An interesting flightless May beetle

I suppose you are an entomologist?

Not quite so ambitious as that, sir. I should like to put my eyes on the individual entitled to that name. No man can be truly called an entomologist, sir; the subject is too vast for any single human intelligence to grasp.

Phyllophaga cribrosa | Gloss Mountain State Park, Major Co., Oklahoma

Phyllophaga cribrosa | Alabaster Cavern State Park, Woodward Co., Oklahoma

I suppose the above quote from The Poet at the Breakfast Table, by Oliver Wendell Holmes, Sr., applies just as well to those who would call themselves coleopterists, for restricting oneself to the study of beetles diminishes by scarcely more than half the known diversity of all insects. I still dare to call myself a coleopterist, but I’m the first to admit that while there are a few groups of beetles that I know very well, there are many more that I know only superficially and some that befuddle me completely. An example of the latter is May beetles (family Scarabaeidae, genus Phyllophaga). With more than 400 species in North America (Ratcliffe & Jameson 2010), it is one of the most speciose genera in our fauna. Not surprisingly, species identifications can be very difficult, oftentimes relying on examination of male and female genitalia.


The flightless adults are most often found on the ground or low vegetation.

Not all species of Phyllophaga, however, are difficult to identify. The species shown here—P. cribrosa—is rather easily recognized within the genus by its oval, convex shape, shining black coloration, cribrose (perforated like a sieve) surface, 10-segmented antennae, and flightless nature. The resemblance to certain darkling beetles (family Tenebrionidae) is striking, although I suspect this may be a result of convergent adaptation to dry habitats rather than mimicry (Kaufman & Eaton 2007). While most species of Phyllophaga are nocturnal and capable of flight, adults of P. cribrosa are flightless and can be found crawling on the ground and clinging to low vegetation during the day. I found these beetles this past June at several locations in northwestern Oklahoma.


The distinct elytral furrows distinguish P. cribrosa from closely related species.

There are at least two other closely related species in the genus (i.e., P. epigaea and P. zavalana) that resemble P. cribrosa; however, both of these species are restricted to Texas, and they lack the distinct longitudinal elytral furrows exhibited by P. cribrosa. The species is said to be an occasionally serious pest of crops (Luginbill & Painter 1953), although I suspect that in most cases this results from new plantings of crops in former grasslands because of the limited dispersal abilities of the beetles.


Eaton, E. R. & K. Kaufman. 2007. Kaufman Field Guide to Insects of North America. Houghton Mifflin Harcourt, 391 pp.

Luginbill, P., Sr. & H. R. Painter. 1953. May beetles of the United States and Canada. U.S. Department of Agriculture Technical Bulletin 1060, 102 pp,

Ratcliffe, B. C. & M. L. Jameson (eds.). 2010. Generic Guide to New World Scarab Beetles (available at: /Guide/index4.htm).

Copyright © Ted C. MacRae 2013

Oklahoma Collecting Trip iReport

I’m back home after my week-long collecting trip to western Oklahoma, and at the risk of sounding hyperbolous I can only describe it as one of the most successful collecting trips I’ve ever had. Seriously! These kinds of trips don’t happen all that often for a variety of reasons—timing is off, rains didn’t happen, weather was uncooperative, etc. etc. Once in a while, though, everything comes together, and this was one of those times. The trip was also a return to my roots so to speak—I’ve been rather distracted in recent years with tiger beetles, but jewel beetles (family Buprestidae) and, to a lesser extent longhorned beetles (family Cerambycidae), are really the primary focus of my taxonomic studies. It had been several years since I’d had a good “jewel beetle trip,” so that was the focus of this trip. In planning the trip, I recalled seeing jewel beetle workings in several woody plant species in the same area during last September’s trip, and the occurrence of May rains seemed to bode well for my early June timing.

Gloss Mountains State Park, Major Co., Oklahoma

Gloss Mountains State Park, Major Co., Oklahoma

My instincts proved to be justified—in seven days in the field I collected an estimated 1000–1500 specimens representing at least two dozen species of Buprestidae and a dozen or more Cerambycidae. More important than the numbers, I collected a number of species in good series that I have either not or only rarely collected before, and in fact the second beetle that I collected turned out to be a new state record! Of course, I also brought along my full-sized camera and associated gear and photographed many of the species that I collected. I will feature these photos in future posts, but for this post I thought it might be fun to give a high level view of the trip illustrated only with photos taken with my iPhone (which I also carry religiously in the field with me). The iPhone is great for quick snaps of scenery and miscellaneous plants and animals for which I don’t feel like breaking out the big camera, or as a prelude to the big camera for something I’d like to share right away on Facebook. Moreover, there are some types of photos (landscapes and wide-angles) that iPhones actually do quite well (as long as there is sufficient light!).

