Group mimicry in Cerambycidae… and more

During last year’s extended visit to Argentina, I had the chance to spend the early part of April in the northern province of Chaco. Though much of this hot, arid plain has been converted to agriculture, remnants of thorn forest remain along fence rows and in small patches of Chaco Forest. Despite the decidedly tropical latitude of the region, however, the profuse bloom of Chilean goldenrod, Solidago chilensis, along these fence rows during the Argentine autumn is reminiscent of crisp fall days here in the eastern U.S., and like the goldenrod here the ubiquitous stands of yellow blossoms stretching across the Chaco Plain are equally attractive to a multitude of insects. Among those insects are the Cerambycidae, or longhorned beetles, and while the eastern U.S. cerambycid fauna of goldenrod boasts only a few (albeit spectacular) species in the genus Megacyllene, the Argentine cerambycid fauna that I found on these flowers included at least three species in various genera belonging to two different tribes.

Rhopalophora collaris (Germar 1824) | Chaco Province, Argentina

Two of the species I saw are shown here, and their similarity of appearance is no coincidence, as both belong to the tribe Rhopalophorini (coming from the Greek words rhopalon = club and phero = to bear, in reference to the distinctly clavate, or club-shaped, legs exhibited by nearly all members of the tribe). In fact, a great many species in this tribe exhibit the same general facies—slender in form and black in coloration with the head and/or pronotum red to some degree. Since all of these species are diurnal (active during the day) and frequently found on flowers, one can assume that the members of this tribe represent an example of what Linsley (1959) called ‘group mimicry.’ In this simple form of Batesian mimicry (harmless mimic with protected model), a group of related species within a genus or even a tribe have a general but nonspecific resemblance to those of some other group of insects—in this case presumably small, flower-visiting wasps. Although the tribe is largely Neotropical, the nominate genus Rhopalophora does extend northward with one eastern U.S. representative, R. longipes. Among the numerous species occurring in South America, the individuals I saw in Argentina can be placed as R. collaris due to the relative lengths of their antennal segments and uniquely shaped pronotum (Napp 2009).

Cosmisoma brullei (Mulsant 1863) | Chaco Province, Argentina

The second species could easily be mistaken for another species of Rhopalophora were it not for the distinct tufts of hair surrounding the middle of the antennae. These tufts immediately identify the beetle as a member of the large, strictly Neotropical genus Cosmisoma (derived from the Greek words kosmos = ornament and soma = body, a direct reference to the tufts adorning the antennae of all members of this genus). Three species of the largely Brazilian genus are known from Argentina, with the black and red coloration of this individual easily identifying it as C. brullei (Bezark 2o13). In the years since this genus was described, additional related genera have been described that bear remarkably similar tufts of hair not on the antennae, but on the elongated hind legs. The great, 19th century naturalist Henry Walter Bates “tried in vain to discover the use of these curious brush-like decorations” (Bates 1863), and nearly a century later Linsley (1959) conceded that their function still remained unknown. Antennal tufts are actually quite common in Cerambycidae, especially in Australia, and while experimental evidence continues (to my knowledge) to be completely lacking, Belt (2004) records observing “Coremia hirtipes” (a synonym of C. plumipes) flourishing its leg tufts in the air (presumably in a manner similar to waving of antennae) and, thus, giving the impression of two black flies hovering above the branch on which the beetle was sitting. This seems also to suggest a function in defense, with the tufts perhaps serving as a distraction to potential predators in much the same way that many butterflies have bright spots near the tail to draw the predator’s attention away from the head.

REFERENCES:

Bates, H. W. 1863. The Naturalist on the River Amazons. Murray, London, 2 vols.

Belt, T. 2004. The Naturalist in Nicaragua. Project Guttenberg eBook.

Bezark, L. G. 2009. A Photographic Catalogue of the Cerambycidae of the World. Available at http://plant.cdfa.ca.gov/byciddb/

Linsley, E. G. 1959. Ecology of Cerambycidae. Annual Review of Entomology 4:99–138.

Napp, D. S. 2009. Revisão das espécies sul-americanas de Rhopalophora (Coleoptera: Cerambycidae). Zoologia (Curitiba) 26(2):343–356.

