Best of BitB 2012

Welcome to the 5th Annual “Best of BitB”, where I pick my favorite photographs from the past year. 2012 was one of the most intensive travel years I’ve ever had—I spent 8 weeks in Argentina from February through April, made separate trips to Puerto Rico and Arkansas in May (bracketing a personal week in California), traveled almost weekly to Illinois and Tennessee from June to September (interrupted by a personal week in Florida in July), toured the southeastern U.S. (Arkansas, Louisiana, Mississippi and Georgia—great food!) in early September, chased tiger beetles in Oklahoma, Texas and Arkansas in late September, went back to Argentina for a week in October, and capped off the travel year by attending the Entomological Society of America Annual Meetings in Knoxville, Tennessee (for the first time in more than 10 years!)—whew! While many would cringe at such a travel load, I am among the lucky few who actually get paid for doing something that is also my hobby—entomology! This gives me ample opportunity to further hone my photography skills (nine of the 13 photos I’ve selected below were actually taken while I was on business travel), resulting in two key accomplishments this year—my first ever photography talk at the ESA’s insect photography symposium and my first commercial sales (look for the BitB commercial site to go online in 2013).

Enough blather! Here are my favorite BitB photographs from 2012. Click the link in the text below the photo to see the original post. I would greatly appreciate knowing if you have a favorite (and why)—your feedback will be enormously helpful to me as I continue to learn and develop as a photographer.  For those interested, here are my previous year picks for 2008, 2009, 2010 and 2011. And, as always, thank you for your readership!

Spintherophyta (?) sp. in flower of Abutilon pauciflorum | Buenos Aires, Argentina

From  (posted 2 Feb). One of my 2012 learnings was that sometimes a photograph that is not so close is more effective than one that is as close as possible. In one of my earlier attempts at “not-so-close” macrophotgraphy, the soft colors of the flower compliment the brash shininess of the tiny leaf beetle that has been feeding on its pollen. Pink lines lead the eye directly to the subject and create a pleasing composition, and pollen grains stuck to the beetle—a distraction in some situations—add to the miniature natural history story of the photo.

Apiomerus flavipennis with stink bug prey and kleptoparasitic flies | Chaco Province, Argentina

From  (posted 11 Mar). I selected this photo solely for the complex natural history story drama it shows—stink bug (Piezodorus guildenii) feeding on soybean becomes prey of an assassin bug (Apiomerus flavipennis), with volatiles from the chemicals it emitted in a vain attempt to defend itself serving as cues to kleptoparasitic flies (families Milichiidae and Chloropidae) that benefit from the assassin bug’s labors.

Planthopper nymph | Buenos Aires Province, Argentina

From  (posted 26 Mar). Another learning that I began putting into practice in 2012 was the use of low perspective for compositional impact. The cryptic coloration of this planthopper nymph (family Fulgoridae) made it almost invisible on the branch on which it was sitting when viewed from a normal “top-down” human perspective. Getting “down under” it, however, brought the nymph to life and emphasized its unusual form.

Megabaris quadriguttatus | Corrientes Province, Argentina

From  (posted 12 Apr). I spent much of 2012 working on the “blue sky background” technique, with these weevils from northern Argentina representing one of my better attempts. Macrophotography of insects with a blue sky background involves setting exposure, ISO, and aperture to achieve two separate exposures—full flash illumination of the subject for maximum depth-of-field, and ambient light from the sky to create a clean, uncluttered, natural-looking background. In this shot I managed to achieve an almost ideal shade of blue to compliment the wild black, white and red colors of the beetles. (My one criticism of the photo is having clipped one of the beetle’s feet.)

Bombylius sp. cf. mexicanus | Scott Co., Missouri

From  (posted 16 May). This photo is unusual if nothing else. Focus, lighting, depth-of-field, and composition are all better than can be hoped for in a single shot, but the subject—perfectly alive—is in a most unusual position. Read the original post to find out how this happened.

