Flower ants? Check again!

Last spring while hiking the North Fork Section of the Ozark Trail in southern Missouri (Howell Co.), I made sure to check the abundant flowering dogwood (Cornus florida) blossoms that were in gorgeous peak bloom at the time (early May). I’ve learned to check flowers of dogwood whenever I can, as they are quite attractive to a variety of insects but especially those groups of longhorned beetles (family Cerambycidae) that tend to frequent flowers as adults. In the case of flowering dogwood, most of the cerambycids that I encounter belong to two genera: Molorchus and Euderces. Both of these genera are known for their great resemblance to small ants, no doubt representing examples of Batesian mimicry (where a harmless species adopts the appearance or warning signals of a harmful species to gain protection from predators).

Tilloclytus geminatus

Tilloclytus geminatus on Cornus florida | North Fork Section, Ozark Trail, Howell Co., Missouri

During this particular hike I was determined to photograph Molorchus bimaculatus, common in Missouri during early spring on a great variety of flowering trees. On this day, however, the tiny (<10 mm length) beetles were rather scarce, and I had been frustrated in my attempts to get good photographs of the few that I had found. I’ve seen enough of these beetles over the years that I can recognize them quickly for what they are without the need to closely examine every “ant” that I see. So when I saw an “ant” that was too big and convex in profile to be Molorchus I almost discounted it as a true ant. Something about it, however, gave me pause, and when I looked closer I saw that it was, indeed, a longhorned beetle. But, it was not Molorchus, nor was it Euderces. Instead, it was the species Tilloclytus geminatus—an exciting find!

Tilloclytus geminatus

Adults in profile greatly resemble ants of the same size.

Tilloclytus geminatus has been recorded only sporadically from across the eastern U.S., where it has been reared from a variety of deciduous hardwoods (Craighead 1923, Rice et al. 1985). Perry (1975) did report rearing this species from Pinus virginiana (along with several other species normally associated with hardwoods); however, that record likely represents an ‘‘overflow’’ host (Hespenheide 1969) that is not typical of the species’ normal host preferences. I myself had never seen the species until the years after I published my checklist of Missouri cerambycids (MacRae 1994), having succeeded in rearing adults from a variety of previously unrecorded hardwood hosts that I collected at several localities across southern Missouri (MacRae & Rice 2007). It remains, for me, an infrequently encountered species—perhaps part of this a result of being overlooked due to its effective ant mimicry.

Tilloclytus geminatus

The anterior, oblique markings give the illusion of a constricted “waist”, while the posterior, transverse markings resemble the “sheen” of a shiny abdomen.

Unlike Molorchus and Euderces, this species has not been frequently associated with flowers as adults. In fact, the only report I am aware of is that of Rice et al. (1985), who reported adults on flowers of hawthorn (Crataegus sp.). Perhaps this additional find on Cornus is indicative of a true adult attraction to flowers by T. geminatus, although a single adult provides only weak support. However, a related ant-mimicking longhorned beetle—Cyrtophorus verrucosus—has been collected on flowers of roughleaf dogwood (Cornus drummondii) (MacRae 1994) as well as flowering dogwood (Scheifer 1998a). The floral attraction of ant-mimicking cerambycids may be more characteristic of species in the subfamily Cerambycinae, as only one flower record exists (Physocarpus opulifolius) for Psenocerus supernotatus (Wheeler & Hoebeke 1985) and none exist for Cyrtinus pygmaeus, both in the subfamily Lamiinae rather than Cerambycinae and the only other true ant-mimicking species in Missouri of which I am aware.

REFERENCES:

Craighead, F. C. 1923. North American cerambycid larvae. A classification and the biology of North American cerambycid larvae. Dominion of Canada, Department of Agriculture, Technical Bulletin No. 27 (new series), 239 pp. [Internet Archive].

Hespenheide, H. A. 1969. Larval feeding site of species of Agrilus (Coleoptera) using
a common host. Oikos 20:558–561 [JSTOR].

MacRae, T. C. 1994. Annotated checklist of the longhorned beetles (Coleoptera: Cerambycidae and Disteniidae) known to occur in Missouri. Insecta Mundi 7(4) (1993):223–252 [pdf].

MacRae, T. C. & M. E. Rice. 2007. Distributional and biological observations on North American Cerambycidae (Coleoptera). The Coleopterists Bulletin 61(2): 227–263 [pdf].

Perry, R. H. 1975. Notes on the long-horned beetles of Virginia, part III (Coleoptera: Cerambycidae). The Coleopterists Bulletin 29(1):59 [JSTOR].

Rice, M. E., R. H. Turnbow Jr. & F. T. Hovore. 1985. Biological and distributional observations on Cerambycidae from the southwestern United States (Coleoptera). The Coleopterists Bulletin 39(1):18–24 [pdf].

