Tuesday Teaser

A recent post by Art Evans at What’s Bugging You reminded me about this photograph that I took some 10 years ago.  This will likely be a difficult challenge, but I’m willing to entertain guesses about its identity and where I found it.  For location, let’s just say I’ve featured quite a few insects from this place in past months – it might take a little digging to figure it out, so first correct answer is worth 4 points.  Knowing this will be key to figuring out its identity.  In that regard, order will be a gimme, so the first person who stumbles upon this will likely earn the measely 2 points available for correctly answering that question.  Family will be more difficult – 4 points if you score first on this one (hint – beware of recent taxonomic changes).  Genus will be a real, though not impossible challenge (in fact, necessary resources to determine this are available online) – a whopping 6 points await the first person to correctly identify that taxon.  Sadly, a definite species name won’t be possible (another clue?), but there is a short list of species that have been described from the general area, so bonus points are available for anyone willing to take on that challenge.  I have no additional pictures of this beast, so look for the answer as a comment to this post in a couple days or so.

Copyright © Ted C. MacRae 2010

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Trilogy of Terror

Last week, Alex at myrmecos tagged me with a fun new meme called These are a few of my favorite stings…. It’s simple – list the things which have stung you (biting doesn’t count), and tag three others for their tales of envenomization. Of course, being the dedicated myrmecologist that he is, Alex leads off with a most impressive list of venomous arthropods, and he selected worthy competition in buzzybeegirl and bugeric.  But me?  I have, for the most part, succeeded in avoiding stings by focusing on a group of insects (beetles) that never evolved such structures.  My domestic list is short and mundane – honey bees, paper wasps, sweat bees, fire ants – and even those not very often.  Alex, however, suspected I might have some tales from exotic lands – thus, I offer the following trilogy and tag Art, Doug, and Kolby.

Tale 1
When I made my first Neotropical collecting excursion some 20 years ago to Ecuador, I was warned by my guide about large, black ants that he called “Congas.” I later learned the species to be what many people call the bullet ant (Paraponera clavata). Now, I’m not an expert on which arthropod truly has the most painful sting, but many people knowledgeable about such matters say it is this species – and I believe them! We were camped out in Sucumbios Province east of Nueva Loja (also called “Lago Agrio”) at an Amazon forest site where recent construction had left rows of month-old slash lining both sides of a 2-km stretch of new road through the forest – can you say woodboring beetles? I roamed up and down that stretch of road, picking a wonderful diversity of longhorned beetles (Cerambycidae) and jewel beetles (Buprestidae) off the slash. At one point, I encountered a whole tree crown laying by the side of the road that required some clambering to get at the beetles crawling on its inner branches. At one point, I braced myself with my arm against a branch and immediately felt an excruciating pain. I looked at my arm and saw one of these large ants clamped onto my arm and quickly slapped it off. I really don’t think words can describe how painful that sting was, and not only did it throb for the rest of the day, but I actually felt sick for the next several days (though I still managed to keep roaming the slash rows). I don’t know if the bullet ant I captured right afterwards was the one that stung me, but I still took great delight in impaling a #2 insect pin through its thorax after I returned home.

Tale 2
Alex mentioned one plant – stinging nettle (Urtica dioica, which also lines one of my favorite mountain bike trails), but I’ve also had a run-in with a much more formidable plant in Mexico. Mala mujer (Cnidoscolus angustidens), which translates as “bad woman” in Spanish, deserves all the respect you can give it. Reported to be one of the most painful stinging nettle-type plants known, it grows commonly from the arid southwest down into the dry, tropical thorn forests of southern Mexico where my colleague Chuck Bellamy and I have made several trips in recent years to search for jewel beetles. One quickly learns to recognize this distinctive euphorbiaceous plant by its green palmate leaves with white veins and thick covering of yellow, stinging trichomes. Unfortunately, in my zeal for beating buprestids from Leucaena diversifolia (netting several of the rare Pelycothorax tylauchenioides and a now paratypical series of what was then an undescribed species of Agrilus), I forgot to maintain my lookout for this common understory plant and got a swipe across the knuckles. Not only did the extreme pain last for hours, but my ring finger began swelling so worrysomely that we stopped in a hospital looking for somebody to cut the ring off. My poor Spanish brought me no sympathy (or service), but fortunately the swelling began subsiding that evening and I didn’t lose my finger. I did, however, live with a rash for the next several days that developed into a hard, purple skin discoloration for the next several weeks.  Bad woman, indeed!

