During the 6-week period from late May to early July this year, I collected ~400 jewel beetle specimens representing at least 20 species (see Working with Cerceris fumipennis—Part 1). A final accounting of the species represented won’t be done until this winter, but the genera represented include Acmaeodera, Actenodes, Agrilus, Anthaxia (Haplanthaxia), Buprestis (Knulliobuprestis), Chrysobothris, Dicerca, Poecilonota, and Spectralia. Perhaps two-thirds of the specimens were “ground-picked”¹, while most of the remaining third were “stolen” directly from wasps by netting wasps in flight as they returned to their nest carrying prey.

¹ It’s not clear to me why I found so many abandoned buprestids at nest entrances. The wasps are known to drop prey when threatened and, rather than search for and relocate the prey, fly off to look for a new beetle(Careless et al. 2009). I observed this myself in several cases when I missed netting the wasp but swiped the net close enough to scare it, at which time it dropped the beetle and flew off (and I popped the beetle in a vial). However, the bulk of the beetles I found on the ground were not only at the nest entrance, but even mixed within the diggings surrounding the nest. My first act when checking each field was to check each nest, pick up any adults lying on top of the burrow diggings, and then carefully spread out the diggings with a knife or trowel to collect the beetles hidden within them. One nest contained as many as 13 Agrlus obsoletoguttatus inside the diggings. I wondered at one point if the wasps were leaving the beetles at the burrow entrance and then digging out the burrow before coming back to retrieve them, but I never actually witnessed this. On the other hand, I observed numerous wasps approaching their burrows while carrying prey, and every time the wasp dropped directly into the burrow. In fact, I could even predict what beetle species I was likely to find inside the nest based on the species I found around the entrance (more on that below).

This ball field with contains several dozen Cerceris fumipennis nests.

There is a third method that I used to collect beetles that I haven’t yet discussed, and that is digging them out of nests. In the latter part of the survey period (late June and early July), beetle numbers dropped rapidly, as did apparent wasp activity. As mentioned in the previous post, this drop off in activity came precisely at the time of season when I have observed buprestid beetle activity to decline in Missouri. As the drop off in activity was taking place, I began wondering what I would find if I tried digging up some of the burrows. Of course, digging up a nest takes much more effort than netting wasps or picking beetles up off of the ground, so it becomes important not only to identify whether a nest actually belongs to C. fumipennis and if it is active and likely to contain freshly captured beetles.  In addition, I observed the burrows of a variety of other insects in these fields as well, some of which are shown here and which might be confused with burrow entrances of C. fumipennis.

Cerceris fumipennis nest with Chrysobothris sp. adult left on diggings.

Cerceris fumipennis burrows exhibit perfectly circular, pencil-sized entrances surrounded by a symmetrical mound of diggings with a fine rather than granular texture. There are other Cerceris species that make nearly identical burrows, but they prey on other insects rather than buprestid beetles. At my site I found C. bicornis, a weevil specialist, almost as common as C. fumipennis. Their burrow entrances on the whole seemed slightly larger, but I could not use this as consistent distinguishing character. What I could use, however, was the presence of weevils rather than buprestids lying on the ground near the nest entrance. (I also observed this species returning to its nest and noted a rather faster, more powerful flight that made them even more difficult to capture than C. fumipennis). In contrast, there can be no doubt that the burrow above, with a buprestid beetle lying on the ground near the entrance, belongs to C. fumipennis.²

² The white plastic tag marks the burrow to facilitate locating nests on subsequent visits. It is secured with a golf tee and also can be rotated so that the hole covers the entrance. The hole is large enough to allow the wasp to leave but too small for a returning wasp to enter while carrying a beetle. The idea was to rotate the tags when I first entered a field to cover all the burrow entrances, watch for wasps returning with prey, and then net the wasps as they tried (in vain) to enter the burrow. However, I never actually observed a wasp trying to enter a covered burrow, even after leaving a field and returning 20–30 minutes later.

I presume this nest to be that of Bembix americana (sand wasp).

For the first few weeks, I thought the burrows such as that shown in the above photo also belonged to C. fumipennis. However, I never found beetles lying on the ground near the entrance, nor did I ever observe a wasp to enter or leave the burrow. I eventually noticed several distinct differences in burrow architecture—the burrow entered the ground at an angle rather than straight down, the diggings were distributed asymmetrically to one side of the entrance, and the latter seemed consistently a little larger than those of C. fumipennis. In addition, these burrows always seemed to be in the sandier portions of the fields. While I never associated any insect directly with these burrows, I did observe sand wasps (perhaps Bembix americana) in the vicinity and have seen similar-looking burrows dug by these wasps at Sand Prairie Conservation Area.