Schiefer, T. L. 1998a. A preliminary list of the Cerambycidae and Disteniidae (Coleoptera) of Mississippi. Transactions of the American Entomological Society 124(2):113–131 [JSTOR].

Wheeler, A. G., Jr. & E. R. Hoebeke. 1985. The insect fauna of ninebark, Physocarpus opulifolius (Rosaceae). Proceedings of the Entomological Society of Washington 87(2):356–370 [BioStor].

© Ted C. MacRae 2015

Just how widespread is Prionus heroicus?

On our insect collecting trip to the western Great Plains last June, Jeff Huether and I encountered several species of beetles in the genus Prionus—longhorned beetles (family Cerambycidae) known commonly as “prionid root borers”. I’ve already recounted our experiences with two of them—P. integer in southeastern Colorado and P. fissicornis in northeastern New Mexico, both of which we encountered unexpectedly and by the “bucketload” using prionic acid baited traps in shortgrass prairie habitats. The third species we encountered was also rather unexpected, so much so that we were not even sure about its identity until we examined the collected specimens more closely. Unlike the two previously mentioned species, however, the behavior of this third species—P. heroicus, especially regarding its attraction to prionic acid, was quite different, and one of the localities where we found it raises some interesting questions regarding its recorded distribution.

Prionus heroicus male

Prionus heroicus male | Mills Canyon, Harding Co., New Mexico.

Prionus heroicus is among the largest species in the genus, and we first encountered it in northeastern New Mexico while searching for suitable habitat around the small town of Mills (Harding Co.) where several grassland-associated species in the genus had been found by other collectors. While we were searching we were intrigued by the USDA Forest Service road sign pointing to “Mills Canyon” and decided to head down the gravel road to see what it looked like. A few miles down the road the endless shortgrass prairie quickly changed to sandstone canyonland with oak/pine/juniper woodland. By now it was late afternoon, and the rich, pulsing buzz of cicadas filling the air as we drove slowly through the area suggested the potential for good insect activity. We stopped to take a closer look at things, and within minutes an enormous prionid beetle came flying up to the car—apparently attracted to the prionic acid lures inside!  We netted the beetle, and shortly afterwards another beetle flew into the vicinity… and another! We hurriedly set out some lure-baited traps, and over the next hour or so we collected close to a dozen beetles. Our first thought, based on their large size and the presence of 12 antennal segments, was that they might be P. californicus. This was a bit of a letdown, as our goal was to find little known and uncommonly encountered Great Plains species, not common and widespread western species like P. californicus. However, the blackish rather than reddish color and relatively broad pronotum eventually convinced us that we had actually found P. heroicus—not a rare species, but also not something we expected to see at this relatively northeastern location compared to its usual haunts in southeastern Arizona.

Prionus heroicus male

Males generally landed in the vicinity of the prionic acid lures but seemed unable to locate its precise whereabouts.

Interestingly, none of the beetles came directly to the lure-baited traps but, rather, flew to the vicinity and then either circled broadly in the air or landed some distance from the trap and ran rapidly on the ground as if searching. We missed a few that we tried to net in mid-air, which responded by immediate straight line flight far away into the distance. Those that we watched while on the ground searched for up to several minutes but seemed unable to locate the lure, eventually taking flight and flying away. This suggests that prionic acid is more important as a component of the female’s calling pheromones for long distance detection, but that other components might be necessary to allow the males to locate females in the immediate vicinity. We returned to the site the following day around noon and were surprised to see males flying to our lures immediately after our arrival, since we had observed P. integer and P. fissicornis activity to occur almost exclusively during the morning and evening hours. Linsley et al. (1961), however, also note that adults of this species fly during the day, as well as at night. Again, none of the beetles seemed capable of finding the precise location of the lure.

Prionus heroicus male

Males rapidly searched the ground after landing but seemed unable to locate the lures and eventually flew off.

The larval habits of P. heroicus are not known, but the closely related P. californicus feeds on roots of living oaks or occasionally in dead oak logs or stumps (Linsley 1962). Prionus heroicus may have similar habits, since adult females have been found in leaf litter at the bases of living oak trees and stumps (Hovore & Giesbert 1976, Skiles 1978). Presumably the Mills Canyon population is associated with Gambel oak (Quercus gambelii), which was the only oak species noted at this location. An interesting defensive behavior was observed for an adult beetle that I kept alive and photographed later in the hotel room. I had put the beetle on a light-colored tray, and while I was getting the camera ready to take some shots the beetle began crawling away. I picked it up and put it back into position, and for a brief period it assumed the rather comical-looking “standing-on-its-butt” pose shown in the accompanying photograph. It maintained this pose for some seconds and then gradually lowered itself in the front before beginning to crawl again. If I poked it before it finished lowering itself it immediately pushed its front all the way back up again and held the pose briefly before beginning to lower itself again. I did not encounter this behavior in the field—beetles on the ground immediately tried to flee when I handled them and took flight if allowed to.