Tale 3
I debated whether to include this experience, but the terror was real so here it is. I wrote about it recently in an article called “Dungers and Chafers – a Trip to South Africa” that appeared in the December 2008 issue of SCARABS Newsletter. Enjoy this excerpt:

After arriving at the park [Borakalalo National Park, North West Province], I could hardly contain myself – I was so anxious to start collecting… We drove through the park for a little bit looking for a good spot to pull over and begin the hunt. After finding such a spot, I grabbed my trusty beating sheet and began doing what I have done so many times before – walking up to a tree, giving a branch a whack with the handle of my net, and hoping to see some prized buprestid laying on the beating sheet. The habitat was ideal for this – dominated by low, spreading acacias such as Acacia tortilis and A. karoo. Buprestids love acacias! I had already learned this in my travels through the American desert southwest and down into Mexico and South America – surely it was the same in South Africa. The first whack yielded nothing – typical. Even when collecting is good, buprestids are never “dripping from the trees,” and often one must literally beat dozens and dozens of trees to really get a good idea of the diversity and abundance of buprestid species that are active in a given area. I whacked a few more trees, with similar results. I then spotted one particularly large acacia tree – something about it said, “beat me!” I walked over to it and gave a branch a whack. All at once, it seemed as though the world was exploding! The air was suddenly abuzz with dozens of large, flying insects, whirring and swirling all around me. My first thought in that initial moment of terror was that I had whacked a hornet’s nest – who knew what kinds of deadly, venomous wasps one might encounter in Africa? Instinctively I ducked and started running, but within a few moments I realized that I was not being chased. Cautiously, I sneaked back towards the tree (after stuffing my heart back down my throat) and realized that they were not hornets after all, but instead beetles. I looked more closely and saw that the tree was literally alive with dozens and dozens of large, green cetoniines resembling our own green June beetle, Cotinis nitida (L.), which seemed to be attracted to the small, white blooms that covered the tree in profusion. I netted a few of the beetles, which I would later determine to represent the common savannah species Dischista cincta (de Geer) (Photo 2). Such was my welcome to Africa, where it seemed the trees literally are ‘dripping’ with beetles!

Disticha cincta (de Geer)

Photo 2. Disticha cincta (de Geer)

Copyright © Ted C. MacRae 2009

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Answers to “Winter botany quiz #2”

Finally, I present to you the answers to “Winter botany quiz #2 “. The delay in providing these answers was two-fold. Firstly, I knew this would be a hard test, so I wanted to give people plenty of time to figure out the answers. Secondly, the answers were delayed an extra day due because of some debate that arose among the experts I consulted about #3 – more on that below. I thank all those who participated, and while there was no clear-cut “winner”, several honorable mentions are deserved:

  • Doug Taron, who was the first to properly deduce the South African nature of these plants.
  • James C. Trager, a myrmecologist (yet still my friend!) who correctly identified the genus of #1.
  • Everyone, for guessing that #2 was “an orchid” – although Tom @ Ohio Nature was the only one to use the formal scientific name for the family, and Doug Taron was the only one to attempt a generic identification (and came close – Oncidium and Ansellia are both assigned to the tribe Cymbidieae in the subfamily Epidendroideae).

#1.  Ornithogalum seineri (family Hyacinthaceae)
Ornithogalum is a large genus occurring mostly in the drier habitats of southern Africa and around the Mediterranean.  The genus and its relatives were formerly included in the Liliaceae (as many of the participants guessed), but the group is now given familial status as the Hyacinthaceae.  This genus contains numerous species of horticultural note.  One is (as James noted) O. umbellatum, or  “star of Bethlehem”, which in North America has escaped cultivation as a garden ornamental and gained status as an invasive weed.  Another is O. longibracteatum (syn. caudatum), a popular houseplant with the common name “pregnant onion”.  This species, native to the Cape and Natal Provinces of South Africa, is easily recognized by its bulb that “gives birth” to tiny replicas of itself just beneath a thin, transparent ‘onion’ skin (as shown in the photo at right from Trans-Pacific Nursery).  At flowering, a long spike grows from the center of the green strap leaves, eventually giving rise to a spearhead of tiny white flowers situated at the end.

