Tuesday, May 2, 2017

A Tropical Monstrosity in Ohio: the Owl Moth

On October 9th of last year, I happened across this beauty in the breezeway of my apartment in Columbus, OH. The specimen was dead, but exceedingly fresh. It had been a chilly night, with a low near 40; she had evidently perished just a few hours beforehand.

Not being a lepidopterist, I really had little idea of what I had found, other than that it was an erebid moth not unlike the Black Witch (Ascalapha odorata). However, further investigation demonstrated that this enormous specimen (wingspan 15 cm., as shown) was in fact a female Owl Moth (Thysania zenobia), along with the Black Witch a member of the Thermesiini. 

Much to my pleasure, the find is unusual. T. zenobia ranges throughout the eastern Americas south to Rio Grande do Sul (Specht et al., 2004),  and occurs in the Greater Antilles and Galapagos Islands (Roque, 1999): according to the Ohio Lepidopterists' Society, it has only been collected in the state on five occasions, none of which occurred in Franklin County (Rings et al., 1992).

Indeed, the animal is a tropical stray. Their caterpillars' cassiine host plants (Specht et al., 2004) do not grow natively in the Lower 48. Early authors even proposed that its appearance in the northern United States was due to human mediation (Felt, 1907). However, given that they have cropped up in good condition in Wisconsin (Ziemer, 1949), New England (Farquhar, 1938) and Ohio (Rings et al., 1992) on occasion, it is now clear that the moths fly northwards under their own power; their native presence in the Galapagos would also indicate excellent dispersal capabilities.

One lepidopterist wrote that he was "unprepared for" collecting a live T. zenobia in Ontario while sugaring for underwings (Catocala spp.): "However, the monster was taken" (Kilman, 1889). I regret that I did not have the same good fortune to see this T. zenobia in life. She would have brought to my mind Albert Giraud's poem cycle "Pierrot Lunaire" (here translated by Cecil Gray):

 And from heaven earthward bound
Downward sink with somber pinions
Unperceived, great hordes of monsters
On the hearts and souls of mankind...
Gloomy, black giant moths.     

Farquhar, D. W. (1938). The Lepidoptera of New England (Doctoral dissertation). Harvard University, Cambridge.

Felt, E. P. (1906). 22nd Report of the State Entomologist on injurious and other insects of the state of New York 1906. New York State Museum Bulletin, 110, 39-44. 

Kilman, A. H. (1889). Correspondence: a rare moth. The Canadian Entomologist, 21(1): 240. 

Rings, R. W.; Metzler, E. H.; Arnold, F. J.; and Harris, D. H. (1992). The Owlet Moths of Ohio: Order Lepidoptera, Family Noctuidae. Bulletin of the Ohio Biological Survey, 9(2), vi+219 pp.

Roque, L. (1999). Two large tropical moths, Thysania zenobia (Noctuidae) and Cocytius antaeus (Sphingidae) colonize the Galapagos Islands. Journal of the Lepidopterists' Society 53(3), 129-130.  

Specht, A.; Silva, E. J. E.; and Link, D. (2004). Noctuídeos (Lepidoptera, Noctuidae) do Museu Entomológico Ceslau Biezanko, Faculdade de Agronomia Eliseu Maciel, Universidade Federal de Pelotas, RS. Revista Brasileira de Agrociência, 10(4), 389-409.

Ziemer, S. E. (1948). Erebus odora and Thysania zenobia in Wisconsin. The Lepidopterist's News, 2(3), 25-36.

Sunday, March 5, 2017

Announcing the Iberobaeniidae

Holotype of Urtea graeca, mustachioed with silly maxillary palporgans (Paulus, 2004)
Undoubtedly, Europe is one of the regions most thoroughly picked-over by descriptive taxonomists, particularly in the case of an insect group as comparatively popular with entomologists as the beetles (Coleoptera). Nonetheless, unexpected oddities in the beetle department do crop up in the western Palearctic from time to time. The incomparable Crowsoniella relicta (placed in a monotypic family) is known from soil samples collected from the base of a single Italian chestnut tree (Pace, 1975), while Urtea graeca (a genus and species known from a single locality in northern Greece) is the only European representative of the oddball Atractocerinae (Lymexylidae), a wood-boring taxon that aside from a single Japanese species (Arractocetus nipponicus) is restricted to the tropics and southern hemisphere (Paulus, 2004).