Copyright © Ted C. MacRae 2013

The gloriously dichromatic Dasymutilla gloriosa

I have a small collection of velvet ants (family Mutillidae) that I’ve accumulated over the years—not through active collection but more as bycatch from my beetle hunting operations. Velvet ants are, of course, not ants at all, but wasps, and as such the females are—like their winged relatives—quite capable of delivering a painful sting if mishandled. They also tend to be seen running rather frenetically across the ground, making them difficult to guide into a collection vial or grab with forceps. You’ve gotta really want ‘em if you want to collect them. Still, even though I don’t study them I find them interesting enough to pick up on occasion, and with most groups outside of my area of focus the hope is that eventually they will end up in the hands of somebody who actively studies the group. Such is now the case with my mutillid collection, which will be shipped this week to another collector specializing in the group. In return I will be filling some holes in European representation of my collection of Cerambycidae.

Dasymutilla gloriosa, female | Brewster Co., Texas

Without question, the most interesting mutillid species that I’ve encountered is Dasymutilla gloriosa. All mutillids are sexually dimorphic, as only the males are winged, but most also tend to be sexually dichromatic to a greater or lesser degree. No species I am aware of takes this to the same level as D. gloriosa! The males (photo below) are rather typically colored compared to other species in the genus, but the females (photo above) are densely covered with long, strikingly white hairs. While this would seem to make them quite conspicuous, the true effect is the exact opposite as they easily confused with fuzzy plant seed. For this reason they are commonly called thistledown velvet ants. I encountered the female in west Texas in 2003 while walking a mountain trail and at first thought it was the fuzzy seed of a creosote bush (Larrea tridentata) being blown by the wind—except there was no wind! It took me a little while looking closely at it before I could figure out what it actually was. This is the only female of this species that I’ve seen in the wild, and I’ll be a little sad to see it sent to another location.

Dasymutilla gloriosa, male | Riverside Co., California

The male also is the only one I’ve encountered—or at least taken the trouble to collect. I would have never suspected this male, which I collected in southern California in 1991, was the same species as the female that I collected many years later. My thanks to Kevin Williams, who provided the identifications for both of these specimens.

Also called the ”thistledown velvet ant”

Copyright © Ted C. MacRae 2012

Gulf Fritillary in southern Missouri

Gulf Fritillary (Agraulis vanillae) | Mississippi Co., Missouri

I’m not sure, but I think this might be the first time I’ve photographed a butterfly caterpillar. Not a bad subject to start with, as few butterflies have caterpillars that are more colorful than the Gulf Fritillary, Agraulis vanillae. A common resident in the southern states and further south, this member of the nymphalid subfamily Heliconiinae is less commonly encountered in Missouri—in fact, I had never seen (or at least noticed) these caterpillars before encountering a few feeding hungrily on the foliage of maypop (Passiflora incarnata) growing in a city park in Missouri’s southeastern lowlands. While the stunning colors of these caterpillars are a delight to human eyes, their function, as in most butterfly caterpillars, is to advertise the unpalatability of their toxin-laced bodies. In the case of this species, the toxins include cyanogenic glycosides that the larvae sequester from the tissues of their host plants (ironically these compounds are supposed to serve the same protective function for the plant that produces them, but butterflies have become master specialists at evolving mechanisms to sidestep toxic impacts).

According to my friends Richard & Joan Heitzman, long-time students of Missouri Lepidoptera, this species is a sporadic migrant that occasionally forms summer colonies in Missouri, especially in the western half of the state, until the first hard freeze destroys the colony (Heitzman & Heitzman 1987).

REFERENCE:

Heitzman, J. R. & J. E. Heitzman. 1987. Butterflies and Moths of Missouri. Missouri Department of Conservation, Jefferson City, 385 pp.

Copyright © Ted C. MacRae 2012

Tracking Tetraopes texanus with Terry

Last month I traveled to Starkville, Mississippi to meet with an academic cooperator at Mississippi State University.  While arranging the trip, I contacted Terry Schiefer (no, not the fashion jewelry designer, but curator at the Mississippi Entomological Museum) to let him know I would be visiting.  Considering that late May should be pretty good insect collecting in that area, I wanted to see if he might be interested in doing a little beetle collecting after I finished up with my meetings.  Terry also specializes in Coleoptera and shares with me an interest in the taxonomy and faunistics of Cerambycidae and Cicindelinae.  I first met Terry some 13 years ago during my previous visit to MSU; I remember ogling at an impressive series of Aegomorphus morrisii, a spectacular species of longhorn beetle that was known at that time by precious few specimens and that he had recently found in Mississippi.  We hadn’t seen each other since but managed to keep in contact with occasional correspondence during the course of our longhorn studies.