While I couldn’t find much information about O. seineri, I did find this spectacular photo of numerous blooming plants in bushveld habitat amongst grazing zebra (photo by ingrid1968 in this post at SANParks.org Forum).  My view of this species was not quite so spectacular, as I saw only the lone plant in the photographs posted earlier.

#2.  Ansellia africana (family Orchidaceae)
Ansellia is an African genus of orchid commonly called Leopard Orchid or African Ansellia.  There is some degree of morphological, geographical and ecological variation in Ansellia populations, with the result that several species, subspecies and varieties have been described.  Flower color varies from pure yellow to variably splotched with brown to almost completely black with finely indicated yellow divisions.  Recent taxonomic work has concluded that there are no discontinuities within the spectra of variations exhibited and the populations are thus attributable to the single, polytopic species, A. africana (Khayota 1999).

Ansellia africana is a large, perennial, epiphytic species that usually grows attached to the branches of tall trees but is sometimes found growing on rocks.  This genus is immediately recognizable by its large, cane-like pseudobulbs that arise from a basal rhizome and is notable for the white, needle-like, upward pointing aerial roots that form a sort of “trash basket” around the clump.  The term is surprisingly appropriate, since the root basket seems to function in catching dropping leaves, flowers and detritus which provide nutrients for the plant as they decay.  This species can grow to enormous size and often forms spectacular clumps, some of which have an estimated weight of more than one ton.

Of the three plants featured in the quiz, this was the one I expected someone would guess, since the species is popularly cultivated by orchid enthusiasts.  Unfortunately, the pressures of wild collection for commercial purposes has caused declines in its population.  The problem is exacerbated by the unsustainable methods use to harvest, transport, and cultivate wild-born plants.  Host trees are usually cut down and sections with the orchid removed, resulting in wholesale destruction of both orchids and hosts. After harvesting, plants are cut up and transported slowly in open handcarts, to be sold along roadsides where they may sit exposed to full sun for days or weeks.  Cutting the clumps damages the roots, and exposure results in dessication, making it difficult for harvested plants to recover once in cultivation.  Plants that do survive harvest and transplant suffer high mortality rates in cultivation due to improper attention to light and moisture regimes.

#3. Adenia sp., poss. glauca (family Passifloraceae)
To be completely honest, not only did I not expect anyone to guess this one, I didn’t think I was even going to be able to provide an answer. I sent the photos to my friend and colleague, George Yatskievych, director of the Flora of Missouri Project (and author of the recently published Steyermark’s Flora of Missouri, 1999 and 2006), who forwarded the photographs to several more colleagues, and at the same time I posted the photos on SANParks.org Forum (a fantastic resource, which I just recently discovered myself, for those interested in South Africa National Parks and their natural history). It took some time for these sources to weigh in with their opinion, which in the end were in agreement that it represented a species of African passion flower in the genus Adenia of the family Passifloraceae (not to be confused with Adenium, a genus of flowering plants in the family Apocynaceae – also occurring in Africa). As for which species, the choices had been narrowed down to either A. glauca or A. fruticosa. According to Imberbe, a photo of the leaves would have been diagnostic, and the flowers are also different (A. glauca has yellow flowers while those of A. fruticosa are green). Fred Dortort, in an article on the University of California at Berkely Botanical Garden website titled, “Passion and Poison“, notes that A. fruticosa has a tall, spindle-shaped caudex topped with a few thin, sparsely-leafed, arching branches, while in A. glauca the caudex is roughly globose and can become quite large. This description seems to favor A. glauca, which Imberbe also noted was known to occur in the area where I took the photographs.