Prionus heroicus male

Defensive posturing as a result of being disturbed.

The story does not end here, however, as this would not be our last encounter with the species. While we were traveling back towards the east, we decided to visit the area around Black Mesa in the extreme northwestern corner of the Oklahoma panhandle. I had been to this site the previous summer, and although conditions were very dry at the time the presence of Gambel oak in the area—perhaps the easternmost outpost of this western species—had me thinking about the possibility of the area serving as an unknown eastern outpost for western species of jewel beetles (family Buprestidae). Conditions were once again dry, and no jewel beetles were found, but this quickly became a moot point as not long after our arrival a large prionid beetle came flying up to me—again in the same manner as P. heroicus at Mills Canyon and in the middle of the afternoon. We decided these, too, represented P. heroicus and remained in the area for the rest of the day, counting at least a dozen males attracted to the vicinity of the lures that we’d set out. As with the males we observed in New Mexico, none seemed able to find the precise location of the lure, and some circling only briefly and then flew off without even landing. Unlike New Mexico, however, we did succeed in trapping a few males in lure-bated traps that we set out overnight and checked the next morning.

Prionus heroicus male

This Prionus heroicus male  near Black Mesa, Oklahoma confirms the presence of this species in the state.

Our capture of P. heroicus at Mills Canyon in northeastern New Mexico and near Black Mesa in northwestern Oklahoma not only seems to suggest an association of this species with Gambel oak, but also confirms the previously questionable occurrence of the species in Oklahoma. Alexander (1958) reported the species (under one of its synonyms, P. tetricus) from Payne Co. in north-central Oklahoma. This record is dubious, since Payne Co. is well east of the 100th meridian and has a decidedly eastern flora, including its oaks. That record has not been mentioned by any subsequent authors addressing the overall distribution of P. heroicus (Linsley 1962, Chemsak 1996). The presence of P. heroicus in the Black Mesa area, with its decidedly western flora, does not necessarily give credence to the Payne Co. record, but it does bring up the question of just how broadly distributed P. heroicus really is. Most records of this species are from mountainous areas of southern Arizona and New Mexico, but more recently it has been recorded from Texas (Big Bend National Park) by Van Pelt (1999, 2007) and extreme northwestern Colorado (Moffat Co.) by Heffern (1998). The latter record is particularly interesting, as this represents a considerable range extension from any previously recorded locality and begs the question regarding its occurrence in other parts of Colorado.

REFERENCES:

Alexander, D. R. 1958. A preliminary survey of the cerambycids (Insecta: Coleoptera) of Oklahoma. Proceedings of the Oklahoma Academy of Science 38:43–52 [pdf].

Chemsak, J. A. 1996. Illustrated Revision of the Cerambycidae of North America. Volume I. Subfamilies Parandrinae, Spondylidinae, Aseminae, Prioninae. Wolfsgarden  Books, Burbank, California, x + 149 pp., plates I–X [description].

Heffern, D. J. 1998. Insects of Western North America 1A Survey of the Cerambycidae (Coleoptera) or Longhorn Beetles of Colorado. Gillette Museum of Arthropod Diversity, Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, i + 32 pp., 1 color plate [summary].

Hovore, F. T. & E. F. Giesbert. 1976. Notes on the ecology and distribution of western Cerambycidae (Coleoptera). The Coleopterists Bulletin 30:350 [JSTOR].

Linsley, E. G. 1962. The Cerambycidae of North America. Part II. Taxonomy and classification of the Parandrinae, Prioninae, Spndylinae, and Aseminae. University of California Publications in Entomology 19:1–102, 1 plate [OCLC WorldCat].

Linsely, E. G., J. N. Knull & M. Statham. 1961. A List of Cerambycidae from the Chiricahua Mountain area, Cochise County, Arizona (Coleoptera). American Museum Novitates 2050:1–34 [pdf].

Skiles, D. D. 1976. Taxonomy and description of two prionine Cerambycidae from southern Arizona: a new species of Stenodontes and new status for Neomallodon arizonicus (Coleoptera). Proceedings of the Entomological Society of Washington 80:407–423 [Biodiversity Heritage Library].

Van Pelt, A. F. (ed.). 1999. Inventory of insects of Big Bend National Park, Texas. Big Bend Natural History Association, Big Bend National Park, and Texas.

Van Pelt, A. F. (ed.). 2007. Inventory of insects of Big Bend National Park, Texas. Report to Big Bend National Park, 204 pp.

© Ted C. MacRae 2015