Male Iberobaenia minuta; scale bar=1 mm.
Last December, an entirely new beetle family was described from the mountains of southern Spain, including the single genus Iberobaenia and two species distinguished upon the basis of the male genitalia (Bocak et al., 2016). What appears to be a third, still-undescribed species is known only from a third-instar larva. The circumstances of their discovery warrant some explanation: the larvae were collected for soil biodiversity surveys in southern Spain and noticed when metagenomic analysis of these samples revealed mitochondrial sequences within the Elateroidea (click beetles and kin) that did not nest within any family; these sequences allowed linkage of these larvae with adult male specimens, which had been collected by sweeping foliage. Interestingly, the relative positioning of protein-coding and ribosomal genes in the beetle's mitogenome had been altered from the putatively ancestral condition, this being the first such rearrangement documented in the Coleoptera (Andujar et al., 2016) and an abnormal occurrence for any insect, which tend to be conservative with respect to mitochondrial gene order (Cameron, 2014).

In appearance, Iberobaenia is clearly elateroid in both known ontogenetic stages. The tiny, soft-bodied male imagos resemble, in their reduced mouthparts and lack of elytral costae or tubercles, other elateroid lineages that have undergone miniaturization relative to the superfamily's norm. Their habitus is most comparable to such genera as Paradrilus (Omalisidae) and Antennolycus (Lycidae), but iberobaeniids differ from these in their hypognathous* mouthparts (in contrast to omalisids), lack of a pronotal margin (differing from lycids and non-paradriline omalisids in this respect), and uncompressed legs (here being distinct from otherwise comparable lycids). Larvae are not dorsoventrally flattened and lack urogomphi (unlike their lycid counterparts), with robust convergent mandibles (Bocak et al., 2016).

Female Duliticola paradoxa (Lycidae): image by Bernard Dupont
A taxonomically comprehensive molecular phylogenetic analysis of the Elateroidea, using two loci each from the nuclear and mitochondrial genomes, retrieved Iberobaenia as the sister group of the Lycidae (net-winged beetles). Male miniaturization in the genus is noteworthy in terms of speculation on iberobaeniid biology. Elateroids with males of this dwarfish, soft-bodied description tend to be sexually dimorphic, and if comparison of Iberobaenia to these better-known cousins is valid, its unknown females are probably larviform crustacean-like things far larger than their paramours. This syndrome has evolved on multiple occasions throughout the Elateroidea, with examples of flightless lycid females looking more like denizens of the Sea of Corruption than inhabitants of the East Indies (see above).

While Iberobaenia is certainly distinctive in that it cannot be confidently placed in any elateroid family, I personally doubt that its assignment to an entirely new one will hold under future acquisition of more genetic data. From a morphological standpoint, the proposed Iberobaeniidae lacks any nomothetic characters, meaning that we must rely upon the phylogenetic analysis of Bocak et al. (2016) for justification of its ranking as family. This phylogeny (with Elaterophanes and Aphodiites, which are respectively the oldest known elateriforms and scarabaeiforms, as calibration priors) suggests that Iberobaenia branched from its most recent common ancestor in the mid-Jurassic: I wonder how much water mitogenomes (from which half of the data used in this analysis were drawn) hold at that timescale, given their famously rapid rate of mutation (Brown et al., 1979). 

As with so many things in our world, more data would be welcome.

*With mouthparts directed downwards.
Superficially cercus-like processes on the terminal segments of some beetle larvae.
One particular feature that is unique to the taxon in question.

Andujar, C.; Arribas, P.; Linard, B.; Kundrata, R.; Bocak, L. and Vogler, A. P. (2016). The mitochondrial genome of Iberobaenia (Coleoptera: Iberobaeniidae): first rearrangement of protein-coding genes in the beetles. Mitochondrial DNA Part A: DNA Mapping, Sequencing, & Analysis; 28(2), 156-158. Retrieved 3/3/17 from http://www.tandfonline.com/doi/abs/10.3109/19401736.2015.1115488?journalCode=imdn21http://www.tandfonline.com/doi/abs/10.3109/19401736.2015.1115488?journalCode=imdn21

Bocak, L.; Kundrata, R.; Fernandez, C. A.; and Vogler, A. P. (2016). The discovery of Iberobaeniidae (Coleoptera: Elateroidea): a new family of beetles from Spain, with immatures detected by environmental DNA sequencing. Proceedings of the Royal Society of Britain, 283, 1-7. Retrieved 3/5/17 from https://pdfs.semanticscholar.org/e4da/1e1282ecb9a5b438ed0de3bb9193dc1ea10e.pdf

Brown, W. M.; George, M. Jr.; and Wilson, A. C. (1979). Rapid evolution of animal mitochondrial DNA. Proc. Nat. Acad. Sci. USA, 79, 1967-1971. 

Cameron, S. L. (2014). Insect mitochondrial genetics: implications for evolution and phylogeny. Annual Review of Entomology, 59, 95-117. 