Species identification aside, the genus Adenia is notable for its bizarre adaptations for water storage. Most of the 100 or so species in this Afrotropical and Indomalaysian genus have underground tubers. Those of species adapted to drier environments have grown proportionately larger, with some turning into above ground caudices that can take several different forms and that, in some species, may reach up to eight feet in diameter and height. Even more notable than these succulent adaptations are the poisonous properties that many plants in the genus possess. Not all species have been analyzed (and I found little or conflicting information about A. glauca and A. fruticosa), but one species in the genus – A. digitata – has gained notoriety as perhaps the most poisonous plant in the world. Two different toxins are found within its tuber, one a cyanogenic glycoside, the other a particularly potent toxin called modeccin. The latter is a 57kD protein that resembles ricin and acts a powerful inhibitor of protein synthesis by binding to ribosomes (Gasperi-Campani et al. 1978). Imberbe, in her comments about the photos I posted on SANParks.org Forum, noted the following about plants in this group:

…take heed of the Afrikaans name “Bobbejaangif” (Baboon poison)… It has been used as a fish poison, as well as in suicide and murder. It causes nausea, fits and liver and kidney damage.


Gasperi-Campani, A., L. Barbieri, E. Lorenzoni, L. Montanaro, S. Sperti, E. Bonetti, & F. Stirpe. 1978. Modeccin, the toxin of Adenia digitata. Biochemistry Journal 174:491-496.

Khayota, B. N. 1999. Notes on systematics, ecology and conservation of Ansellia (Orchidaceae), pp. 423-425. In: J. Timberlake & S. Kativu (eds.), African Plants: Biodiversity, Taxonomy and Uses, Royal Botanic Gardens, Kew.

Copyright © Ted C. MacRae 2009

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Winter botany quiz #2

In the first winter botany quiz, I learned that I have some rather astute botanists amongst my readership. They were not only able to quickly identify to species every plant I had pictured but also identify their commonality, sometimes from quite afar. As a result, this one is harder.  I use the term “winter botany quiz” in the broadest possible sense – just because it’s winter here doesn’t mean it’s cold everywhere! All of the photos were taken in the same general (for now unspecified) locality during late November and early December (this paragraph simply reeks of clues!).

To give everyone a fair chance, I’ve turned on comment moderation so people can submit their answers without seeing what has already been submitted.  I’ll remove moderation after a couple days or so.  First one with all the right answers wins the admiration and jealousy of their peers!




#1B - closeup of flowers in #1A




#2B - closeup of flowers of #2A


#3A - the vine, not the trees


#3B - closeup of vine base

Copyright © Ted C. MacRae 2009

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Magodo – giant twig wilter


In previous posts, I have highlighted some of the insects I observed on a trip to South Africa in November-December 1999.  All of the photos I have shown to this point were taken at Borakalalo National Park in North West Province or at the Geelhoutbos farm of Susan Strauss below the Waterberg Range in the formerly Northern, now Limpopo Province.  Both of these locations are deep inside the bushveld, providing ample opportunity to observe an incredible diversity of insect life.  This is not to say that insects, even spectacular ones, cannot be found in more urban areas.  During the weekend between those two mini-expeditions, I stayed with my friend and colleague, Chuck Bellamy, at his home in Pretoria, a beautiful city with lovely architecture, elegant gardens… and some very impressive bugs!  The bug in this photo was found on a tree in a shrubby enclave, and at well over 35 mm in length it is easily the largest leaf-footed bug (order Hemiptera, family Coreidae) that I have ever seen.  Its chunky build, velvety black coloration with thin yellow lines along the sides and down the center of the thorax, and greatly enlarged hind femora quickly led me to a provisional identification of a male Petascelis remipes, or giant twig wilter.  This ID was confirmed by my friend and colleague Harry Brailovsky, an entomologist at UNAM (Universidad Nacional Autónoma de México) and world expert on Coreidae (and who, incidentally, just recently published a review of this Afrotropical genus – Brailovsky 2008).

