Assassin ate

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I came upon this interesting scene last month while hiking through Allen David Broussard Catfish Creek Preserve State Park, which preserves some of the highest quality remnants of sand scrub habitat on the Lake Wales Ridge of central Florida. The spider seems to be Peucetia viridans (green lynx spider), widespread across the southern U.S. and distinguished by its bright transparent green color with red spots and black spines (Emerton 1961). These largest of North American lynx spiders hunt diurnally on low shrubs with an agility excelled only by the jumping spiders (Salticidae) and aggressively attack their insect prey. In this case, the prey is one of the so-called “bee assassins” of the genus Apiomerus (Hemiptera: Reduviidae). The common and generic names of these insects both derive from their habit of preying upon bees, not only on flowers but also by ambushing them at nest entrances, although other insects are preyed upon as well. Ironically, this particular assassin himself got ate.

An interesting situation was uncovered while I tried to determine which species of Apiomerus was represented by the prey. By virtue of its pale ventrals with the front and hind margins black, it keys to A. spissipes in a literature-based key to Florida Reduviidae (Bierle et al. 2002) – one of two species considered widely distributed across the eastern U.S. In reality, however, it appears that this individual represents another species named almost 30 years ago but which remains officially undescribed. As explained in this BugGuide post by Daniel Swanson, the genus was revised by Berkeley grad student Sigurd Leopold Szerlip in partial fulfillment of the requirements for a Ph.D., who proposed a number of taxonomic acts including the description of 19 new species. Among these were eastern U.S. populations to which the name A. spissipes had been applied, with those in Florida being described as the new species “A. floridensis“. However, dissertations do not meet the criteria of publication according to Article 8 of the International Code of Zoological Nomenclature (ICZN 1999), and none of the dissertation was formally published. Thus, “A. floridensis” remains an invalid, unpublished name.  This is a most unfortunate situation, as Swanson considers the dissertation to be well done.  It is not only names, but important information about life histories and detailed genitalic studies that remain unavailable to the scientific community as well.  What are the nomenclatural impacts of this work remaining unpublished?  Is this as much a failure by the advising professor as by Szerlip himself?  What ethical considerations would need to be addressed in order for it to be published in absentia, or is this even possible?

Photo details: Canon 100mm macro lens on Canon EOS 50D (manual mode), ISO-100, 1/250 sec, f/13, MT-24EX flash 1/2 power through diffuser caps.

REFERENCES:

Bierle, S., E. Dunn, S. Frederick, S. Garrett, J. Harbison, D. Hoel, B. Ley and S. Weihman. 2002. A literature-based key to Reduviidae (Heteroptera) of Florida (assassin bugs, and thread-legged bugs). Unpublished manuscript, University of Florida, Department of Entomology and Nematolgy, Insect Classification ENY 4161/6166, 18 pp.

Emerton, J. H. 1961. The Common Spiders of the United States. Dover Publications, Inc., N.Y., xx + 227 pp.

International Commission on Zoological Nomenclature [ICZN]. 1999. International Code of Zoological Nomenclature, 4th Edition. The International Trust for Zoological Nomenclature, c/o Natural History Museum, London. xxix + 306 pp.

Szerlip, S. L. 1980. Biosystematic revision of the genus Apiomerus (Hemiptera: Reduviidae) in North and Central America. Unpublished Ph.D. thesis, University of California, Berkley, CA.