Gloss Mountains State Park, Major Co., Oklahoma

Atop the main mesa at Gloss Mountains State Park, Major Co., Oklahoma

My first destination was Gloss Mountains State Park (Major Co.), a stunning system of gypsum-capped, red-clay mesas. I’ve already found a number of rare tiger beetles here such as Cylindera celeripes (Swift Tiger Beetle), Amblycheila cylindriformis (Great Plains Giant Tiger Beetle) and Dromochorus pruinina (Frosted Dromo Tiger Beetle), and in the past two falls I’ve found two interesting jewel beetle records: Chrysobothris octocola as a new state record, and Acmaeodera macra as a northern range extension. On this trip, I started out beating the mesquite  (Prosopis glandulosa) and immediately got the longhorned beetle Plionoma suturalis—a new state record! They were super abundant on the mesquite, and I collected several dozen specimens along with numerous C. octocola as well. I then moved over to the red-cedar (Juniperus virginiana), which was showing a high incidence of branch dieback, and collected nice series of several buprestids, including what I believe to be Chrysobothis ignicollis and C. texanus. Up on top of the mesa there are small stands of hackberry (Celtis laevigata) and soapberry (Sapindus saponaria), both of which are very good hosts for Buprestidae. Not much was on the soapberry, but I beat large series of several Buprestidae from the hackberry, including what I believe to be Chrysobothris caddo and—the real prize—Paratyndaris prosopis! My old friend C. celeripes was also out in abundance, so I collected a series to add to my previous vouchers from this site. Back down below, I marveled at a juvenile western diamondback rattlesnake (Crotalus atrox) in the area where I found some more A. cylindriformis larval burrows. Daylight ran out before I could dig them up, and after 11 hours in the field I was exhausted, so I returned the next morning and got one 1st- and two 3rd-instar larvae and went back up on top of the mesa and beat several more P. prosopis from the hackberry.

Bullsnake (Pituophis catenifer sayi) | Alabaster Cavern State Park, Woodward Co., Oklahoma

Bullsnake (Pituophis catenifer sayi) | Alabaster Cavern State Park, Woodward Co., Oklahoma

My second stop was at Alabaster Cavern State Park (Woodward Co.), where C. celeripes was again abundant on the gypsum-clay exposures surrounding an impressive gorge thought to be a collapsed cave complex. I focused on beating hackberry because of the success with buprestids on this plant at Gloss Mountains SP, and although they were not quite as abundant here as at Gloss Mountains I still managed to end up with good series of C. caddo and several species of Agrilus. Because I had spent the morning at Gloss Mountains, I had only a partial day to explore Alabaster Caverns and, still getting used to the weight of the camera bag on my back, decided to leave the big camera in the car. This was a mistake, as I encountered my first ever bullsnake (Pituophis catenifer sayi) and had to settle for iPhone photos of this species—the photo above being the best of the bunch. An approaching storm put an end to my second day after another 10 hours in the field, and I drove an hour to Woodward.

Moneilema sp. on Opuntia phaecantha | Alabaster Cavern State Park, Woodward Co., Oklahoma

Moneilema sp. on Opuntia phaecantha | Alabaster Cavern State Park, Woodward Co., Oklahoma

My third day started out at nearby Boiling Springs State Park, a riparian oasis on sandy alluvium alongside the nearby Cimarron River. The woodlands are dominated by hackberry and American elm, and although a few buprestids were beaten from hackberry and honey locust (Gleditisia triacanthos), the numbers and diversity were not enough to hold my interest in the spot. After lunch, I decided to return to Alabaster Caverns SP and explore some other areas I had not had a chance to explore during the previous partial day. It’s a good thing that I did, as I ended up finding a nice population of longhorned cactus beetles in the genus Moneilema associated with prickly pear cactus (Opuntia phaecantha). I collected a nice series of adults and also learned a few lessons in how to photograph these beetles on their viciously protective host plants. The photo above gives a taste of what will come in the photos that I took with the big camera. After eight hours in the field and darkness falling, I drove two hours to Forgan in Beaver Co.

Beaver Dunes State Park, Beaver Co., Oklahoma

Beaver Dunes State Park, Beaver Co., Oklahoma

Day 4 in the field started out cold and ominous, having stormed heavily during the previous night and with thick clouds still hanging in the sky. I feared the day might be a wash but decided to venture to Beaver Dunes State Park anyway and take my chances (beating can still be productive even in cold weather as long as the foliage is not wet). It’s a good thing that I did, as the buprestids were as numerous as I’ve ever seen them. The park’s central feature is a system of barren sand dunes that are frequented by ORV enthusiasts and surrounded by hackberry woodlands. The park also has a reservoir and campground, around which are growing a number of cottonwoods (Populus deltoides).

Hackberry Bend Campground, Beaver Dunes State Park, Beaver Co., Oklahoma

Hackberry Bend Campground, Beaver Dunes State Park, Beaver Co., Oklahoma

These hackberrys and cottonwoods proved to be extraordinarily productive. On the former I collected large series of several species of Chrysobothris and Agrilus, and while I collected fewer Buprestidae on the latter, these included Agrilus quadriguttatus and Poecilonota cyanipes! The latter species I had never collected until last year (from Cerceris fumipennis wasps), and beating the lower branches of the declining cottonwoods produced a series of about a dozen specimens. I also got one specimen on black willow (Salix nigra), along with a few Chrysobothris sp. and what I take to be Agrilus politus. Also in a low branch of one of the cottonwoods was a bird’s nest with a single egg that, according to Facebook comments, either represents the American Robin or a Gray Catbird. (I returned the next day and saw two eggs in the same nest.)

American Robin or Gray Catbird nest w/ egg | Beaver Dunes State Park, Beaver Co., Oklahoma

American Robin or Gray Catbird nest w/ egg | Beaver Dunes State Park, Beaver Co., Oklahoma

As the day drew to a close, I found two interesting longhorned beetle species at the edge of the dunes: one large, powdery gray Tetraopes sp. on milkweed (Asclepias sp.), and huge numbers of Batyle ignicollis evidently perched on the yellow spiked inflorescence of an as yet undetermined plant. I have seen this species on many occasions, but always in low numbers, yet here were literally hundreds of individuals on the plants, all having assumed a characteristic pose on the inflorescence suggesting that they had bedded down for the night. I only spent eight hours in the field on this day because of the late start, and as darkness approached I began the two-hour drive to Boise City.