Pace, R. (1975). An exceptional endogeous beetle: Crowsoniella relicta n. gen. n. sp. of Archostemata Tetraphaleridae from central Italy. Bolletino del Museo Civico di Storia Naturale, Verona; 2, 445-458.

Paulus, H. F. (2004). Urtea graeca nov. gen. et nov. spec., der erste Vertreter tropischen Atractocerinae in Europa sowie eine Beschreibung von Hymaloxylon aspoecki nov. spec. aus Yunnan (China) (Coleoptera, Cucujiformia, Lymexylidae, Atractocerinae nov. status). Denisia, 277-290. Retrieved 2/11/17 from http://www.zobodat.at/pdf/DENISIA_0013_0277-0290.pdf 

Saturday, January 28, 2017

A Flagrant Exercise in Laziness Concerning Chalcidoids

After several years of deferrals, I finally got around this May to penning a post for the Ohio State University's Triplehorn Insect Collection. I'll take this opportunity to shamelessly self-plagiarize.

I have always been fascinated by insects, but it was not until, at age 15, I took a week-long field insect taxonomy course at the Ohio State University’s Stone Laboratory. There, I learned the conventions of arthropod collection and preservation and something of proper curatorial practices. Ever since, I have steadily accumulated a collection with pretensions made modest by the limited resources of a teenager; and collection and identification remain an exceedingly enjoyable activity for me.

Therefore, it was only natural that I gravitated to the Triplehorn Insect Collection upon commencing my undergraduate career. Sorting unidentified specimens was easily my favorite task there. Although the collection contains the full range of insect diversity, those specimens that I was tasked with identifying almost always belonged to the order Hymenoptera—often casually referred to as “ants, bees, and wasps”, but technically including far more taxa than simply those that happen to have colloquial names. Being one of the four most diverse insect orders, the variety of Hymenoptera is considerable: and I encountered much of their phylogenetic span through this process, while becoming intimately familiar with Goulet and Huber’s tome Hymenoptera of the World: an Identification Guide to Families (1993) (PDF), the monochromatic line drawings within which—printed on thick, coarse paper—have caused hymenopterists to nickname it “the coloring book”.

Paratype of Dicopomorpha echmepterygis, male (Huber & Noyes, 2013)
As an exemplar of the many taxa with which I thereby became familiar, I have chosen to briefly discuss the superfamily Chalcidoidea herein. These parasitoid wasps are one of those aforementioned many prominent insect taxa that have no name in the vernacular—understandable, given that the vast majority of these particular parasitoids are a few millimeters in length or less. (Indeed, the smallest insect known to science—the 0.13-mm.-long male of Dicopomorpha echmepterygis [Mockford, 1997]—is a chalcidoid.)

Unidentified pteromalid from the Amazon Basin, photographed by yours truly (©OSU)
This diminution has also resulted in a lack of taxonomic attention from entomologists, and chalcidoid systematics is by consequence a frustratingly opaque matter—something one is immediately impressed with while attempting to identify the miniscule things: keys are peppered with qualifiers like “usually” and “most”, not to mention annotated with lengthy footnotes elucidating the exceptions to each couplet. The fundamental problem at hand, as Goulet and Huber point out, is that chalcidoid families are often defined by combinations of characters, as opposed to singular traits that are unique to that taxa and none other (autapomorphies, in cladistic terms). This has resulted in a superfamily littered with taxa whose boundaries are under constant debate (e.g., the Agaonidae) or that do not hold up to scrutiny whatsoever (the grossest wastebasket taxon of flagrant wastebasket taxa, the Pteromalidae).

Female Sycoryctes cyathistipula (Agaonidae: Sycoryctinae) (©Iziko Museums of South Africa)
Chalcidoids are hardly deserving of this neglect, considering their ecological and numerical diversity (they possibly constitute 10% of all insect species; Noyes, 2003). I would have impartially respected this significance regardless of my work at the Collection, but parsing through unit tray after unit tray of nigh-microscopic specimens representing untold numbers of species—each one a chalcidoid—gave me a concrete grasp of that abstraction.

I still have strong visual impressions of many of them: the subtly turquoise, spatula-shaped abdomen I swiftly came to associate with the Tetracampidae; the minute serrations on the inner rims of a stocky chalcidid’s femora, making its thighs appear like chitinous razors; the oar-like forewings of many an insubstantial mymarid, fringed with haloes of setae; the metallic, spindle-shaped abdomen that accounted for two-thirds the length of a sycoryctine. I am not the only one to have thought them often quite showy under sufficient magnification: Alexandre A. Girault, a notoriously verbose chalcidologist, spoke of the tiny wasps as “gem-like inhabitants of the woodlands, by most never seen or dreamt of” (Thomer & Twidale, 2014).

