2019 Arizona/New Mexico/California Insect Collecting Trip iReport

This is the eighth “Collecting Trip iReport”; this one covering a one-week trip to southern Arizona, New Mexico, and California from September 7–14, 2019 with meloid/cerambycid-enthusiast Jeff Huether. Jeff has been a frequent collecting trip partner during recent years, this being our sixth joint outing since 2012. Our initial objective on this trip was to collect cerambycid beetles of the genus Crossidius occurring across southern Arizona/California—part of a larger effort to sample as many of the named subspecific taxa as possible from multiple locations (including type locations when possible) for future molecular studies. We had good success, though we did not collect every taxon that we were after (we were a tad early in soCal). Also, the fact that we had Crossidius as our primary goal did not mean that we would not concurrently be on the lookout for buprestids (me), meloids (Jeff), or other cerambycids (both of us)—and in that regard we were also successful.

As with all previous “iReports” in this series, this report is illustrated exclusively with iPhone photographs (thus the term “iReport”), with previous versions including the following:
2013 Oklahoma
2013 Great Basin
2014 Great Plains
2015 Texas
2018 New Mexico/Texas
2018 Arizona
2019 Arkansas/Oklahoma


Day 1 – Dripping Springs Mountains, Arizona
First stop of the trip, and we’re heading east to Safford. As soon as we got east of Superior up into the mountains we saw a place where Heterotheca subaxillaris was in bloom abundantly along the roadsides and pulled over. There were four species of Acmaeodera on the flowers, and I also found a fifth species on the flower of a small white aster. Nice first stop for the trip!

Dripping Spring Mountains.
Acmaeodera gibbula on flower of Heterotheca subaxillaris.
Acmaeodera rubronotata on flower of Heterotheca subaxillaris.
Acmaeodera alicia on flower of Heterotheca subaxillaris.

3.6 mi NW Bylas on US-70, Arizona
Continuing our way to Safford, Jeff saw some patches of sunflower and wanted to look for Epicauta phoenix. I found the first two (but not in sunflower), and then Jeff found two more. As we were walking back to the car I noticed a Crossidius suturalis sitting on Isocoma tenuisecta that was not quite in bloom, and then another nearby on the same plant. We searched the area again, but the only plants were those few right around the car.

I’ve never seen an orange jumping spider (family Salticidae) before!

5.7 mi NE Safford, Arizona
After getting a hotel in Safford, we had time to come back to a spot where Jeff had collected Epicauta phoenix back in July. We found quite a few (see photo) on plants nearby the original collection spot. Looking around more I found an Acmaeodera convicta perched on the tip of a shrub—first time I’ve collected this species! There were several species of tenebrionids crawling on the ground, perhaps prompted to activity by cooling temps as rain whipped up in the distance. I kept one eye on the skies and the other on the plants and eventually found two more A. convicta perched together on the same type of shrub just as rain began pelting my back. We made a quick dash back to the car and called an end to Day 1 in Arizona.

Epicauta phoenix (order Coleoptera, family Meloidae).

Day 2 – 1.9 mi S Artesia, Arizona
We started seeing Isocoma tenuisecta just coming into bloom as we headed south of town so stopped to see if we could find any Crossidius. I looked at a lot of plants before finding a single C. suturalis sitting on one of the non-blooming plants and in the meantime found one Trichodes peninsularis and a fair number of Zonitis dunniana on the flowers. Looking around on other plants, I found one large Chrysobothris sp. (not C. octocola, but longer and narrower) on the branch of a living Acacia constricta [Edit: this is C. knulli—a new one for me!] and one Acmaeodera disjuncta, several more Z. dunniana and T. peninsularis on flowers of Hymenothrix wislizeni. Finally, I did some sweeping of the bunch grasses in the area and got a nice series of what I presume to be Agrilus rubrovittatus—first time I’ve collected that species!

Crossidius suturalis on pre-blooming Isocoma tenuisecta.
Trichodes peninsularis on Isocoma tenuisecta.
Zonitis dunniana on flowers of Isocoma tenuisecta.

17.7 mi S Artesia, Arizona
Another stop with both Isocoma tenuisecta and also Gutierrezia microcephala coming into bloom. We immediately began finding Crossidius pulchellus on the latter and eventually collected a good series of them and also Trichodes peninsularis off the plants When I returned to the first plant we had checked (in fullest flower), a Lampetis webbii landed on it right in front of me! I eventually found C. suturalis on Isocoma tenuisecta, as well as Trichodes sp. and a few C. pulchellus. There was a tall-stemmed malvaceous shrub off which I got a male/female pair of Tylosis maculata, and sweeping produced a couple more Agrilus rubronotata, a few more T. peninsularis, and one Acmaeodera scalaris. I saw a couple of Acmaeodera disjuncta on Baileya multiradiata flowers but missed them both!

