or anuses) of termites isn’t quite fair; according to Nauer et al. (2018), half of the methane produced by at least some species of termites in the process of breaking down cellulose is oxidized before it’s even emitted. As it turns out, the walls of the mounds of these species house methanotrophic bacteria that metabolize the methane before it can be released into the atmosphere. So, much of what people believe to be true about termites isn’t, and, beyond those few facts, most people don’t really know anything else about them. That said, I’m not sure that greater familiarity with termite lifestyle, particularly their social behavior, would necessarily lead to greater respect and admiration. Autothysis (explosive defecation; Berenbaum 2017), phragmosis (use of the enlarged head to block entry holes in the nest wall; Scholtz et al. 2008), and proctodeal feeding (anus-to-mouth food delivery), all hallmarks of termite social life, sound more like a list of disorders in a psychology textbook rather than social behaviors worthy of emulation. All of the admittedly limited pop-culture depictions of termites I’ve been able to find include eating wood as a plot element. Oddly, however, a disproportionate number of those depictions suggest that termites don’t even belong on Earth in the first place. For example, there’s the 1952 Woody Woodpecker cartoon Termites from Mars, in which Earth is invaded by squadrons of alien termites who proceed to consume every available piece of furniture until Woody Woodpecker figures out how to capture them (“turning the tables” on them, as it were). There’s also the episode from the 1980 reboot of the television cartoon show Super Friends (based on the DC Comics franchise), titled “Termites from Venus,” in which Earth is threatened by an invading force of termites. With respect to features, the 1995 film Alien Apocalypse depicts a takeover of Earth by alien termites, who enslave humans in order to harvest wood, supplies of which they’ve depleted on their home planet. In contrast with the earlier shorts, I don’t recall that the planet of origin for these alien isopteran invaders is identified. Now that I think about it, given the relationship between termites and methane, the possibility exists that they might have come from Pluto—maybe the presence of an alien termite civilization could explain how those methane snowdrifts got there in the first place. May Berenbaum is a professor and head of the Department of Entomology, University of Illinois, 320 Morrill Hall, 505 South Goodwin Avenue, Urbana, IL 61801. Currently, she is studying the chemical aspects of interaction between herbivorous insects and their hosts. References Cited Barney , S.A. , W.J. Lewis , J.A. Beach , and O. Berghof . 2006 . The etymologies of Isidore of Seville . Cambridge University Press , NY . Berenbaum , M.R. 2000 . Buzzwords: a scientist muses on sex, bugs, and rock ‘n’ roll . Joseph Henry Press , Washington, DC . Berenbaum , M.R. 2017 . Buzzwords: explosive issues . American Entomologist 63 : 1 – 5 . Google Scholar Crossref Search ADS Borner , C. 1904 . Zur Systematik der Hexapoden . Zoologischer Anzeiger 27 : 511 – 533 . Brullé , G.A. 1832 . Expédition Scientifique de Morée. Section des Sciences Physiques. Tome III. Partie 1. Zoologie. Deuxième Section–Des Animaux Articulés. Levrault, Paris (cited in Wheeler 1904) . Eggleton , P. , G. Beccaloni , and D. Inward . 2007 . Response to Lo et al . Biology Letters . 3 : 564 – 565 . Google Scholar Crossref Search ADS Engel , M.S. 2011 . Family-group names for termites (Isoptera), redux . Zookeys 148 : 171 – 184 . Google Scholar Crossref Search ADS Govorushko , S. 2018 . Economic and ecological importance of termites: a global review . Entomological Science 2018. doi:10.1111/ens.12328. Hagen , H. 1858 . Monographie der Termiten . Linnaea Entomologica 12 : 1 – 342 (cited in Wheeler 1904). Inward , D. , G. Beccaloni , and P. Eggleton . 2007 . Death of an order: a comprehensive molecular phylogenetic study confirms that termites are eusocial cockroaches . Biology Letters 3 : 331 – 335 . Google Scholar Crossref Search ADS PubMed Lo , N. , M.S. Engel , S. Cameron , C.A. Nalepa , G. Tokuda , D. Grimaldi , O. Kitade , K. Krishna , K.-D. Klass , K. Maekawa , T. Miura , and G.J. Thompson . 2007 . Save Isoptera: a comment on Inward et al. Proceedings of the Royal Society 3: Biology Letters . https://royalsocietypublishing.org/doi/full/10.1098/rsbl.2007.0264. McKitterick , F.A. 1964 . A contribution to the understanding of cockroach-termite affinities . Annals of the Entomological Society of America 58 : 18 – 22 . Google Scholar Crossref Search ADS Nauer , P.A. , L.B. Hutley , and S.K. Arndt . 2018 . Termite mounds mitigate half of termite methane emissions . Proceedings of the National Academy of Sciences 115 : 13306 – 13311 . Google Scholar Crossref Search ADS Rust M.K. , and N.-Y. Su . 2012 . Managing social insects of urban importance . Annual Review of Entomology 57 : 355 – 375 . Google Scholar Crossref Search ADS PubMed Scholtz , O.L. , N. MacLeod , and P. Eggleton . 2008 . Termite soldier defence strategies: a reassessment of Prestwich’s classification and an examination of the evolution of defence morphology using extended eigenshape analyses of head morphology . Zoological Journal of the Linnean Society 153 : 631 – 650 . Google Scholar Crossref Search ADS Telfer , M.W. , E.J.R. Parteli , J. Radebaugh , R.A. Beyer , and T. Bertran . 2018 . Dunes on Pluto . Science 360 : 992 – 997 . Google Scholar Crossref Search ADS PubMed Wheeler , W.M. 1904 . The phylogeny of termites . Biological Bulletin 8 : 29 – 37 . Google Scholar Crossref Search ADS © The Author 2019. Published by Oxford University Press on behalf of the Entomological Society of America. This article is published and distributed under the terms of the Oxford University Press, Standard Journals Publication Model (https://academic.oup.com/journals/pages/open_access/funder_policies/chorus/standard_publication_model)
TI - Out of Order
JF - American Entomologist
DO - 10.1093/ae/tmz012
DA - 2019-03-12
UR - https://www.deepdyve.com/lp/oxford-university-press/out-of-order-84wIKzfsdR
SP - 3
VL - 65
IS - 1
DP - DeepDyve
ER -