Perisphaerus sp. (a pill roach) | Vietnam (captive individual)

From  (posted 27 May). White-box photography is an excellent technique for clean, uncluttered photographs of insects, but it also isolates them from their natural surroundings and limits their natural history appeal. The best white-box photos are those that highlight a key feature or behavior of the subject—in this case a pill roach’s comically conglobulating defensive posture.

Micronaspis floridana (Florida intertidal firefly) larva | Pinellas Co., Florida

From  (posted 31 July). Here is another photo whose back story played a big part in its selection. This firefly larva not only represents a rare Florida-endemic species but was also first seen by my then 12-year old nephew, who willingly accompanied me through a dark, spooky salt marsh in the middle of a humid Florida night to see what he could learn. The lesson here for budding natural historians (and old-timers like me) cannot be overstated!

Arctosa littoralis (beach wolf spider) | Lewis Co., Missouri

From  (posted 23 Aug—prelude to  posted 28 Aug). Those who follow this blog know of my obsession with close-up portraits, and while tiger beetles are the subjects I most commonly photograph in this manner, I am always on the lookout for good subjects in other taxa. This wolf spider “face” almost looks human, with “two” eyes, two “nostrils” and a shiny upper lip above huge (albeit hairy) buck teeth! It’s enough fill-the-frame spidery goodness to melt (or explode) the heart of even the most ardent arachnophobe!

Anticarsia gemmatalis (velvetbean caterpillar) egg on soybean leaf

From Life at 8X—Guide to lepidopteran eggs on soybean (posted 3 Sep). “Life at 8X” was a new series I introduced this year, featuring insects photographed at magnifications testing the upper limit of my equipment and photographic skills. Diffraction is the chief difficulty with magnifications as high as this and is the primary flaw in the above photograph. Nevertheless, such view of a moth egg on the underside of a soybean leaf provides a spectacular view of the otherwise unseen micro-world that lives right beneath our noses.

Megacyllene decora (amorpha borer) on snakeroot flowers | Mississippi Co., Missouri

From  (posted 12 Sep). This second example of “blue sky background” was taken later in the year and was considerably more difficult to capture than the first because of the larger size of the subject and resulting need for a longer focal length macro lens. Getting a well-lit, focused, and composed image with a desirable shade of blue in the background depended not only on finding the proper camera settings, but also secure body and camera bracing techniques for this completely hand-held shot.

Cicindelidia politula politula (Limestone Tiger Beetle) | Montague Co., Texas

From  (posted 28 Sep). I will go ahead and say it—this is my favorite photograph of 2012. As discussed under the first entry, panning back from the subject can allow for some very interesting compositions. This photo combines charismatic pose by a wary subject with panning back and low perspective to create an image that scores high in both natural history and aesthetic appeal.

Calosoma sayi (black caterpillar hunter) | New Madrid Co., Missouri

From Black is beautiful! (posted 7 Nov). Of course, close-as-possible can also be used to create striking photos, especially if the subject exhibits features that are best seen up close. Anything with jaws fits the bill in my book, and highlighting the mandibular sculpturing of this caterpillar hunter (a type of ground beetle) required precise angling of the flash heads for maximum effect.

Cicindela repanda (Bronze Tiger Beetle) | St. Louis Co., Missouri

From  (12 Nov). This final selection is not a rare species, but it is as close as I have come to what I consider the “perfect” tiger beetle macrophotograph—a close, low angle, lateral profile of an adult in full-stilt posture (a thermoregulatory behavior), well lit, perfectly focused, and with a dynamic but pleasingly blurred background. It’s a perfect storm of a photo that took the better part of two hours to achieve—rarely do all of these elements come together in a hand-held photograph of an unconfined tiger beetle in its native habitat.

Well, there you have it. I hope you’ve enjoyed my selections, and again please do let me know if you have a personal favorite. See you in 2013!

Copyright © Ted C. MacRae 2012

More Eocene insects

Most of the Green River Formation (GRF) insect fossils that I have on loan clearly represent either beetles (order Coleoptera) or flies (order Diptera). I’ve already shown a few of the latter (fungus gnat, midge), as well as some that don’t belong to either order (ant, cricket?). Here are a few more that seem identifiable to order, but family-level identification is less certain. Thoughts from the readership would be most welcome.