Me & Terry Schiefer | Noxubee National Wildlife Refuge, May 2011.

Terry was more than happy to go beetle collecting with me, and among the possibilities that he mentioned when I arrived at the museum was nearby Noxubee National Wildlife Refuge.  I had done a little collecting there on my last visit, but I was especially intrigued when he mentioned the local population of an uncommon milkweed beetle species, Tetraopes texanus, that he had reported in one of the refuge’s prairie remnants (Schiefer 1998).  I have only seen this species once, up here in in east-central Missouri and which I reported as the species’ northernmost known population (MacRae 1994).  My more recent attempts to find this species have not been successful, so I was excited at the chance to see this longhorned species once again.

We arrived at the prairie with plenty of daylight to spare and began walking through the area where Asclepias viridis (its presumed host in Mississippi; in Missouri I found it on Asclepias viridiflora) was growing.  Typically milkweed beetles are quite approachable, having nothing to fear from predators by virtue of the cardiac glycosides that they sequester in their bodies from their milkweed foodplants and advertise so conspicuously with their bright red and black coloration.  Thus, we were looking for beetles sitting brazenly on the plants, but none were seen.  Eventually, Terry saw one in flight, and then I saw one in flight as well.  For some time, this was the only way we were seeing the beetles, and only by slowing down and scanning the prairie vegetation more carefully and deliberately did we begin to see the adults sitting on vegetation.  Interestingly, very few of them were seen actually sitting on milkweed plants.  Rather, they were on all manner of other plants, and they were very quick to take flight on our approach.  This was playing havoc with my desire to get field photographs of the beetles, especially field photographs on the host.  I decided that any photograph—host plant or not—was better than none, so I began attempting some shots.  My first one didn’t work out so well:

Finally I was able to get one of the beetle sitting on a plant, but the dorsal characters can’t be seen, nor is there anything about the photo that allows the species to be distinguished as T. texanus (the abruptly attenuate last antennomere distinguishes it from similar-appearing species):

Progress—more of the dorsal surface can be seen in the photo below, and the beetle is actually sitting on a milkweed plant.  However, the antennal tips are still frustratingly out of focus.  Note the completely divided upper and lower lobes of the eye—Tetraopes beetles give new meaning to the term “four-eyes”:

I chased beetle after beetle in flight, endlessly zigzagging across the prairie in what had to be a spectacle to any unknown observer.  Eventually, we found a beetle sitting on its host plant, and it remained calm during my deliberate approach.  I circled around for a good view of the dorsal surface and snapped off an apparent winner—everything in focus, good composition… but arghh, the antennal tips were clipped!

I kept at it and was about to back off a bit on the magnification and switch to landscape mode so I could get the full antennae in the frame when the beetle turned in a most fortuitous manner—nicely positioning its distinctive antennal tip right in front of a bright green leaf for contrast.  My friends, I present Tetraopes texanus on its presumed host plant, Asclepias viridis!

Terry and I were both puzzled by the flighty, nervous behavior that the beetles were exhibiting.  Neither of us had seen such behavior with milkweed beetles before, and I’m not sure I can offer any explanation for such.  I’d be interested in hearing any ideas you might have.

My thanks to Terry for showing me a few of his favorite spots (allowing me to collect a few choice species of longhorns), and to my co-worker/colleague Jeff Haines for indulging my desire mix a little beetle collecting into the business trip.  I hope they enjoyed it as much as I did.

REFERENCES:

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

Schiefer, T.L. 1998. Disjunct distribution of Cerambycidae (Coleoptera) in the black belt prairie and Jackson prairie in Mississippi and Alabama. The Coleopterists Bulletin 52(3):278–284.

Copyright © Ted C. MacRae 2011

Bichos Argentinos #13 – Spotted Maize Beetle

Astylus atromaculatus (spotted maize beetle) | Inés Indart, Argentina.