Black Mesa landscape

Sculpted sandstone landscape in the vicinity of Black Mesa State Park, Cimarron Co., Oklahoma

The final two days in the field were supposed to be spent exploring the area around Black Mesa in the extreme northwest corner of Oklahoma, and another hour of driving was needed to get to the area from Boise City. I first went to Black Mesa State Park, and while the landscape was stunning (see above) the area was extremely dry. I feared the collecting would not be at all productive in this area but wanted to give the area a good effort before making a call. As I approached the entrance to the park, I saw a jeep parked by the side of the road with a license plate that read “Schinia,” which I recognized as a genus of noctuid moths that are very popular with collectors. I pulled over and talked to the driver, who was indeed a lepidopterist from Denver and had just arrived himself. We talked and exchanged contact information, and learning of my interest in beetles he directed me to a small stand of Gambel oak (Quercus gambelii) and one-seed juniper (Juniperus monosperma) on a sculpted sandstone escarpment not far from the park. I found the spot, and although I beat three Chrysobothris sp. from the first juniper tree that I whacked, another hour of beating produced only one more beetle from the juniper and nothing from the oak. I returned to the spot where we had met and encountered him again on his way out! We stopped and chatted again and found a few specimens of what I take to be Typocerus confluens on the yellow asters, but by then I was having my doubts about staying in the area. I told him I was going to check out a ravine in the park and then decide.

Petrified forest | Black Mesa State Park, Cimarron Co., Oklahoma

Petrified forest | Black Mesa State Park, Cimarron Co., Oklahoma

The petrified forest ended up being the only interesting thing I found in the ravine—the area was so dry that I think even the real trees were almost petrified! At any rate, it was clear that I was not going to have much success in this area. I looked at my watch, knowing that it would take three hours to drive back to Beaver Dunes, and estimated that if I left now I could get in about three hours of collecting at Beaver Dunes where I’d had so much success the previous day. Thus, I did what I rarely do on a collecting trip—drive during the afternoon!

Beaver Dune

The main dune at Beaver Dunes State Park, Beaver Co., Oklahoma.

A chunky grasshopper nymph inhabiting the dune

A chunky grasshopper nymph inhabiting the main dune.

I arrived back at Beaver Dunes with several hours of daylight still remaining, so I decided to take a look around the main dunes before heading towards the woody plants. I’ve actually visited Beaver Dunes previously, on the tail end of a fall tiger beetle trip in 2011. At that time I had seen only the rather common and widespread species Cicindela formosa (Big Sand Tiger Beetle) and C. scutellaris (Festive Tiger Beetle), but I thought there could still be a chance to see the much less common C. lengi (Blowout Tiger Beetle). Early June, however, is a little late to see the spring tigers, and in fact I saw only a single C. formosa. Nevertheless, I find dune habitats irresistible—alien habitats occupied by strange plants and animals, and I spent a bit of time exploring the main dune before heading back towards where I had collected so many Buprestidae the previous day.

Low water levels in the reservoir at Beaver Dunes are a result of three years of drought.

Low water levels in the reservoir at Beaver Dunes are a result of three years of drought.

Western Oklahoma, like many parts of the central U.S., has suffered rather severe drought conditions for the past several years. This was evident not only in the large amount of branch dieback seen in the woody vegetation of the area (and probably a contributor to my success at collecting Buprestidae) but also the very low water level in the park reservoir. In the photo above the small cottonwood saplings in the foreground and large cottonwood trees in the left background indicate the normal water level. Cottonwoods, of course, like to keep their feet wet, and the trees around this reservoir—left high and dry by the drought—have responded with major branch dieback and lots of subsequent adventitious sprouting at the bases of the main branches. It was from this adventitious growth that I had beaten most of the Poecilonota cyanipes that I collected the previous day, so I repeated the cottonwood circuit in the hopes of collecting more. Not only did I collect more, but I collected twice as many as the previous day, so I ended up with a very nice series of more than two dozen individuals of the species from the two days collecting. I also did a little more beating of the hackberry trees which had produced well the previous day and collected several more Chrysobothris caddoC. purpureovittata, and Agrilus spp. such as A. leconteiA. paracelti, and perhaps others. When I arrived I was unsure whether I would stay here the following day, but eventually I decided I had sampled the area about as well as I could and that I would go back to the Gloss Mountains for my last day in Oklahoma. Thus, as the day began to wane I began hiking back to the car and spent the next two hours driving back to Woodward to spend the night.

Steep slope below the main mesa | Gloss Mountains State Park, Major Co., Oklahoma

Steep slope below the main mesa | Gloss Mountains State Park, Major Co., Oklahoma

Echinocereus sp. | Gloss Mountains State Park, Major Co., Oklahoma

Echinocereus sp. | Gloss Mountains State Park, Major Co., Oklahoma

Arriving at the Gloss Mountains the next morning was like coming home! I’ve spent so much time at this place and found so many great insects, yet every time I come here I find something new. Today, however, my goals were more modest—I wanted to improve on my series of Paratyndaris prosopis and Chrysobothris texanus, so I focused most of my time beating the hackberry and juniper on top of the mesa and continued beating the juniper down below as well. Success! I collected four more Paratyndaris off of the hackberry, but the C. texanus were far more abundant on this day than they were earlier in the week—I probably got another two dozen individuals of this species. Of course, I also got distracted taking photographs of a number of things, so the day went far more quickly than I realized. I wanted to leave around 6 pm and get in about three hours of driving so that I would have time to make it into Missouri the next morning and have a nice chunk of time to collect before finishing the drive and arriving home on Saturday night. It was actually closer to 7:30 pm before I hit the road, the reason for the delay being the subject of a future post (I will say that BioQuip’s extendable net handle comes in handy for much more than collecting tiger beetles!).