This fossil shows an aggregation of insects that I believe represent some kind of beetle. Based on shape and size (16.7 mm length) I’m guessing perhaps either a diving beetle (family Dytiscidae) or whirligig beetle (family Gyrinidae). These are both aquatic families, although only the former is among the beetle families recorded from the GRF by Wilson (1978).

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There are two insect fossils on this specimen, but the closeup is the one near the center of the rock. It is tiny (3.5 mm in length), and at first I thought it might be a fly (order Diptera). However, dipterist Chris Borkent thinks it might be a small hymenopteran (bee?) because it has what looks to be long multi-segmented antennae. The only bee family recorded for the GRF by Wilson (1978) is Anthophoridae (now included within Apidae), of which this fossil clearly is not a representative. There are six other hymenopteran families recorded in that work, of which Tenthredinidae is the only one that seems plausible. Of course, it could represent a family not recorded by Wilson (1978). Collected along Hwy 139 in Douglas Pass (Garfield Co., Colorado).

Here is a closeup of the other fossil (far right in photo above). This looks to me like a brachyceran fly, and I’ve sent a high resolution version of the image to Chris Borkent to see what he thinks.

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The label accompanying this fossil indicates “Mosquito (?),” but to my eye this looks like a true bug (order Hemiptera). It is small—only 5.9 mm in length—and has the gestalt of a plant but (family Miridae) or seed bug (family Lygaeidae). GRF fossils representing the latter but not the former were recorded by Wilson (1978). Also collected along Hwy 139 in Douglas Pass (Garfield Co., Colorado).


Wilson, M. V. H. 1978. Paleogene insect faunas of western North America. Quaestiones Entomologicae 14(1):13–34.

Copyright © Ted C. MacRae 2012

Virtual Mantle 2012

In recent years a truly delightful custom has developed among taxonomic entomologists—Photoshopped e-Christmas cards featuring their favorite insects! This should not be a surprising development—connectivity among scientists from around the world is at an all-time high, as is their access to digital cameras and imaging software. And, of course, what better subject for such cards could one ask for than insects (especially beetles, I might add)? It’s hard to resist anthropomorphizing them, with their buggy eyes and comical expressions, but their outright beauty makes them perfect also as pure ornaments. I did my first one last year (Santa Jaws), and for this year’s version I created Buprestis saintnicholasii. Following are the e-Christmas cards on my virtual mantle this year, received from beetle taxonomists as far and wide as Arizona, California, Oregon, Bulgaria, Germany and Romania!

I really enjoy getting these—if I’m not already on your e-Christmas card mailing list please consider adding me in 2013!


Chuck & Rose Bellamy – Sacramento, California


Mark Kalashian – Yerevan, Armenia


Hans Mühle – Munich, Germany


Adrian Ruicanescu – Cluj-Napoca, Romania


Bill Warner – Phoenix, Arizona


Bill Warner – Phoenix, Arizona


Rick & Kathy Westcott – Salem, Oregon


Victor Gashtarov – Sofia, Bulgaria

Copyright © Ted C. MacRae 2012

The 2nd-oldest Known Myrmicine Ant

Among the 20 or so insects represented in the Green River Formation (GRF) fossils that I currently have on loan, this rather obvious ant (family Formicidae) is the only one that is firmly assignable to the order Hymenoptera (wasps, bees and ants). This is not surprising, as hymenopterans are not well represented among GRF insect fossils. In fact, of the 300+ insect species that have been described from GRF deposits (Wilson 1978), more than two-thirds belong to just three orders—Diptera (flies), Coleoptera (beetles) and Hemiptera (true bugs). Hymenoptera, on the other hand, comprise only 4% of GRF fossils (Dlussky & Rasnitsyn 2002). I presume these numbers are more a result of taphonomic (fossil formation) bias than a true reflection of insect diversity in western North America during the Middle Eocene (47–52 mya).

cf. Myrmecites rotundiceps | fossil impression from the Green River Formation (45 mya, middle Eocene)

cf. Myrmecites rotundiceps (length = 6.7 mm).