One of the most common insects encountered in agricultural fields in Argentina is Asylus atromaculatus (spotted maize beetle).  This native species can also be found further north in Bolivia and Brazil, and as implied by its common name it is frequently encountered in maize fields.  The species, however, is also common on soybean, on which the individual in the above photo (and mating pair in the previous post) were found.  Looking like some strange cross between a checkered beetle (family Cleridae) and a blister beetle (family Meloidae), it is actually a member of the Melyridae (soft-wing flower beetles)—placed with the Cleridae in the superfamily Clerioidea.

Despite its abundance (and the resultant attention it gets from growers), the pollen feeding adults are of little economic importance.  It’s easy to see, however, why this species gets so much attention from growers—during January through March the adults occur in tremendous numbers, congregating on a wide variety of flowering plants, but especially corn. Their large numbers are an impressive sight, with literally dozens to even hundreds of adults occurring on a single plant. Tassles—the source of corn pollen—are highly preferred, but when populations are heavy the silks and any exposed ears are also popular congregation sites. Despite their numbers, the impact of the beetles on yield is rarely sufficient to warrant the cost of control measures.

Whatever economic impact the species might have is actually due more the larvae—hidden within soil—than to the super-abundant and highly conspicuous adults. Feeding primarily on decaying plant matter within the soil, larvae do occasionally attack newly planted corn, either before or just after germination. Their attacks are more common in dry years and in severe cases can lead to the need to replant a field. This seems to be more common in South Africa, where the species was introduced in the early 1900s, than in its native distribution in South America.

Whenever I see a ubiquitous, diurnal, brightly and contrastingly colored insect, the first suspicion that comes to my mind is aposematic (warning) coloration and chemical defense against predation. There seems to have been some investigation into the toxicity of this species (Kellerman et al. 1972), and in South Africa they have been implicated in poisoning of livestock when accidentally ingested with forage (Bellamy 1985).  Few other reports of toxicity by beetles in this family are known, but four species of the genus Choresine have been shown to produce high levels of batrachotoxin alkaloids—these are the same toxins found in the skin of poison-dart frogs of the genus Phyllobates (Dumbacher 2004).  The frogs are unable to synthesize these toxins themselves, thus, it is presumed that they sequester these compounds from their diet—whether it is from some species of Melyridae remains to be determined.

Congratulations to Alex Wild and Max Barclay, who both answered the call to ID Challenge #8 and correctly determined all taxa from order to species.  Alex, by way of submitting his ID first, gets a bonus point and leads the current BitB Challenge session with 9 points.  Thanks to the rest who played along as well—see my response to your comments for your points earnings.

REFERENCES:

Bellamy, C. L. 1985.  Cleroidea, pp. 237–241.  In: Scholtz, C. H. and E. Holm (Eds.), Insects of Southern Africa, Butterworths, Durban.

Dumbacher, W. A., S. R. Derrickson, A. Samuelson, T. F. Spande and J. W. Daly. 2004.  Melyrid beetles (Choresine): a putative source for the batrachotoxin alkaloids found in poison-dart frogs and toxic passerine birds.  Proceedings of the National Academy of Sciences, USA 101(45):15857–15860.

Kellerman, T. S., T. F. Adelaar and J. A. Minne. 1972. The toxicity of the pollen beetle Astylus atromaculatus Blanch. Journal of the South African Veterinary Medical Association 43(4):377–381.

Copyright © Ted C. MacRae 2011

Monday Moth – Polka-dot Wasp Moth

Syntomeida epilais - polka-dot wasp moth

It’s been a while since I’ve done a Monday Moth post, so I thought I’d feature one of the prettier specimens in my very limited Lepidoptera collection.  This is Syntomeida epilais (polka-dot wasp moth), one of four species in the genus that occurs in the United States.  This particular specimen was collected by me way back in the mid-1980s (I was not quite yet the discriminating beetle collector that I am now) in Everglades National Park (yes, I had a permit).  The bright, contrasting coloration obviously screams aposematism (warning coloration), and in fact the tissues of the adult moths of this species are chock-full of several cardiac glycosides sequestered by the larva from its now preferred food plant, oleander (Nerium oleander).  Add to it their somewhat wasp-like appearance, and there should be no question to any would-be predator that these moths are a bad idea.  Wasp moths are related at the tribal level to another group of wasp-like moths called maidens which are restricted to the Old World.  I featured one of these from South Africa last year in the post, Monday Moth – Simple Maiden (Amata simplex).