Stagmomantis limbata (bordered mantis) on Gutierrezia microcephala.
Lampetis webbii on Gutierrezia microcephala.

1.1 mi N Rodeo, New Mexico
We slipped just inside the New Mexico border to visit the area around the type locality of Crossidius hurdi. We found a spot where there were good stands of Isocoma tenuisecta along the roadsides and checked them out. Like the other spots today they were just starting to come into bloom, and rain had just moved through the area. We found perhaps 20 Crossidius individuals total, and honestly they were so variable that I don’t know whether they represent C. suturalis, C. hurdi, or both! [Edit: they are all C. suturalis] I also collected one Sphaenothecus bivittatus and several individuals each of three species of clerids on the flowers of these plants. A male Oncideres rhodosticta was found on the twig of Prosopis glandulosa, and I also found a cool meloid that I’ve never seen before—Megetra punctata!

A particularly well-marked female Crossidius suturalis on flowers of Isocoma tenuisecta.
The coloration of Megetra punctata screams “Don’t eat me or you’ll be sorry!”
Oncideres rhodosticta on Prosopis glandulosa (mesquite).
Mule deerly departed.
Fence row to the Chiricahua Mountains.

Willcox Playa, Arizona
We plan to visit Willcox Playa tomorrow (my inaugural visit!), but we had some time at the end of the day and decided to come take a look. There were some stands of Isocoma tenuisecta at the north end of the playa, and I found just a couple of Crossidius individuals on them, presumably C. suturalis, but it looks like they are bedding down for the evening. Also got a couple of Enoclerus sp. on the flowers.

North end of Willcox Playa.

Day 3 – 8.4 mi SE Willcox, Arizona
On our way towards the Chiricahua Mountains to see if we can find any Crossidius host plant stands. We found patches of Isocoma tenuisecta and Gutierrezia microcephala along Hwy 186 southeast of town—the former was just coming into bloom, but there were plenty of Acmaeodera (scalaris, disjuncta, and amplicollis) on the flowers, including on the unopened heads. We found perhaps a dozen Crossidius suturalis on them as well, and Jeff found one small female that looks like C, hurdi [Edit: it is C. suturalis]. I looked at a lot of Gutierrezia before finding a single C. pulchellus sitting on one of the plants. The same diversity of Acmaeodera as well as a few A. gibbula and T. peninsularis was also found on flowers of Hymenothrix wislizeni, and I took a series of about 10 specimens of what I looks like A. parkeri on flowers of what appears to be Stephanomeria pauciflora. There were also some tiny membracine treehoppers on a thorny shrub (maybe Condalia?) being tended by ants—both adults and young, and I collected a few of the adults.

Acmaeodera amplicollis on flowers of Isocoma tenuisecta.
Acmaeodera disjuncta on flowers of Isocoma tenuisecta.
Acmaeodera scalaris on flowers of Hymenothrix wislizeni.

Jct AZ-186 & AZ-181, Arizona
After passing over a small range towards the Chiricahuas we didn’t see any Isocoma tenuisecta until we got to Hwy 181. There were some Baccharis sarothroides at the junction also, so we stopped and looked around. The Isocoma was just barely coming into bloom, but I found two Crossidius on them—one male C. suturalis and one small female that may be C. hurdi [Edit: nope, it is C. sururalis]. Heterotheca subaxillaris was in bloom abundantly, but there were no Acmaeodera on them and the area in general looked quite dry. I did find two A. decipiens on Sphaeralcea sp., and in the way back to the car I spotted a huge Lampetis webbii hanging on Ericameria nauseosa (which we’re not even close to blooming)—surely an incidental record.

Chiricahua Mountains in the distance.

4.1 mi SE Willcox, Arizona
We came back towards town where things seemed to be further along and found stands of Isocoma tenuisecta in full bloom at the junction of Blue Sky Rd (a classic Arizona collecting locality). Crossidius suturalis were out in numbers on the flowers! Every now and then I got one that seemed too heavily maculated, making me think it could be C hurdi, but in the end I decided that all represented C. suturalis.

Crossidius suturalis mating pair on flowers of Isocoma tenuisecta. Note the difference in antennal length between the male (top) and female.