According to Picker et al. (2002), these insects are found on plants in the genus Combretum.  Like most species in the family, they have scent (“stink”) glands that provide defensive capabilities. Adults are gregarious and bold, walking towards intruders with antennae vibrating when disturbed, and they are apparently capable of squirting their defensive secretions for some distance.  The nymphs are black as well but futher advertise their noxiousness with warning coloration of red spots on a whitish background. Interestingly, and despite their powerful chemical defenses, this species is considered a delicacy in parts of Mozambique where it is known as Magodo.  In a post called Insects for Dinner (in a blog with the eerily similar title, Beating about the Bush), Bart Wursten of Gorongosa National Park in Mozambique describes how local folk burn small patches of the grassland in which these insects are found to smoke them out and catch them.  The Magodo hunters kill the bugs by breaking off the head and removing the scent glands, which releases a very strong almond-like smell.  In doing this, the locals are able to catch considerable quantities of the bugs, which they eat with supper.

Lest you believe such practices are an anomaly, van Huis (2003) has compiled a list of about 250 insect species used as food in sub-Saharan Africa.  Lepidoptera, Orthoptera and Coleoptera represented the bulk (78%) of species eaten, with Isoptera, Hemiptera, Hymenoptera, Diptera and Odonota making up the rest.  Several examples of toxic insects and the traditional methods used to remove the poisons were given.  It was noted that whether or not insects are eaten depends not only on taste and nutritional value, but also on customs and ethnic preferences or prohibitions.  I’m not one to shy away from the thought of eating insects – after all, shrimp are just bugs that live in water, and insects rank far lower in ‘slime factor’ than many other invertebrates (e.g., oysters) that enjoy great popularity in our culture.  I’ve eaten roasted beet armyworm (Spodoptera exigua) pupae and munched on chocolate covered ants, but that’s kid stuff – the armyworms tasted like the soy sauce in which they were roasted, and the ants tasted like, well… chocolate.  I did once eat a softshelled crab (alive!), and I actually hope to one day taste the enormous grub of the giant metallic ceiba borer, Euchroma gigantea, eaten by indigenous cultures in Central and South America.   Still, I think I’d need a lot of faith in my chef’s scent gland removal prowess before I started scarfing Magodo down like popcorn.

What insects have you eaten?


Brailovsky, H.  2008. Notes on the genus Petascelis Signoret and description of one new species (Hemiptera: Heteroptera: Coreidae: Coreinae: Petascelini).  Zootaxa 1749:18–26.

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

van Huis, A. 2003. Insects as food in sub-Saharan Africa. Insect Science and Its Application 23(3):163-185.

Copyright © Ted C. MacRae 2009

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Ship-timber beetle

Atractocerus brevicornis

One of the more unusual, and enigmatic, beetles that I encountered in South Africa was this beetle in the pantropical genus Atractocerus.  Placed in the family Lymexylidae (ship-timber beetles), species in this genus look less like beetles than they do large flying ants or strange damselflies due to their highly reduced elytra that expose their greatly elongated abdomen and leave the hind wings uncovered.  The hind wings also are unusual in that they are held fan-like in repose rather than folded as in most other beetles.  Atractocerus brevicornis is the only species in the genus found in Africa (Scholtz & Holm 1985).

Adults are attracted to light at night, as was this individual that came to our ultraviolet light at Geelhoutbos farm below the Waterberg Range in Limpopo Province. Nothing is known about the biology of Atractocerus, but larvae of other genera are reported to bore into hardwoods and palm stems (Picker et al. 2002). Larvae of the genera Lymexylon and Melittomma are believed to form symbiotic associations with ambrosia fungi that grow on the walls of their galleries (Young, 2002).  Adult females deposit fungal spores in a sticky matrix when they lay their eggs, and the hatching larvae carry the spores into wood on their bodies.  The large eyes of Atractocerus, however, suggest a predatory lifestyle. The common name of the family originates from a northern European species that has in the past been a destructive pest of ship timbers.