If the cardiac gycosides stored in the tissues of this moth aren’t enough to cause gastric distress, trying to digest the higher taxonomic history of this group surely will.  Back in my school days, this moth belonged to the family “Ctenuchidae.”  As best I understand it, this group was later subsumed into the tiger moth family “Arctiidae” – itself later subsumed within the borg of all moth families, the Noctuidae.  In the most recent classification I’ve found, the arctiine moths have been pulled back out of the Noctuidae and combined with the former “Lymantriidae” (propelled to infamy by the gypsy moth) to form the family Erebidae (Lafontaine and Schmidt 2010).  Are you ready to purge yet? It’s still not clear to me whether this latest incarnation represents a consensus monophyletic unit, but it really doesn’t matter – whenever I see wasp moths, maidens, and especially the ctenucha moths that are so common in my area on goldenrod flowers during the fall, “ctenuchid” will still be the first name that comes to my mind.

REFERENCE:

Lafontaine, J. D. and B. C. Schmidt.  2010.  Annotated check list of the Noctuoidea (Insecta, Lepidoptera) of North America north of Mexico.  ZooKeys 40:1–239.  doi: 10.3897/zookeys.40.414

Copyright © Ted C. MacRae 2011



Brazil Bugs #1

ID Challenge #3 update: I knew this would be a hard one, and so far nobody has figured this one out (only one commenter got the right order!).  I’ve released the comments gotten so far so you can see where things stand, and maybe with the additional information (and my pointing out an important clue) it will be enough for one of you to arrive at a full ID.  Further comments are still being moderated until I decide to close the challenge.  Right now the points are there for the taking!

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This past Monday I embarked on an extended business trip to Brazil.  “Wow, Brazil!” – you say, and while getting to travel to an exotic tropical locality on my company’s dime definitely rocks, I do have to justify the trip by actually working.  Add that to the time involved with planes, automobiles, and hotel transfers, and there is precious little time for more esoteric activities such as photographing insects.  It is Brazil, however, and summer at that, so whatever time does become available over the next ten days, I’ll be on it!  I’m stationed in Campinas, about 2 hours north of São Paulo, and this evening I had my first opportunity to break out the camera and do a little exploring around the hotel grounds during the waning hours of daylight. I’m not normally one to take short walks just looking for any random insect to photograph, but hey – it’s Brazil!  I also don’t normally like to post photographs of insects without knowing much about them – especially their identity. But hey, it’s Brazil!  I think I’ll be lucky to figure out most things to family, although I might be able to drill down a little further on occasion.  With that prelude, I hope you’ll indulge me these random postings over the next 10 days or so, primarily photographs but perhaps accompanied by a little bit of text.

Coreidae?

At first I thought this was a member of the hemipteran family Pyrrhocoridae (red bugs), as some North American species have similar coloration; however, members of that family apparently lack ocelli, which this insect clearly possesses. This would seem to indicate instead some species of Lygaeidae (seed bugs), but the forewing membrane with numerous veins arising from a transverse basal vein and presence of what appears to be a distinct metathoracic scent gland opening suggest instead some “unleaf-footed” species of leaf-footed bug (Coreidae). Whatever its identity, I would imagine it is quite distasteful, based on what clearly seems to be aposematic coloration and the fact that there were numbers of these bugs hanging out quite conspicuously on these flowers.

Same species?

This individual was differently colored than the others, but otherwise it seemed structurally and behaviorally identical.  Is it merely a highly melanized individual?  Maybe a case of sexual dimorphism, and I only saw this one individual of one of the sexes?  Maybe it truly is a different species – it is Brazil, afterall!

Pseudoplusia includens?

Daylight began to slip away much too soon, and I was about to pack it up when  I noticed some blurs at the flowers.  I realized that moths had begun to visit the flowers in the obscurity of dusk and became determined to photograph one, despite the fact that they never actually landed on the flowers but hovered in front of them instead.  It was quite difficult to even get them framed in the viewfinder, and on those few occasions when I managed to do this it was all but impossible to spend any time trying to focus – I used the lamp on the flash unit to help me see the moth, then just framed and quickly took the shot.  This one actually turned out not too bad – a little bit of blur in the wings but otherwise acceptable enough to let me tentatively identify it as Pseudoplusia includens (soybean looper).  Brazil is well on its way to becoming the world’s largest producer of soybean, and the caterpillars of these moths are enjoying the bounty!