Dolomite glades | Hercules Glades Wilderness, Taney Co., Missouri

Dolomite glades | Hercules Glades Wilderness, Taney Co., Missouri

Long Creek | Hercules Glades Wilderness, Taney Co., Missouri

Long Creek | Hercules Glades Wilderness, Taney Co., Missouri

For my last day of collecting, I decided to stop by at one of my favorite spots in the White River Hills of extreme southwestern Missouri—Hercules Glades Wilderness in the Mark Twain National Forest. I’ve been to this place a number of times over the years, but in recent years my visits have usually been late in the season to look for the always thrilling to see Cicindelidia obsoleta vulturina (Prairie Tiger Beetle). It had actually been about 25 years since I’d visited these glades during the spring, and because of the success I’d had collecting in Oklahoma I was really optimistic that I would find the same here. Sadly (and inexplicably), insect activity was very low, and it didn’t take long for this to become apparent as branch after branch that I beat along the trail through the dry-mesic forest down to Long Creek yielded nothing. By the time I got to the creek I still had not collected a single beetle. A consolation prize was found along the creek, as beating the ninebark (Physocarpos opulifolius) produced a few specimens of the pretty little Dicerca pugionata, and a couple more consolation prizes were found further up the trail approaching the main glade when I saw a Cylindera unipunctata (One-spotted Tiger Beetle) run across the trail and then beat a single Agrilus fuscipennis from a small persimmon (Diospyros virginiana) tree at the edge of the glades. It had been about 25 years since I last collected the latter species, so I was very happy to see it, but no more were seen despite beating every persimmon tree that I saw during the rest of the day. At the end of the day, I had hiked seven miles and collected only six beetles—a rather inauspicious ending to what was otherwise a wonderfully successful trip.

A rare ''selfie''

The author takes a rare ”selfie” at Gloss Mountains State Park.

Arriving back at the car at the end of the day on the last day of an extended collecting trip is always a little depressing—despite the vagaries of travel, cheap hotel beds, meals on the go, and general exhaustion, I’m never happier than I am when I am in the field. Still, the success that I’d had during this trip did much to ease my depression, and arriving home late that night and seeing my girls again (who waited up for me!) finished off any remaining depression.

© Ted C. MacRae 2013

My favorite of Missouri’s milkweeds

Milkweeds of the genus Asclepias are among my favorite plants, although I’m not fully sure why that is the case. Sure, their blooms are conspicuous and colorful, but so are those of many other plants. Perhaps one reason is their status as hosts for milkweed beetles (genus Tetraopes, family Cerambycidae). Four species of these beetles occur in Missouri, including the rare T. texanus. Another reason might be their diversity—in Missouri alone there are 16 different species, ranging from the ubiquitous common milkweed (A. syriaca) to the federally endangered Mead’s milkweed (A. meadii). The latter is one of six milkweed species occurring in Missouri that I have not yet seen, so I suppose I should withhold judgement until I’ve succeeding in finding all 16 species. Nevertheless, I would have to say that clasping milkweed (A. amplexicaulis) has to be my favorite of Missouri’s milkweeds.

Clasping milkweed (Asclepias amplexicaulis) | Sand Prairie Conservation Area, Scott Co., Missouri

Clasping milkweed (Asclepias amplexicaulis) | Sand Prairie Conservation Area, Scott Co., Missouri

Clasping milkweed (also known as sand milkweed—not to be confused with A. arenaria occurring further west in the Great Plains) is said to occur sporadically throughout Missouri in prairies, glades, rocky open woods, roadsides, and railroads. However, I have seen this species only a few times—all in dry sand habitats in the southeastern Mississippi Alluvial Plain (or, the “bootheel” as we say here in Missouruh). Until a few  years ago the only time I had ever seen this plant was many years in an eroded sandy opening on Crowley’s Ridge (an elevated ridge of alluvium and loess deposited during the last glacial maximum). Those plants were not in flower, but their was no mistaking their identity due to their erect stems and broad, cordate-clasping, tomentulose leaves with wavy margins. I would see this plant again a few years ago during my first visit to Sand Prairie Conservation Area, and although I would see it again on many subsequent visits, at no time did I succeed in seeing the blooms.

This species is characterized by broad, clasping, tomentulose leaves with wavy margins.

Broad, clasping, tomentulose leaves with wavy margins.

Finally, last year, I returned to Sand Prairie during late April (a weather-delayed installment of my Annual-Birthday-First-Bug-Collecting-Trip-of-the-Year). I had actually gone there to photograph Missouri’s unique intergrade population of the Festive Tiger Beetle (Cicindela scutellaris), but the weather was cool and the beetles apparently had decided to remain in their burrows. A bad day of collecting, however, is still better than a good day of just about anything else—perhaps because there are almost always consolation prizes, and my consolation prize on this day was my first sight of clasping milkweed plants in full bloom.