Suffice it to say, without my work at the Collection, I would not have seen nor dreamt of so many chalcidoids.
Huber, J. T. and Noyes, J. S. (2013). A new genus and species of fairyfly, Tinkerbella nana (Hymenoptera, Mymaridae), with comments on its sister genus Kikiki , and discussion on small size limits in arthropods. Journal of Hymenoptera Research, 32, 17-44. Retrieved 1/6/17 from http://jhr.pensoft.net/articles.php?id=1635

Mockford, E. L. (1997). A new species of Dicopomorpha (Hymenoptera: Mymaridae) with diminutive, apterous males. Annals of the Entomological Society of America, 90, 115-120.

Noyes, J. S. (2003). Universal Chalcidoidea Database. Retrieved 5/18/16 from http://www.nhm.ac.uk/our-science/data/chalcidoids/introduction.html
Thomer, A. K. and Twidale, M. B. (2014). How Databases Learn. In: iConference 2014 Proceedings (pp. 827-833). Retrieved 4/8/15 from https://www.ideals.illinois.edu/bitstream/handle/2142/47268/409_ready.pdf?sequence=2

Wednesday, July 27, 2016

Yet More Haidomyrmecine Strangeness

Recently, I ran across the artwork shown below, depicting an ant (Formicidae) by the name of Ceratomyrmex ellenbergi; according to the author, the creature is known in fossil form from mid-Cretaceous Burmese amber. Since I found the supposed reconstruction on a forum nominally devoted to speculative evolution, I naturally assumed the insect to be fictitious. Not biologically implausible, by any means: but that seta-covered projection on the head, nearly touched by vertical scythe-mandibles? Give me a break. Clearly, this was merely an admirable thought experiment, too outlandish to be genuine.

Impression of Ceratomyrmex ellenbergi by Joschua Knüppe
Of course, much to my pleasure and chagrin, Ceratomyrmex is quite real (Perrichot et al., 2016): and no artistic license was taken. It is a member of the Haidomyrmecini, a basal tribe of the Formicidae known only from the mid-Cretaceous, and already characterized by unique cephalic equipment consisting of a setose median lobe (which could be interpreted as an extension the clypeus; Barden & Grimaldi, 2013) directly betwixt the antennae, scythe-shaped mandibles thatuniquely among antsswing on a vertical axis, and a singularly elongated face to accommodate these mandibles. 

It appears that Ceratomyrmex represents an exaggeration of these haidomyrmecine traits. The elevated setose portion of the clypeus* is extended forwards and to a tremendous length (hence the nickname "unicorn ant") and has been warped into a spatulate shape, with pegs restricted to its spoon-like tip and setae running the process' length. Just below the base of the antennal sockets emanate trigger hairs, which are correspondingly longer than in any other haidomyrmecine: indeed, they are more like whiskers than anything else (Perrichot et al., 2016). 

Anochetus sp. (Ponerinae) with mandibles in striking position; photographed by Alex Wild (who else?)
As the term "trigger" would suggest, it has been imputed that haidomyrmecines were ambush predators with a "trap-jaw" strategy, much like living Anochetus or Myrmoteras; dissimilar to the likes of these, however, the mandibles would have snapped upwards upon triggering, pressing prey items against the spatulate clypeal process, the setae upon which would have provided gripping capability. While the majority of modern ants which conform to a "trap-jaw" habitus are generalist predators, the fact that the armature of Ceratomyrmex would have been useless for capturing prey items below a certain size suggests a considerable degree of dietary specialization (Perrichot et al., 2016). Of course, it is anyone's guess as toward what prey items this remarkable specialization was directed.

Without a doubt, we can add the "unicorn ant" to the long list of insects that we wish still lived  on this planet.

*An insect mouthpart articulating with the frons (i.e., face), from which emanates the labrum.

Barden, P. and Grimaldi, D. (2013). A new genus of highly specialized ants in Cretaceous Burmese amber (Hymenoptera: Formicidae). Zootaxa, 3681(4), 405-412. Retrieved 12/24/15 from http://www.mapress.com/zootaxa/2013/f/zt03681p412.pdf 

Perrichot, V.; Wang, B.; and Engel, M. S. (2016). Extreme Morphogenesis and Ecological Specialization among Cretaceous Basal Ants. Current Biology, 26, 1468-1472. Retrieved 7/27/16 from https://www.researchgate.net/publication/303544013_Extreme_Morphogenesis_and_Ecological_Specialization_among_Cretaceous_Basal_Ants