Ants in particular have been poorly represented by GRF deposits. Only four named species were known until Dlussky & Rasnitsyn (2002) reviewed available GRF fossils and increased the number to 18 (15 described as new, one older name placed in synonymy). Diagnoses, line drawings, and keys to all covered subfamilies, genera and species provide one of the best treatments to GRF insect fossils that I’ve come across. According to that work, the fossil in these photos seems comparable to the description and illustration given for Myrmecites rotundiceps, a unique fossil with the general appearance of ants in the subfamily Myrmicinae but differing from all known Eocene and New World fossil ants by its very short, two-segmented waist. The only difference I noted was size—6.7 mm length for my fossil versus 5.5 mm for the holotype (see figure below). Of course, I’m more comfortable identifying extant Coleoptera than extinct Formicidae, so I contacted senior author Gennady M. Dlussky to see if he agreed with my opinion. He graciously sent the following reply:

I agree that specimen on your photo is very similar to Myrmecites rotundiceps. It is larger (holotype is 5.5 mm long), but it may be normal variability. I cannot see another differences.

Myrmecites rotundiceps, holotype (Gennady & Rasnitsyn 2002)

Myrmecites rotundiceps Gennady & Rasnitsyn 2002, holotype (reproduced from Gennady & Rasnitysyn 2002)

If correctly assigned, M. rotundiceps is the second oldest known member of the subfamily Myrmicinae—the oldest being Eocenidris crassa from Middle Eocene Arkansas amber (45 mya). In fact, the only older ant fossil of any kind in North America is Formicium barryi (Carpenter) from Early Eocene deposits in Tennessee (wing only). [Edit: this is actually the only older Paleocene ant fossil—there are some Cretaceous-aged fossils such as Sphecomyrma freyi (thanks James Trager).] Since myrmicine fossils of comparable age are lacking from other parts of the world, this might suggest a North American origin for the subfamily; however, it could also be an artifact of incomplete knowledge of ants from older deposits in other parts of the world. Myrmicine ants make their first Eurasian appearance in Late Eocene Baltic amber deposits (40 mya) and become more numerous in North America during the early Oligocene (Florissant shales of Colorado, 33 mya). (Dlussky & Rasnitsyn (2002) consider the Middle–Late Eocene ant fauna to represent the beginnings of the modern ant fauna, with extant genera becoming numerous and extinct genera waning but still differing by the preponderance of species in the subfamily Dolichoderinae over Formicinae and Myrmicinae.


USA: Colorado, Rio Blanco Co., Parachute Creek Member.

The photo above shows the entire fossil-bearing rock (also bearing the putative orthopteran posted earlier).

My thanks to Gennady Dlussky and James Trager for offering their opinions on the possible identity of this fossil.


Dlussky, G. M. & A. P. Rasnitsyn. 2002. Ants (Hymenoptera: Formicidae) of Formation Green River and some other Middle Eocene deposits of North America. Russian Entomological Journal 11(4):411–436.

Wilson, M. V. H. 1978. Paleogene insect faunas of western North America. Quaestiones Entomologicae 14(1):13–34.

Copyright © Ted C. MacRae 2012

T.G.I.Flyday: Tachinomyia sp.

You would think that somebody with enough patience to photograph tiger beetles would be equally patient with “calyptrate” flies, but for me such is not the case. It’s not that I don’t find them interesting (although, really, what insect group can match the diversity, polytopism, ecological extremism and behavioral charisma of tiger beetles?), but their flighty, frenetic behavior and difficult taxonomy are just a bit too much for me. After all, why invest the time it would take to get a good photograph of something that, in the end will probably be unidentifiable.


Tachinid fly (prob. Tachinomyia sp.) | Wayne Co., Missouri

Last April as I was hiking a woodland trail in Sam A. Baker State Park in southeastern Missouri’s Ozark Highlands, I saw this decent-sized fly bumbling across the trail. I knelt to look at it more closely, and though it tried to flee, it seemed too weak and uncoordinated to take flight. It was in beautiful condition—a perfect specimen, and I surmised that it represented a newly emerged adult that had not yet hardened sufficiently to withstand the rigors of flight. I was fascinated by its distinctive, orange tarsal pads and the white “beard” around its head, and the ability to coax the fly onto a leaf and hold the leave in whatever position I desired was all the enticement I needed to spend a little time with it. Out of the several shots that I took, these two are my favorites.