A single inflorescence atops each stem.

A single inflorescence atops each stem.

I may not be exactly sure why I like milkweeds so much, but I think I now know why I like clasping milkweed above all others. The softly colored green and pink blossoms are exquisite, to be sure, but more importantly the species is firmly linked in my mind to one of my favorite Missouri habitats. I imagine that clasping milkweed might be an attractive, if somewhat gangly, addition to a native wildflower garden. However, I’m not sure I would enjoy cultivated plants in my garden as much as I do seeing wild plants in one of Missouri’s rarest and most endangered natural communities.

Sand Prairie Conservation Area, Scott Co., Missouri

Sand Prairie Conservation Area, Scott Co., Missouri

Copyright © Ted C. MacRae 2013

Rush skeletonplant pea gall wasp

Lygodesmia juncea with galls of Antistrophus lygodesmiaepisum (Hymenoptera: Cynipidae) on stem.

The Loess Hills landform along the western edge of Iowa and extreme northwestern Missouri is home to a unique assemblage of plants and animals.  The majority of these are associated with loess hilltop prairies – grassland remnants that have their origins in the hypsithermal maximum of several thousand years ago and that persist as small relicts on the landform’s steep, dry, south- and west-facing slopes.  Many of the plants and animals found in these grassland remnants are more typically found further west in the Great Plains, but hang on in the Loess Hills as hypsithermal relicts.

Antistrophus lygodesmiaepisum galls on stem of Lygodesmia juncea.

One such hypsithermal relict is rush skeletonplant, Lygodesmia juncea, a wirey, leafless-looking plant in the family Asteraceae¹.  More common in the Great Plains, this plant occurs in Missouri only on these loess hilltop prairie remnants.  The first time one encounters this plant, they are left with the impression that the plant bears small, pea-like fruiting structures along the length of its stem.  These are not fruiting structures, however, but galls made by the cynipid wasp Antistrophus lygodesmiaepisum.  Although this insect does not have a common name, it is associated exclusively with L. juncea, as suggested by its specific epithet (which also alludes to the pea-like galls with the suffix -pisum), so I see no reason why this wasp cannot be called the “rush skeletonplant pea gall wasp.”  Some sources variably misspell the genus as Anistrophus (without the first “t”) or the species name as simply pisum, a synonym first introduced by Ashmead in the late 19th century a few years after the species was described (I made both mistakes [and also erroneously referred to L. juncea as skeletonweed] in one of my earliest posts: The Loess Hills in Missouri).  It would seem that Antistrophus lygodesmiaepisum is the correct name, according to Pickering (2009).

¹ Not to be confused with rush skeletonweed, Chondrilla juncea – also in the Asteraceae, which despite the similarity of common names, specific epithet, and general appearance (except with yellow flowers) is an altogether different plant that was introduced from the Mediterranean Region and is now considered an invasive weed in much of the Great Plains.

Antistrophus lygodesmiaepisum larva in gall on stem of Lygodesmia juncea.

Rush skeletonplant exudes a latex-like sap when damaged, making it unpalatable to most grazers – this latex-like sap can be seen when the galls made by the wasps are cut open.  Cynipid wasps are the second most diverse group of gall-making insects behind the gall midges, and many species are mono- or oligophagous (Ronquist and Liljeblad 2001), meaning that they are associated exclusively with a single plant species or group of closely related species.  Antistrophus lygodesmiaepisum is one such monophagous species, thus its occurrence in Missouri, like that of L. juncea, is restricted to the tiny loess hilltop prairie remnants in extreme northwestern Missouri.  In recent years, these prairie relicts have suffered heavily from conversion to agriculture, abusive grazing, and suppression of fire that has led to invasion by woody and exotic plants.  In Missouri, only about 50 acres of loess hilltop prairie remain, and only half of these are in conservation ownership, making it among the most critically imperiled of natural communities in Missouri.  While lacking the conservation charisma of L. juncea and the dozen or so other plants and vertebrates that are restricted in Missouri to these prairie remnants, A. lygodesmiaepisum nevertheless deserves equal consideration as a Missouri species of conservation concern.

I knew this would be a difficult ID Challenge and am quite impressed that at least a few people figured out at least the correct genus.  Tim Eisele scored 8 points in this challenge to not only take the win but also move way up into a 3-way tie for 4th place in the overalls.  Ben Coulter continues to be Mr. Consistency, earning 4 points for 2nd place and retaining his overall lead by an almost insurmountable margin (see what happens when you play every game!).  JasonC beat out the other contenders for the final podium spot on the basis of a bonus point, but the hot contest continues to be the battle for 2nd place overall.  Janet Creamer still holds it at 14 pts, but there is a veritable gaggle of contenders nipping at her heals – the next few challenges could be interesting.

Photo 1: Canon 100mm macro lens (ISO 100, 1/250 sec, f/2.8).
Photos 2-3: Canon MP-E 65mm 1-5X macro lens (ISO 100, 1/250 sec, f/14).
All photos: Canon 50D , Canon MT-24EX flash w/ Sto-Fen + GFPuffer diffusers. Typical post-processing (levels, minor cropping, unsharp mask).


Pickering, J.  2009.  Database of Hymenoptera in America north of Mexico. (accessed 20 Jan 2011).

Ronquist, F. and J. Liljeblad.  2001.  Evolution of the gall wasp-host plant association.  Evolution 55(12):2503–2522.