Atractocerus species are rarely encountered and therefore, not well studied. Their evolutionary history is still unknown; however, the oldest known lymexylid fossil is a very primitive member of the genus Atractocerus preserved in 100 myo Burmese amber (Grimwold & Engel 2005). Thus, the lineage containing these beetles had already appeared by the mid-Cretaceous and may have originated as early as the Jurassic, a fact that has earned them the moniker “living fossils.” These beetles were once thought to be among the most primitive of all Coleoptera – their simple wing venation, almost undifferentiated antennae and tarsi, and naked abdomen being likened to a supposed neuropteran common ancestor. Most authors now consider the family to be most closely related to the cucujiform groups Cleroidea and Cucujoidea (Wheeler 1986, Young 2002), although some have placed it in the Elateriformia near the Lampyridae based on wing venation. Scholtz and Holm (1985) accepted a cucujiform placement but related the group to the Styopidae (twisted-winged parasites), apparently due to the similarity of their highly reduced forwings.  At the same time, they acknowledged the many morphological and behavioral differences between the two groups, the latter itself being the subject of much evolutionary debate due to disagreement about whether the reduced forwings of male stylopids are truly homologous to the elytra of Coleoptera (some have even suggested homology with the halteres of Diptera that were switched from the metathorax to the mesothorax as a result of homeotic mutation). Most authors now place this latter group in the separate order Strepsiptera.


Grimaldi, D. and M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press, New York, xv + 755 pp.

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

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

Wheeler, Q. D. 1986. Revision of the genera of Lymexylidae (Coleoptera: Cucujiformia). Bulletin of the American Museum of Natural History 183:113-210.

Young, D. K.  2002. 71. Lymexylidae Fleming 1821, pp. 261-262.  In: R. H. Arnett and M. C. Thomas [eds.], American Beetles, Volume 2, Polyphaga: Scarabaeoidea Through Curculionoidea, CRC Press, Boca Raton,880 pp.

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.


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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Bizarre, beautiful extremes

No niche, it seems goes unfilled. Specialization is likely to be pushed to bizarre, beautiful extremes.–E. O. Wilson, The Diversity of Life

Wilson didn’t mention treehoppers specifically when he made the above quote, referring to the exuberance of extreme behavioral and morphological adaptations seen in the biota of the tropics, but he could have just as easily led off with them.  Treehoppers (order Hemiptera, family Membracidae) are well-known for their variety of oddly grotesque shapes resulting from a curiously inflated pronotum – presumably having evolved to resemble thorns and buds on their host plants, or the ants that vigorously defend numerous treehopper species in exchange for their sweet honeydew, or perhaps to aid in the dispersal of volatile sex pheromones (an attractive hypothesis but lacking experimental support). Despite inordinate attention in relation to their low economic importance, it remains that the pronotal modifications of many treehoppers are so bizarre that they continue to defy any logical explanation.

I must admit that, despite my passion for beetles, treehoppers were my first love.  (Well, actually anything that I could bring home from my solo wanderings in the urban woodlands and vacant lots near my childhood home and keep alive in a terrarium was my first true love, but from an academic standpoint, treehoppers were the first group to arouse my taxonomic interest as I began my transformation from child collector to serious student.) I had just begun graduate school in the Enns Entomology Museum under the late hemipterist Tom Yonke to conduct leafhopper host preference and life history studies, and although far more Cornell drawers in the museum contained Cicadellidae, it was the treehopper drawers that I found myself rifling through each afternoon after completing the day’s thesis duties. Despite their lesser number, the treehopper drawers had recently benefited from the attentions of a previous student, Dennis Kopp, whose efforts during his time at the museum concentrated on collecting treehoppers from throughout Missouri and culminated in the four-part publication, The Treehoppers of Missouri (1973-1974). I was enamored by these little beasts – specifically by their exaggerated pronotum – and started collecting them whenever I could on my forays around the state surveying for leafhoppers. They were closely enough related to leafhoppers to make them relevant to my work, only cooler – like leafhoppers on steroids! With The Treehoppers of Missouri as my bible and my desk located a half dozen footsteps from the largest treehopper collection within a several hundred mile radius, I delved into their taxonomy and, for a time, considered a career as a professional membracid taxonomist.