Family Crambidae, possibly Herpetogramma phaeopteralis (ID by Chris Grinter)

I saw one last moth before the final traces of daylight disappeared – I don’t have any idea about its identity, but it’s a pretty picture nonetheless.  It has the generic look of the large family Noctuidae, so that’s what I’m gonna go with until somebody tells me differently.  Edit 1/20/11 – Somebody just told me differently!  According to Chris Grinter, this is a species of Crambidae, possibly Herpetogramma phaeopteralis (tropical sod webworm).  Thanks, Chris!

I don’t know the name of the plant whose flowers these insects were visiting, but the hotel staff has promised to ask their gardener in the morning and let me know – now that’s service!

Copyright © Ted C. MacRae 2011

Desmocerus palliatus – elderberry borer

Desmocerus palliatus (elderberry borer), Squaw Creek Natl. Wildlife Refuge, Missouri.

Last June I made two trips to the Loess Hills in northwestern Missouri to survey additional sites for Cylindera celeripes (swift tiger beetle), which my colleague Chris Brown and I had discovered in some of the area’s few remaining loess hilltop prairie remnants the previous year. One of these potential new sites was  Squaw Creek National Wildlife Refuge, where a few tiny slivers of hilltop prairie can still be found on the fingers of loess bluffs that border the refuge’s several thousand acres of restored wetlands that famously host large concentrations of snow geese and bald eagles during the fall and spring migrations.  On the first visit, I had arranged to meet with Corey Kudrna, Refuge Operations Specialist, who was kind enough to take several hours out of his day to personally guide me to each of the site’s loess hilltop prairie remnants. 

As we crossed the highway right-of-way at the base of the bluffs on our way to the one of the remnants, we passed through a large patch of common elderberry, Sambucus nigra ssp. canadensis.  Anytime I see patches of this plant, especially in June, I immediately think of Desmocerus palliatus (elderberry borer) – a spectacularly colored longhorned beetle (family Cerambycidae) that breeds exclusively in the living stems and roots of this plant.  It is not a particularly rare species, but for some reason I have not had much success in finding this species.  In my close to three decades of collecting beetles, I had encountered perhaps a half dozen individuals – never more than two at the same time.  Still, when I get the chance to look at elderberry I look for this beetle, and when I did so this time I was delighted to see one within a few moments of entering the patch.  I was ecstatic when I saw another one almost immediately after the first, and I was stunned when I realized that they were all around me!  Good fortune continued on my subsequent visit two weeks later, when I was able to spend a little more time trying to get a good field photograph.  Wind was a problem, the beetles were easily alarmed, and their tendency to rest in the upper reaches of the plant made it difficult to brace myself and the camera while shooting, making this a rather difficult subject to get a good photograph of.  The photo shown here is literally the last of around two dozen that I took and is the only one that I really like.

Many cerambycid beetles are mimics of other more noxious species, mostly ants and wasps.  However, elderberry borers appear to be the exception in that they are themselves noxious.  The cobalt blue and bright orange coloration of the adults screams aposematic (warning) coloration, and it is reasonable to assume that they accumulate in their bodies for defensive purposes the cyanogenic glucosides produced by elderberry plants (Huxel 2000).  Even their movements are those of a chemically protected model - lumbering and clumsy, without the alert evasiveness usually seen with other flower longhorn species.  Presumably this species participates in a Müllerian mimicry complex involving netwinged beetles (family Lycidae, particularly species in the genus Calopteron) and perhaps Pyromorpha dimidiata (orange-patched smoky moth, family Zygaenidae) as well, and it may serve as a Batesian model for the equally colorful but completely innocuous Lycomorpha pholus (black-and-yellow lichen moth, family Arctiidae).

Photo Details: Canon 50D w/ 100mm macro lens (ISO 100, 1/250 sec, f/10), Canon MT-24EX flash w/ Sto-Fen diffusers.

REFERENCE:

Huxel, G. R.  2000.  The effect of the Argentine ant on the threatened valley elderberry longhorn beetle.  Biological Invasions 2:81–85.

Copyright © Ted C. MacRae 2010