Willcox Playa, Arizona
We went to the Playa to see if there were any tiger beetles to be had. I hiked to the edge of the Playa, and within a few minutes I saw a Cicindela pimeriana—just the second one I’ve encountered (the first was last night at gas station lights)! With that promise of more, I hiked the entire playa edge and never saw another one! I only saw one other tiger beetle—Cylindera lemniscata—seems I’m a bit late in the season for the Willcox Playa tiger beetles. Nevertheless, it’s a cool place and was fun to see. I’ll definitely be back during the summer, not just for here but for nearby Blue Sky Rd. Arriving back at the car, I did find one Moneilema sp. (I think M. appressum) on cholla. There were Crossidius suturalis abundant on the Isocoma tenuisecta, which, like the last spot, was in full bloom, but I’d gotten my fill of them at the previous spot and didn’t collect any.

Stalking tiger beetles.
Jumping spider out in the playa.
Eking out a living.
Cow tracks.
These Gnathium sp. were the tiniest blister beetles I’ve ever seen.

Willcox, Arizona (epilogue)
Collecting the insects from the field is only the beginning. Each night they must be processed for storage until they can be mounted once back in the lab.

Processing the day’s catch.

Day 4 – Santa Rita Mountains, Box Canyon, Arizona
We passed through Box Canyon on our way to Madera Canyon, so we decided to stop near the dry falls where last year I’d collected such a nice diversity of Acmaeodera spp. on flowers of Allionia incarnata. There was evidence of recent rain, and we found the patch nicely in bloom with four species (scalaris, decipiens, cazieri, and parkeri) on the flowers. Nearby in the wash before it crossed the road was a yellow composite (Xanthisma gracile), from which I collected the first three as well as gibbula, rubronotata, and disjuncta. Euphoria verticalis scarabs we’re flying plentifully around the flowers also—first time I’ve seen the species.

Allionia incarnata (trailing four o’clock) blooming the canyon slope.

Flats below Madera Canyon, Arizona
There are records of Deltaspis tumacacorii from Madera Canton at Proctor Rd collected on Croton, so we stopped by on our way south to give it a try. This seems to be a rather hard-to-find bug, so I didn’t have high expectations, and that’s a good thing because I didn’t see the beetle nor anything that even remotely resembled Croton. I ended up checking out the desert broom (Baccharis sarothroides) in the area on which I’d collected Stenaspis verticalis arizonensis and Tragidion spp. (also without high expectations). There were some interesting congregations of Euphoria leucographa feeding at sap flows on the stems and a few Stenaspis solitaria but otherwise litttle of note. I did find one Hippomelas planicauda hanger-on on a low fabaceous shrub (not Mimosa biuncifera), and inspecting the Gutierrezia microcephala plants revealed nothing but a single Acmaeodera rubronotata.

Stenaspis solitaria on Baccharis sarothroides (desert broom).
Peucetia viridans (green lynx spider) feeding on Acanthocephala thomasi twice its size!
Euphoria leucographa and a Polistes paper wasp feeding at a sap flow on Baccharis sarothroides (desert broom).
Euphoria leucographa on Baccharis sarothroides (desert broom).
Taenipoda eques (lubber grasshopper). The striking coloration is a warning to potential predators that it is chemically protected.

Madera Canyon Rd, Arizona
We stopped real quick down the road on the way out of Madera Canyon because we saw stands of Isocoma tenuisecta, although they were still just shy of blooming. We looked at quite a few and found a single Crossidius suturalis—probably we are a tad early, and the area looks like it could use a good rain to pop things out and bring the Isocoma into bloom. We also saw low plants that could be the Croton that Deltaspis tumacacorii has been found on [Edit: I do not believe these are the plants, as they are too low]. Would be good to revisit this spot after a good rain!

The author with Ferocactus wislizenii (fishhook barrel cactus)—also called “compass barrel” due to its habit of leaning to the south.