You got it—the BitB face shot!

Of course, just being able to photograph the fly was only half the battle—there still remained the matter of its identification. It seemed “tachinidish” to me at the time, but a little digging revealed that there are species of Sarcophagidae look very similar to tachinids, the difference being the presence or absence of a visible postscutellum. My photos don’t show this character, and I quickly became overwhelmed when I tried scanning through BugGuide photos for these two families. Nevertheless, I’m a persistent sort, and after winnowing out the numerous unlikely choices I finally settled on not only family Tachinidae, but possibly Tachinomyia sp. (tribe Exoristini). I was tempted to go out on a limb and post the ID here unvetted, but I chickened out and and sent the photos to fly guy Norman Woodley at the Systematic Entomology Lab in Washington, DC. Norm supported my identification and wrote back:

I think it is probably Tachinomyia.  It would be better to have a wing view as well as the hind end to be absolutely positive, but I’m reasonably sure that’s what it is.  It’s a male.  Some species are active in the spring, so that fits with your data as well.

Copyright © Ted C. MacRae 2012

A 50-million-year-old midge

Several of the insect fossils collected from the Green River Formation (45–50 mya) that I am photographing appear to be flies, and specifically members of the “primitive” suborder Nematocera. This is not surprising, as the G.R. Formation of Colorado, Utah, and Wyoming, is composed of shales derived from volcanic ash sediments that were laid down in a system of large, shallow lakes. Most (all?) nematoceran flies are aquatic to some degree in the larval stage, thus the adults are also closely associated with such habitats for mating and egg laying.

Diptera: Chironomidae | USA: Colorado, Garfield, Hwy 139, Douglas Pass

Diptera: Chironomidae | USA: Colorado, Garfield, Hwy 139, Douglas Pass

This particular fossil looked to me a lot like the more elegantly preserved fossil of another fly that I posted a few days ago, which at the time I thought represented a member of the family Mycetophilidae (fungus gnats) or Sciaridae (black-winged fungus gnats). Several knowledgable specialists offered their opinions in comments at this site and at Facebook’s Diptera forum (my thanks to all who offered their opinion), with most settling on Mycetophilidae and Vlad Blagoderov further suggesting subfamily Mycetophilinae. The fossil posted here seemed to me to represent a dorsal view of the same species, but, of course, I’m a coleopterist—so what do I know? Indeed, dipterist Dr. Chris Borkent believes this is actually a species of Chironomidae (common name simply “midges”)—also a nematoceran but differing from Mycetophilidae by their longer front tarsi and longer, relatively narrower wings. Males of the family have thickly plumose (“feathery”) antennae, which are not visible in this specimen and thus suggesting it might be a female. I wouldn’t doubt Chris’ identification  for a second, as he comes from good stock—his father is Art Borkent, a world expert on several families of nematoceran dipterans. Art also agreed after seeing the photo that it looked like a female chironomid midge, so that is what I am going with. Thank you, Chris and Art, for your help in identifying this fossil!

Complete fossil specimen (63 mm x 52 mm maximum each axis).

Copyright © Ted C. MacRae 2012

Eye to eye to eye to eye with a tiger beetle larva

After the past few years of hunting tiger beetles, I’ve learned not only how to find the larval burrows but—at least for most of the species occurring in Missouri—how to identify the larvae in the field. While conclusive identifications rely upon morphological characters, a preliminary field ID is often possible based on a combination of burrow size, placement, soil type and knowledge of which species are likely to occur in a given habitat. Tiger beetle larvae don’t have the same aesthetic appeal to many people that the adults have, and for this reason many species remain undescribed in the larval stage—even the well-studied North American fauna has only about 60% of its species with the larval stages described (Pearson et al. 2006). Nevertheless, the ability to find, collect and rear tiger beetle larva remains an important part of my studies because it not only expands my survey power (most tiger beetles have more restricted temporal occurrence as adults than as larvae) but can also lead to novel findings such as previously undescribed larvae and unknown parasitoid associations.

