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 that—uniquely among ants—swing 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.
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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

News of a Trigonalid

A pot of Eumenes sp., collected by Troy Bartlett in Duluth, GA

And now, to demonstrate that I am not (yet) deceased, a post.

Back on April 30th, I found an adjoining pair of potter wasp nests (likely belonging to Eumenes sp.) while on a collecting trip to Zaleski State Forest with the Ohio State University's undergraduate entomology club. At the recommendation of one of my fellow students, I saved the stout clay pots in hopes that something more interesting than the offspring of their creator would emerge.

My hope paid off just one day shy of a month later, when two distinctly non-eumenine wasps emerged, one from each pot: both larvae had been parasitized (females of the Eumenini lay a single egg in each jar; Hermes et al., 2015). Much to my pleasure, the parasitoids I had unwittingly collected were Lycogaster pullata, a member of the small, seldom-encountered family Trigonalidae.

Four genera of trigonalids are present in North America (Murphy et al., 2009). They tend to be moderately sized, chunky wasps with unreduced wing venation and >16 flagellomeres, distinguished from other apocritans of this description by finger-like ventral projections at their tarsomeres' apices and reduced ovipositors in females (Goulet & Huber, 1993). Distinctive enough to warrant their own superfamily, trigonalids are currently regarded as most akin to the Megalyroidea (Heraty et al., 2011).

The reduced ovipositor is a consequence of the wasps' distinctive biology, the convolutions of which are at least partially responsible for their rarity: rather than lay eggs directly upon or within their desired hosts, trigonalid females oviposit large quantities of minute eggs in foliage, where they are ingested by herbivorous sawfly and lepidopteran larvae. This strategy is not unique, but trigonalids add a bizarre wrinkle in that they are for the most part obligate hyperparasitoids, attacking ichneumonid or tachinid larvae that are themselves parasitoids of the insect that originally ingested the eggs (Murphy et al., 2009).

A Lycogaster pullata that emerged from the above pot

L. pullata has been reported both as a hyperparasitoid, and as facultatively employing a different stratagem—the one employed by the particular specimen I collected (Smith, 1996). Namely, it (and other trigonalids) may parasitize the larvae of vespid wasps (including eumenines) that provision their nests with caterpillars, infesting these hosts through inadvertent larval ingestion of minute trigonalid larvae that had hatched within said caterpillars. Evidently, one should thoroughly chew one's food.

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Goulet, H and Huber, J. T. (1993). Hymenoptera of the World: an Identification Guide to Families. Ottawa: Agriculture Canada.

Heraty, J.; Ronquist, F.; Carpenter, J. M.; Hawks, D.; Schulmeister, S.; Dowling, A. P.; Murray, D.; Munro, J.; Wheeler, W. C.; Schiff, N.; and Sharkey, M. (2011). Evolution of the hymenopteran megaradiation. Molecular Phylogenetics & Evolution, 60, 73-88. 

Hermes, M. G.; Araujo, G. and Antonini, Y. (2015). On the nesting biology of eumenine wasps yet again: Minixi brasilianum (de Saussure) is a builder and a renter… at the same time! (Hymenoptera, Vespidae, Eumeninae). Revista Brasileira de Entomologia, 59(2), 121-142. Retrieved 5/30/16 from http://www.sciencedirect.com/science/article/pii/S0085562615000424


Murphy, S. M.; Lill, J. T.; and Smith, D. R. (2009). A scattershot approach to host location: uncovering the unique life history of the trigonalid hyperparasitoid Orthogonalys pulchella (Cresson). American Entomologist, 55, 82-87. Retrieved 5/29/16 from https://www.researchgate.net/publication/221959174_A_scattershot_approach_to_host_location_uncovering_the_unique_life_history_of_the_trigonalid_hyperparasitoid_Orthogonalys_pulchella_Cresson


Smith, D. R. (1996). Trigonalyidae (Hymenoptera) in the eastern United States: seasonal flight activity, distributions, hosts. Proceedings of the Entomological Society of Washington, 98(1), 109-118.