Fast forward nearly 30 years, and my involvement as a taxonomist is neither professional nor deals with membracids. Beetles have taken over as my focal taxon, and I conduct these studies strictly as an avocation. Still, I continue to collect treehoppers as I encounter them, and although such efforts have been largely opportunistic, I’ve managed to assemble a fairly diverse little collection of these insects as a result of my broad travels. Much of this has occurred in the New World tropics, and it is this region that is the center of diversity for the family Membracidae (fossil evidence suggests that subfamily diversification and subsequent New World radiation began during Tertiary isolation about 65 million years ago after South America separated from Africa, since only the primitive subfamily Centrotinae occurs in both the Old and the New Worlds – all other subfamilies are restricted the New World (Wood 1993)).  Every now and then, as I accumulate enough material to fill a Schmidt box, I sit down and study what I’ve collected, comparing it to my meager literature to attempt identifications.  For material I collect in eastern North America, this works fairly well, as there have been a number of publications covering different parts of this area.  Outside of this area, however, my only hope is to entice one of the few existing membracid specialists into agreeing to look at what I’ve accumulated and ask for their help in providing names, in exchange for which they will be granted retention privileges to benefit their research.

idd-treehoppersMost recently, I was able to convince Illinois Natural History Survey entomologist Chris Dietrich to take a look at the material I had accumulated during the past ten years or so, which included many specimens from Mexico and a smattering from other world areas, including South Africa. Chris did his doctoral work at North Carolina State University under “Mr. Membracid” himself, Lewis Deitz, and has since been conducting evolutionary and phylogenetic studies on Membracidae and the related Cicadomorpha. I recently received this material back from Chris (photo above), the majority of which he had been able to identify to species – only a few specimens in the more problematic genera were left with a generic ID.


Campylocentrus sp. (Mexico: Oaxaca)


Hyphinoe obliqua (Mexico: Oaxaca)


Poppea setosa (Mexico: Puebla)


Umbonia reclinata (Mexico: Oaxaca)


Umbonia crassicornis male (Mexico: Puebla)


Umbonia crassicornis female (Mexico: Puebla)

The selection of photos here show a sampling of some of the more interesting forms contained within this batch of newly identified material – all of which hail from southern Mexico. Campylocentrus sp. is an example of the primitive subfamily Centrotinae, distinguished among most membracid subfamilies by the exposed scutellum (not covered by the expanded pronotum).  Hyphinoe obliqua is an example of the largely Neotropical subfamily Darninae, while Poppea setosa represents one of the more bizarre ant-mimicking species of the subfamily Smiliinae.  Umbonia is a diverse genus in the subfamily Membracinae, occurring from the southern U.S. south into South America. Umbonia crassicornis is one of the most commonly encountered species in this genus, with the photos here showing the high degree of sexual dimorphism it exhibits.  As membracids go, these species are quite large (10 mm in length from frons to wing apex for Campylocentrus sp. and P. setosa, a slightly larger 11-13 mm for the others); however, the many smaller species in this family are no less extraordinarily ornamented.  I’ve also included a photo (below) of one of the drawers from the main collection after incorporating the newly identified material – this drawer represents about half of my treehopper collection, with the largely Nearctic tribe Smiliini and the primitive family Aetalionidae contained in another drawer. In all, the material contained one new subfamily, six new tribes, 13 new genera¹, and 30 new species for my collection. For those with an appetite for brutally technical text, a checklist of the species identified, arranged in my best attempt at their current higher classification, is appended below (any treehopper specialist who happens upon this should feel free to set the record straight on any errors). For each species, the country of origin (and state for U.S. specimens) is indicated along with the number of specimens, and higher taxa new to my collection are indicated with an asterisk(*). Don’t worry, I didn’t type this up just to post it here – it’s a cut/paste job from my newly updated collection inventory for Membracoidea. Happy reading!

¹Wildly off topic, and perhaps of interest only to me, but two of the genera represented in the material are homonyms of plant genera: Oxyrhachis is also a Madagascan genus of Poaceae, and Campylocentrus is a Neotropical genus of Orchidaceae. Scientific names of plants and animals are governed by separate ruling bodies (ICBN and ICZN, respectively), neither of which specifically prohibit (but do recommend against) creating inter-code homonyms. The number of such homonyms is surprisingly high – almost 9,000 generic names have been used in both zoology and botany (13% of the total in botany) (source).  Fortunately, there is only one known case of plant/animal homonymy fr BOTH genus- and species-level names – Pieris napi japonica for a subspecies of the gray-veined white butterfly (Pieridae) and Pieris japonica for the popular ornamental plant Japanese andromeda (Ericaceae).