Tetracha virginica 3rd instar larva | Mississippi Co., Missouri

Tetracha virginica 3rd instar larva | Mississippi Co., Missouri

This larva was dug from its burrow in bottomland forest habitat in the southeastern lowlands of Missouri. However, before I even saw the larva I knew it belonged to the genus Tetracha and probably represented the species T. virginica (Virginia Metallic Tiger Beetle, according to Erwin & Pearson 2008). How did I know this? First, the size of the burrow (~8 mm in diameter) excluded all but one other non-Tetracha species known to occur in Missouri—Cicindelidia obsoleta vulturina (Prairie Tiger Beetle), a species known to occur only in the dry, rocky, dolomite glades in the White River Hills region of extreme southwestern Missouri. Secondly, while T. carolina (Carolina Metallic Tiger Beetle) is also found in southeastern Missouri, that species has been associated almost exclusively with treeless habitats—at least in southeastern Missouri (K. Fothergill, personal communication). Since the burrow from which this individual was dug was found in wet, bottomland forest, chances were high that it instead represented T. virginica.

Simple, thorn-like outer hooks with much smaller inner hooks distinguish Tetracha larvae from other tiger beetle genera.

Simple, thorn-like outer hooks with much smaller inner hooks distinguish larvae of Tetracha.

Notwithstanding the circumstantial evidence, there are morphological characters that also distinguish both the genus and the species of this larva. Of primary importance are the hooks and setae on the prominent “hump” of the fifth abdominal segment. This hump is braced against the vertical wall of the larval burrow as it sits at the entrance waiting for passing prey. Once the prey is seized, the hump armature provides traction against the burrow wall, preventing the struggling prey from pulling the tiger beetle larva out of its burrow (where it would not only be ineffectual as a predator but also highly vulnerable to predation itself). Tiger beetle larvae can often be distinguished at the generic level by the shape and size of the main hooks. Tetracha larvae have four hooks—two outer and two inner—that are simple and thorn-like, with the inner hooks much smaller than and placed much closer to the outer hooks than to each other (other genera either have six hooks, or they have the outer pair 1) highly curved or 2) the inner pair larger and nearly as close to each other as to the outer hooks). There are also fine details of the pattern of the setae (smaller hairs) on the hump that identify this larva as T. virginica, but the presence of numerous hairs over the surface of the abdominal segments is a much easier character to see in the field (see first photo).

Note the white-margined pronotum and nearly equal sized simple eyes.

Note also the white-margined pronotum and nearly equal sized simple eyes.

Finally, there is that head—two pairs of large, simple eyes sitting behind gaping, cocked jaws that give them an oh so alien aspect! An often metallic, shield-like pronotum sitting behind the head, both used in concert to seal the burrow entrance as the larva lies in wait, serve to complete the alien ensemble but also offer clues to the larva’s identity. All larvae of Tetracha and closely related genera bear a distinctive rim of white around the pronotal margin, making them instantly recognizable even while still sitting in their burrow. Also useful is the relative size of the eyes, which in the case of Tetracha the second pair of eyes are nearly as large as the first pair (Amblycheila and Omus have the second pair distinctly smaller than the first, while Cicindela and related genera also have the eyes more nearly equal-sized).

P.S. This is what I was photographing when my friend Kent Fothergill surreptitiously took this photograph of me!


Erwin, T. L. and D. L. Pearson. 2008. A Treatise on the Western Hemisphere Caraboidea (Coleoptera). Their classification, distributions, and ways of life. Volume II (Carabidae-Nebriiformes 2-Cicindelitae). Pensoft Series Faunistica 84. Pensoft Publishers, Sofia, 400 pp.

Pearson, D. L., C. B. Knisley and C. J. Kazilek. 2006. A Field Guide to the Tiger Beetles of the United States and Canada. Oxford University Press, New York, 227 pp.

Copyright © Ted C. MacRae 2012