Copyright © Ted C. MacRae 2011

Cicindela nebraskana – Prairie Long-lipped Tiger Beetle

Cicindela nebraskana - the prairie long-lipped tiger beetle

We were only halfway through Day #1 of five days in the field and had already achieved Goal #1 of the trip.  Despite that, it took a few hours before Chris and I were ready to tear ourselves away from our first stop in Fall River Co., South Dakota, where we were treated to the sight of glittering, wine-red adults of Cicindela pulchra bejeweling the charcoal-colored shale slopes.  However, the list of species that we wanted to see over the next several days was long, and eventually our pulchra-fever abated (barely) enough to head south to the Pine Ridge in Sioux Co., Nebraska to look for A-list Species #2 – Cicindela nebraskana.  Sioux Co., Nebraska is the type locality for this species (thus the name), but in reality it is a more western species whose distribution just barely sneaks into the northwestern corner of Nebraska (Pearson et al. 2006, Spomer et al. 2008).  I first saw this species at this very site two years ago, seeing only a handful of individuals and managing one harshly-sunlit, point-and-shoot image of one of them.  To my knowledge, this remains the only known field photograph of this species.

Shortgrass prairie atop the Pine Ridge, Sioux Co., Nebraska (photo taken September 2008).

This time, with a Canon 50D camera and 100mm macro lens in my backpack, I was much better equipped for vastly improved field photographs, but in contrast to the numerous individuals of C. pulchra that we saw earlier in the day, only a single C. nebraskana would turn up after intensive searching by Chris, Matt Brust, and myself in the vast shortgrass prairie sitting at the type locality atop the Pine Ridge.  I didn’t find it – Matt did – and the general rule with rare tigers is to capture the first individual rather than try to photograph it.  If no others are seen, photographing it later in a terrarium of native soil is better than trying to photograph it in the field and risk letting it escape.  Matt gave it to Chris, and at the end of the day when we realized we were not going to see another one, we prepared a terrarium of native soil, taking care to keep the surface as intact as possible so that an accurate replication of the field situation could be created when we photographed it later.

Tiger beetles "hunker down" when fatigued.

Although I prefer actual field photographs, the nice thing about photographing tiger beetles in confinement is… well, they don’t run away!  That’s not to say it is easy.  While they do settle down if left undisturbed for a while, once you start messing with them they quickly become agitated and start running in circles around the terrarium perimeter.  Much finger prodding is necessary to get them away from the edge and into a good spot for photographs, and rarely do they stay put for long.  When they finally do settle down, they tend to “hunker down” in a most unflattering pose (as above) – lacking the appearance of alertness that gives the true field photos their life.

A ferocious pose is struck after judicious prodding of the face and touching of the antennae.

I’m a persistent (syn. stubborn) sort, however, and I’ve learned that I can wear them down and poke and prod them out of their hunker.  Just a light poke at the face will often make them back up and lift their front slightly – poke again and they often open their jaws half-cocked – a light touch on the tip of one antennae and they’ll turn slightly.  With practice and patience, hunkered down beetles can be coaxed into some remarkably aggressive-looking poses.  I like the last of these photos in particular because the oblique, jaws half-cocked pose shows off two nice features of this species – the quite long labrum (upper lip) compared to most other tiger beetle species, and the bright white labrum and mandibles of the males of this species (in females they are partially or completely dark).  The long labrum and jaws give this species a very long-faced appearance that distinguishes it immediately from the black morphs of Cicindela purpurea audubonii that occur with much greater frequency in the same habitats as C. nebraskana.

Photo Details:
1: Canon 50D w/ MP-E 65mm 1-5X macro lens (ISO 100, 1/250 sec, f/13), Canon MT-24EX flash w/ Sto-Fen + GFPuffer diffusers.
2: Panasonic DMC-FX3 (ISO 100, 1/400 sec, f/5.6), natural light.
3-4: Canon 50D w/ 100mm macro lens (ISO 100, 1/250 sec, f/16), Canon MT-24EX flash w/ Sto-Fen + GFPuffer diffusers.
All photos: Typical post-processing (levels, minor cropping, unsharp mask).


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.

Spomer, S. M., M. L. Brust, D. C. Backlund and S. Weins.  2008. Tiger Beetles of South Dakota & Nebraska.University of Nebraska, Department of Entomology, Lincoln, 60 pp.

Copyright © Ted C. MacRae 2010

The real Eleodes suturalis

I recently posted a photograph of a clown beetle (Eleodes hispilabris) (family Tenebrionidae) that I found last July in the Glass Mountains of northwestern Oklahoma.  I had encountered that individual while stumbling through the mixed-grass prairie in the middle of the night in search of the Great Plains giant tiger beetle (Amblycheila cylindriformis).  Although I eventually found the latter species, it took a few hours, during which time I was forced to examine numerous individuals of another clown beetle, Eleodes suturalis – perhaps the most conspicuously common clown beetle in the Great Plains.  I didn’t bother to take photographs of them, focused as I was on my tiger beetle search and owing to the fact that this was not the first time I’d encountered that species in abundance (the first time being many, many years ago as they crossed the highway en masse just a few miles north of the Glass Mountains in Barber Co. Kansas).  In fact, I was becoming rather annoyed with them due to their great similarity in size and coloration to the object of my desire¹, and only when I found the previously photographed individual doing the defensive “head stand” so characteristic of the group did I relent and break out the camera for a series of shots (not easy in the dark of night).