Kopp, D. D. and T. R. Yonke. 1973-1974. The treehoppers of Missouri: Parts 1-4. Journal of the Kansas Entomological Society 46(1):42-64; 46(3):375-421; 46(3):375-421; 47(1):80-130.

Wood, T. K. 1999. Diversity in the New World Membracidae. Annual Review of Entomology 38:409-435.


*Campylocentrus curvidens (Fairmaire) [Mexico] – 4
Campylocentrus sp. [Mexico] – 1

*Umfilianus declivus Distant [South Africa] – 3

*Oxyrhachis latipes (Buckton) [South Africa] – 1

Platycentrus acuticornis Stål [Mexico] – 11
Platycentrus obtusicornis Stål [Mexico] – 3
Platycentrus brevicornis Van Duzee [USA: California] – 7
Tylocentrus reticulatus Van Duzee [Mexico] – 4

*Stalobelus sp. [South Africa] – 1


*Dysyncritus sp. [Argentina] – 1


Aconophora sp. female [Mexico] – 1
*Guayaquila xiphias (Fabricius) [Argentina] – 7

Platycotis vittata (Fabricius) [USA: Arizona, California] – 3
Umbonia crassicornis (Amyot & Serville) [Mexico] – 73
Umbonia reclinata (Germar) [Mexico] – 8

Enchenopa binotata complex [Mexico] – 1
Enchenopa sp. [Argentina] – 6

Subfamily DARNINAE

Stictopelta nova Goding [Mexico] – 9
Stictopelta marmorata Goding [USA: Texas] – 1
Stictopelta pulchella Ball [Mexico] – 11
Stictopelta varians Fowler [Mexico] – 3
Stictopelta sp. [USA: Arizona, California] – 5
Stictopelta sp. [Mexico] – 5
Stictopelta spp. [Argentina] – 6
*Sundarion apicalis (Germar) [Argentina] – 2

*Hyphenoe obliqua (Walker) [Mexico] – 1


Vanduzeea triguttata (Burmeister) [USA: Arizona] – 2

Ceresa nigripectus Remes-Lenicov [Argentina] – 3
Ceresa piramidatis Remes-Lenicov [Argentina] – 4
Ceresa ustulata Fairmaire [Argentina] – 1
Ceresa sp. female [Argentina] – 1
Poppea setosa Fowler [Mexico] – 11
Tortistilus sp. [USA: California] – 1

*Bilimekia styliformis Fowler [Mexico] – 3
Polyglypta costata Burmeister [Mexico] – 18

Cyrtolobus acutus Van Duzee [USA: New Mexico] – 1
Cyrtolobus fuscipennis Van Duzee [USA: North Carolina] – 1
Cyrtolobus pallidifrontis Emmons [USA: North Carolina] – 1
Cyrtolobus vanduzei Goding [USA: California] – 4
Cyrtolobus sp. [USA: Arizona] – 2
*Evashmeadea carinata Stål [USA: Arizona] – 4
*Grandolobus grandis (Van Duzee) [USA: Arizona] – 1
Ophiderma sp. [Mexico] – 1
Palonica portola Ball [USA: California] – 4
Telamona decora Ball [USA: Missouri] – 4
Telamona sp. [USA: Texas] – 1
*Telamonanthe rileyi Goding [USA: Texas] – 2
*Telonaca alta Funkhouser [USA: Texas] – 1
Xantholobus sp. [Mexico] – 1


Microcentrus perditus (Amyot & Serville) [USA: Texas] – 1
Microcentrus proximus (Fowler) [Mexico] – 1


Aetalion nervosopunctatum nervosopunctatum Signoret [Mexico] – 9
Aetalion nervosopunctatum minor Fowler [USA: Arizona] – 2
Aetalion reticulatum (Linnaeus) [Argentina, Uruguay] – 26

Copyright © Ted C. MacRae 2009

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