Copyright © Ted C. MacRae 2009

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Millipede assassin bug

Ectrichodia crux

I continue the hemipteran theme begun in the last post with this photograph I took in South Africa below the Waterberg Range in Northern (now Limpopo) Province. I recognized them as members of the family Reduviidae (assassin bugs), and since to my knowledge species in this family are exclusively predaceous (except for the so-called “kissing bugs” of the mostly Neotropical subfamily Triatominae, large distinctive bugs that feed exclusively on vertebrate blood), I found what I took to be a case of scavenging on a dead millipede to be rather curious.  It had rained the previous evening, resulting in a burst of millipede (and insect) activity that night, and this scene was rather commonly encountered the following morning. Of course, appearances can be deceiving, and it turns out that I actually was witnessing predation – and a most unusual case at that.  The individuals in this photo represent Ectrichodia crux (millipede assassin bug), a common species in many parts of southern Africa.  Although nearly 500 species of assassin bugs are known from the region (Reavell 2000), E. crux is easily recognizable due to its large size (adults measure up to 22 mm in length), stout form, and coloration – shiny black, with a distinctive black cross incised on its dull yellow thorax and with yellow abdominal margins (Picker et al. 2002). The nymphs as well are distinctive – bright red with black wing pads. Clearly, these insects are advertising something.

Ectrichodia crux belongs to the subfamily Ectrichodiinae, noted for their aposematic coloration – often red or yellow and black or metallic blue, and as specialist predators of Diplopoda (Heteropteran Systematics Lab @ UCR).  Species in this subfamily are most commonly found in leaf litter, hiding during the day under stones or amongst debris and leaving their shelters at night in search of millipedes (Scholtz and Holm 1985). They are ambush predators that slowly approach their prey before quickly grabbing the millipede and piercing the body with their proboscis, or “beak.”  Saliva containing paralytic toxins and cytolytic enzymes is injected into the body of the millipede to subdue the prey and initiate digestion of the body contents, which are then imbibed by the gregariously feeding assassin bugs.

Millipedes employ powerful chemical defenses – primarily benzoquinones and sometimes hydrogen cyanide gas as well, which are discharged from specialized glands along the millipede’s body – to protect themselves from predation.  Thus, specialized predation of millipedes is a niche that has been exploited by relatively few predators, and little is known about the mechanisms used for circumventing these defenses. The recently reported millipede specialist, Deltochilum valgum (order Coleoptera, family Scarabaeidae), has been observed killing its prey by violently decapitating and disarticulating it before feeding on the body contents (Larsen et al. 2009, summary here); however, the exact manner by which the beetle avoids or withstands the millipede’s chemical discharges remains unknown.  For ambush predators such as Ectrichodia crux and other ectrichodiines, a strategy similar to that described for another millipede specialist predator, larvae of the phengodid beetle, Phengodes laticollis (order Coleoptera, family Phengodidae), might be employed. This species subdues its millipede prey by piercing thinner regions of the millipede’s integument (e.g., intersegmental membranes on the ventral surface) with its hollow sickle-shaped mandibles and apparently injecting gastric fluids that abruptly paralyze the millipede, thereby preventing it from discharging its gland contents (Eisner et al. 1998).  These undischarged benzoquinones remain confined to the glands and are prevented from diffusing into the body cavity by the glands’ impervious cuticular lining, thus allowing the phengodid larva to safely imbibe the liquified systemic contents of the immobilized millipede.

REFERENCES:

Eisner, T., M. Eisner, A. B. Attygalle, M. Deyrup and J. Meinwald. 1998. Rendering the inedible edible: Circumvention of a millipede’s chemical defense by a predaceous beetle larva (Phengodidae).  Proceedings of the National Academy of Sciences USA 95(3):1108–1113.

Larsen, T. H., A. Lopera, A. Forsyth and F. Génier. 2009. From coprophagy to predation: a dung beetle that kills millipedes. Biology Letters DOI:10.1098/rsbl.2008.0654.

Picker, M., C. Griffiths and A. Weaving. 2002. Field Guide to Insects of South Africa. Struik Publishers, Cape Town, 444 pp.

Reavell, P. E. 2000. The assassinbugs (Hemiptera: Reduviidae) of South Africa. http://oldwww.ru.ac.za/academic/departments/zooento/Martin/reduviidae.html#ectrichodiinae.

Scholtz, C. H. and E. Holm (eds.). 1985. Insects of Southern Africa. Butterworths, Durbin, South Africa, 502 pp.

Copyright © Ted C. MacRae 2009

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