¹ Wrigley (2008) even suggested a mimetic association for Amblycheila cylindriformis and Eleodes suturalis due to their similarity in size, shape and coloration (black with a reddish-brown sutural stripe).

Of course, that individual turned out not to be E. suturalis, but the closely related species E. hispilabris, a fact that I did not realize until several days later as I was examining the photographs more closely. Fortunately, I happened to bring home with me a live individual of what truly represents E. suturalis, which I show in these photographs.  I’m not sure exactly why I brought a live one home with me – I’ve done more and more of this in recent years, mostly just to observe them and see what they do.²  I think in this case, I was intrigued by the possible mimetic association between this species and A. cylindriformis and wanted an individual for comparison with the several live A. cylindriformis individuals that I also brought back with me.

² The singular focus on collecting “specimens” that I had during my younger years seems to be giving way to a desire to know more about species as living entities and not just their external morphology.

Unlike E. hispilabris (my identification of which I only consider tentative), there can be little doubt that the individual in these photographs represents E. suturalis.  No other clown beetle in the Great Plains exhibits the sharply laterally carinate elytra and broadly explanate (spread outward flatly) pronotum (Bernett 2008).  The reddish-brown sutural stripe of the distinctly flattened elytra is also commonly seen in this species, although occasional individuals of a few other clown beetle species exhibit the stripe as well (including E. hispilabris, which likely was the reason I assumed it represented E. suturalis).  All of the characters mentioned above can be seen in the photographs shown here.  However, I nevertheless find the photos rather unsatisfying.  If you think you know why, feel free to comment, otherwise you can wait for the “better” photos…

Photo Details: Canon 50D w/ 100mm macro lens (ISO 100, 1/250 sec, f/18), Canon MT-24EX flash w/ Sto-Fen diffusers. Typical post-processing (levels, minor cropping, unsharp mask).


Bernett, A. 2008. The genus Eleodes Eschscholtz (Coleoptera: Tenebrionidae) of eastern Colorado. Journal of the Kansas Entomological Society 81(4):377–391.

Wrigley, R. A.  2008. Insect collecting in Mid-western USA, July 2007.  The Entomological Society of Manitoba Newsletter 35(2):5–9.

Copyright © Ted C. MacRae 2010

North America’s largest centipede

As I prowled the remote mixed-grass prairie of northwestern Oklahoma in the middle of the night, an enormous, serpentine figure emerged frenetically from a clump of grass and clambered up the banks of the draw I was exploring.  Although I was still hoping for my first glimpse of the Great Plains giant tiger beetle, I was keeping a watchful eye out for anything that moved within the illuminated tunnel of my headlamp due to the potential for encountering prairie rattlesnakes (perhaps the most aggressive of North America’s species).  This was clearly no snake, but at up to 8″, Scolopendra heros (giant desert centipede) easily matches some smaller snakes in length.  Also called the giant Sonoran centipede and the giant North American centipede, it is North America’s largest representative of this class of arthropods (although consider its South American relative, S. gigantea – the Peruvian or Amazonian giant centipede, whose lengths of up to 12″ make it the largest centipede in the world).

Although I had never before seen this species alive, I recognized it instantly for what it was.  Many years ago I was scouting the extreme southwest corner of Missouri for stands of soapberry (Sapindus saponaria), a small tree that just sneaks inside Missouri at the northeasternmost limit of its distribution, in hopes of finding dead branches that might be infested with jewel beetles normally found in Texas.  I had heard that these centipedes also reach their northeastern extent in southwestern Missouri, and just a few miles from the Arkansas and Oklahoma borders I found a road-killed specimen.  I stood there dejected looking at it – too flattened to even try to salvage for the record.

Centipedes, of course, comprise the class Chilopoda, which is divided into four orders.  The giant centipedes (21 species native to North America) are placed in the order Scolopendromorpha, distinguished by having 21 or 23 pairs of legs and (usually) four small, individual ocelli on each side of the head (best seen in bottom photo).  The three other orders of centipedes either lack eyes (Geophilomorpha) or possess compound eyes (Scutigeromorpha and Lithobiomorpha).  These latter two orders also have only 15 pairs of legs (shouldn’t they thus be called “quindecipedes”?).  Among the scolopendromorphs, S. heros is easily distinguished by its very large size and distinctive coloration.  This coloration varies greatly across its range, resulting in the designation of three (likely taxonomically meaningless) subspecies.  This individual would be considered S. h. castaneiceps (red-headed centipede) due to its black trunk with the head and first few trunk segments red and the legs yellow.  As we have noted before, such striking coloration of black and yellow or red nearly always indicates an aposematic or warning function for a species possessing effective antipredatory capabilities – in this case a toxic and very painful bite.

The individual in these photographs is not the first one I saw that night, but the second.  I had no container on hand to hold the first one and not even any forceps with which to handle it – I had to watch in frustration as it clambered up the side of the draw and disappear into the darkness of the night.  Only after I returned to the truck to retrieve a small, plastic terrarium (to fill with dirt for the giant tiger beetles that I now possessed) did I luck into seeing a second individual, which I coaxed carefully into the container.  It almost escaped me yet again – I left the container on the kitchen table when I returned home, only to find the container knocked onto the floor the next morning and the lid askew.  I figured the centipede was long gone and hoped that whichever of our three cats that knocked the container off the table didn’t experience its painful bite.  That evening, I noticed all three cats sitting in a semi-circle, staring at a paper shredder kept up against the wall in the kitchen.  I knew immediately what had so captured their interest and peeked behind the shredder to see the centipede pressed up against the wall. The centipede had lost one of its terminal legs but seemed otherwise none the worse for wear – its terrarium now sits safely in my cat-free office, and every few days it enjoys a nice, fat Manduca larva for lunch.