Larval burrows of Cicindelidia punctulata and other tiger beetles lack diggings around the entrance.

Tiger beetle larval burrows might also be confused with C. fumipennis burrows, especially after rain or high winds which can wash/blow away the diggings from around the entrance. I found adults of the punctured tiger beetle, Cicindelidia punctulata, fairly commonly at the site and presume the numerous tiger beetle larval burrows that were also present belong to that species. Larval tiger beetles burrows also enter the ground straight down and are, at first appearance, also perfectly round, but they are usually a little too small for C. fumipennis (those of Tetracha spp. being an exception)—the presumed C. punctulata burrow in the above photo measures about 5 mm in diameter. In addition, closer examination reveals a slight “D” shape to the burrow entrance (upper right in the above photo—the tiger beetle larva rests its jaws against the flat side) and, more distinctively, beveling of the ground around the rim of the burrow entrance. Cerceris fumipennis nests lack the slight D-shape and distinctive beveling.

Use a grass stem as a guide while carefully digging away the surrounding soil.

Years of practice digging up tiger beetle burrows prepared me well for my first attempts at digging up C. fumipennis burrows. While it might seem an easy task to follow a hole into the ground while digging soil away from it, in practice the burrow can be quickly lost after even a few inches due to falling soil covering the hole and making it impossible to relocate. I use a thin, flexible but sturdy grass stem to preserve the burrow path, inserting the stem into the burrow and down as far as it will go and then removing the soil carefully from around the hole with a knife or trowel. I try to avoid letting soil fall over the hole by prying the soil away from the hole, but if the hole does get covered the grass stem allows it to be easily relocated.

This nest contained a single Buprestis rufipes…

Cerceris fumipennis burrows are not very deep—only 10–15 cm, and angle to one side a few cm below the surface before leveling out near the bottom. I noticed the nest in the above photo because I saw a wasp fly into it. When I went over to look at it I found a Buprestis rufipes lying on the ground near the entrance and so decided to dig it up. As I expected, I found another B. rufipes at the bottom of the burrow (two above photos courtesy of Madison MacRae).

…while this one contained a cache of seven Agrilus quadriguttatus.

The above photo shows a cache of seven Agrilus quadriguttatus that I found at the bottom of another burrow. In this case, the prey is rather small compared to large prey such as Buprestis and Dicerca. While nests provisioned with species in these latter genera often contained only a single beetle in them, I nearly always found multiple beetles in nests provisioned with the smaller Agrilus species. One nest contained as many as 13 Agrilus obsoletoguttatus, among the smallest of the species I found utilized by C. fumipennis at this site.

Buprestidae taken from five different Cerceris fumipennis nests.

Some of the nests I dug up contained multiple species of beetles, but far more commonly I found only a single species in a given nest. The photo above shows the diversity and number of beetles found on one date after digging up five different nests. From top left the beetles are: 1) 1 Buprestis rufipes; 2) 2 Agrilus quadriguttatus and 1 A. obsoletoguttatus; 3) 2 A. quadriguttatus and 1 A. obsoletoguttatus; 4) 8 A. obsoletoguttatus; and 5) 2 Poecilonota cyanipes, 2 A. quadriguttatus, and 1 A. pseudofallax. It would make sense for wasps to provision nests with greater numbers of smaller beetles to ensure adequate food for their larvae to complete development. How the wasps actually locate their prey, and why this species has specialized almost exclusively on buprestid beetles, is a mystery (at least to me); however (and here comes the speculation du jour), I suspect the wasps may have keyed in on volatiles used by the beetles—either those released by suitable hosts or by each other to facilitate mate location. Use of buprestid pheromones or freshly dead host volatiles would allow wasps to more efficiently locate buprestid prey and, once locating a source (a tree harboring a particular beetle species), could return repeatedly to provision their nest fully. It seems less likely that wasps rely exclusively on visual location of prey, as this would involve a large amount of random searching through trees and passing up numerous, seemingly equally suitable prey.

REFERENCE:

Careless, P. D., S. A. Marshal, B. D. Gill, E. Appleton, R, Favrin & T. Kimoto. 2009. Cerceris fumipennis—a biosurveillance tool for emerald ash borer. Canadian Food Inspection Agency, 16 pp.

Copyright © Ted C. MacRae 2012