There are a number of online “fact sheets” on this species, mostly regarding care in captivity for this uncommon but desirable species.  I highly recommend this one by Jeffrey K. Barnes of the University of Arkansas for its comprehensiveness and science-focus.

Photo Details: Canon 50D (ISO 100, 1/250 sec) w/ Canon MT-24EX flash in white box.
Photos 1-2: Canon 100mm macro lens (f22), indirect flash.
Photo 3: Canon MP-E 65mm 1-5X macro lens (f/13), direct flash w/ Sto-Fen + GFPuffer diffusers.
Post-processing: levels, minor cropping, unsharp mask.

Copyright © Ted C. MacRae 2010

Flaming the debate

Eastern redcedar encroaching loess hilltop prairie, a critically imperiled natural community in Missouri.

ResearchBlogging.orgAs my interest in prairie insects has increased over the past few years, so has my interest in their conservation. Many insects are restricted to prairies through dependence upon prairie plants or their unique physical and trophic characteristics. Thus, preservation of not only prairie plants but their insect associates as well is a major goal of conservationists.  The task is daunting – for example only ~1% of tallgrass prairie remains in the central U.S., the rest long ago converted to agriculture or otherwise irreparably altered.  Prairies are dynamic natural communities that rely upon disturbance – this need to “disturb to preserve” creates an oxymoronic conundrum for restoration ecologists that is made even more difficult by the fragmented nature of today’s prairie landscape.  The situation here in Missouri is even more difficult, as nearly all of our grassland preserves (tallgrass prairie, sand prairie, loess hilltop prairie and glades) are exceedingly small and highly disjunct relicts not connected as parts of larger systems.

In recent years, prescribed burning has become the management tactic of choice for restoring and maintaining grassland preserves.  There are good reasons for this – not only are increased floral diversity and reversal of woody encroachment well-documented responses to fire, but burning is also highly cost-effective (a critical consideration in today’s climate of shrinking public budgets).  As the use of prescribed burning on grassland preserves has become widely adopted, however, concerns about the impacts of fire on invertebrate populations have been raised.  The subject is now an area of intense research, but studies are hampered by the limited availability of large, long-unburned tracts of native prairie, and no scientific consensus has yet emerged.  Regrettably, the debate has polarized into “pro-” and “anti-fire” camps that seem unable to communicate with each other constructively.  This is unfortunate, since both ends of the spectrum offer ideas that could be used to achieve the goal of preserving prairie remnants while mitigating concerns about invertebrate impacts.  I have previously expressed my own views on the subject, a position that I suspect some might mistakenly characterize as “anti-fire.”  While I do support the use of prescribed burning, I do not support its use with no consideration of other prairie management strategies such as haying and light grazing (not to be confused with the heavy, abusive, unmanaged kind of grazing that has degraded so much of our landscape).  All of these tools (as well as parcels that receive no management at all) have potential value in prairie management and should be considered.

Those interested in potential fire impacts on prairie invertebrates will be interested in this latest salvo by Scott Swengel and colleagues, who used metadata analysis to correlate declines of prairie butterflies in the Midwest with the widespread adoption of prescribed burning as a management tactic.  The authors present convincing evidence that tallgrass prairie butterfly populations are not co-evolved with fire regimes currently used for prairie management, although their conclusions will no doubt be challenged.  Nevertheless, until a firmer scientific consensus can be achieved, prudence should dictate some measure of caution in the use of fire as an exclusive prairie management tactic.

Dear Colleagues:

We are pleased to announce a new article by Scott Swengel, Dennis Schlicht, Frank Olsen, and Ann Swengel, based on long-term data that has just been published online,  Declines of prairie butterflies in the midwestern USA.  This paper is available free from Springer Open Choice at or by going to the Journal of  Insect Conservation Online First section and scanning through the articles in ascending number order until getting to articles posted 13 August 2010.

The trends of tallgrass prairie skippers shown here, although disastrous, underestimate the decline in Iowa and Minnesota for several reasons:

  1. In statistical testing we only include sites with adequate data for testing, which eliminates many sites from inclusion that had 100% declines of a specialist we know about.
  2. Nearly all sites with long time series were the top sites to begin with, which are likely to take a longer time to show large declines than medium or low-quality sites.
  3. Recent government sponsored surveys not included here show another round of huge declines for Poweshiek Skipperling in Iowa and Minnesota.
  4. Some species went undetectable by the late 1980s and early 1990s, so didn’t register as a presence when the study began.  Hence, they cannot show a decline since then.

Some good news is that conservation based on existing knowledge of specialists’ management responses gets far better results (as shown by Regal Fritillaries and Karner Blues in Wisconsin than typical management.  So declines like this are not inevitable.

The Ecological Interpretations and Conservation Conclusion section of Discussion contain some of our new insights explaining the observed about land-use effects on prairies and butterflies.

Scott Swengel

My thanks to Scott Swengel for giving me permission to reprint his introduction.


Swengel, S. R., D. Schlicht, F. Olsen & A. B. Swengel. 2010. Declines of prairie butterflies in the midwestern USA Journal of Insect Conservation: DOI 10.1007/s10841-010-9323-1.

Copyright © Ted C. MacRae 2010