Tumacacori Mountains, Walker Canyon, Arizona
Our second shot at Deltaspis tumacacorii, which has also been taken in Walker Canyon. We found thick stands of knee-high flowers that we immediately took to be the Croton—just as described by our contact—on which the beetles have been taken. However, we quickly began doubting that ID and decided the plant must be some type of composite. That would make more sense from a host plant standpoint, as all known host plants for Crossidius spp. are composites (subsequently determined to be Pseudognaphalium leucocephalum, family Asteraceae). We looked at the dense stands for quite some time but didn’t see any beetles (or much of anything else) before deciding that we were probably too early—had the beetles already emerged we would have at least found some stragglers. I did take a few Acmaeodera on the flowers (scalaris and rubronotata), as well as a large cantharid (Chauliognathus profundus). I also took single A. amplicollis and A. rubronotata individuals off of a large helianthoid composite (Viguiera cordifolia) and one A. rubronotata on a small yellow composite (Xanthisma gracile). There were a multitude of darkling beetles crawling in the ground—in one spot I saw five individuals of several species all within a one-square foot area. We’ll have one more shot at D. tumacacorii tomorrow at Kitt Peak.

Pseudognaphalium leucocephalum (white rabbit-tobacco) blooms profusely in the dry wash.
Cantharid vs. cantharid! Chauliognathus profundus (right) feeds on a C. lewisi that it has captured.

Day 5 – Pan Tak, Arizona (road to Kitt Peak)
Today’s destination is Kitt Peak to look for Deltaspis tumacacorii and Acmaeodera resplendens, but at the entrance we saw some Isocoma tenuisecta just coming into bloom and decided to check it out. We found a half-dozen Crossidius suturalis but had to really work for them. Alliona incarnata was also nicely in bloom, but I got only one Acmaeodera parkeri? and one A. alicia off of the flowers. There was some Gutierrezia microcephala present, also not quite in bloom, off of which Jeff got a pair of C. suturalis and gave me one. Kinda dry but lots of flowers—wish there would have been more beetles coming to them.

Moneilema sp. on Cylindropuntia imbricata.

Kitt Peak National Observatory, Arizona
Our last chance to find Deltaspis tumacacorii, and I also got a tip that Acmaeodera resplendens has also been taken up here. We immediately found several species of Acmaeodera (amabilis, amplicollis, decipiens, and rubronotata) abundantly on several composite flowers—Heliomeris longifolia, Heterotheca fulcrata, Hymenothrix wrightii, Solidago velutina, and Gutierrezia microcephala, and I found a single A. solitaria on a pink malvaceous flower, but no A. resplendens. We also searched thoroughly for any Croton-like plant for D. tumacacorii but found nothing. The Kitt Peak records of that species are older than the Walker Canyon, Peña Blanca, and Madera Canyon records, and most of the records seem to be in August rather than September, so I suspect we are a bit late for both the species and its host plant. My plan at this point is to return sometime during the middle of August and enlist the help of the source of one of the recent records to accompany me.

View north from Kitt Peak National Observatory.
Multiple species of Acmaeodera visiting flower of Heliomeris longifolia.
Acmaeodera amabilis on flower of Heliomeris longifolia.

Road to Kitt Peak, Arizona
We had noticed Gutierrezia microcephala and some other yellow composites in bloom about halfway up the mountain on our way to Kitt Peak and decided then to stop and take a look around on the way down. I took “down” the mountain, Jeff took “up.” I hadn’t walked very far when I saw what I at first thought was the cantharid Chauliognathus profundus (which I had seen yesterday at Walker Canyon preying on another cantharid) on G. microcephala flowers, but something about it gave me pause—it was too cylindrical and robust. I leaned closer to get a better look and realized it was a cerambycid—one that I did not recognize, a beautiful orange color with black elytral apices and pronotal spots! I quickly grabbed it with my right hand, immediately saw another elsewhere on the bush and grabbed it with my left hand, and as I stood there trying to fumble a vial out of my pack to put them in I saw a third individual taking flight from the bush and spiraling into the air and out of reach! I shouted out to Jeff, who came down to where I was, and showed him what I’d found, and together we decided that it must be Mannophorus forreri—a very uncommonly encountered species and more than adequate consolation for not finding Deltaspis tumacacorii earlier in the day. We spent the next hour searching up and down the roadsides, and I ended up with two more individuals from Gutierrezia flowers and two from Heterotheca fulcrata. Jeff found an additional individual on flowers of Thelesperma sp. I also picked up a few black and white Enoclerus sp., one on flowers of G. microcephala and a mating pair on flowers of Acacia berlandieri. We have a long drive to California in front of us now, and it sure is good going into the drive with such a great find under our belts.

Lower slopes of the road to Kitt Peak Observatory.
Thasus neocalifornicus (giant mesquite bugs) congregate on their host plant (Prosopis glandulosa).

Day 6 – Cajon Pass, California
Finally made it into California! Once we turned off the interstate, we made a quick stop to look at the roadside habitat where we spotted a good stand of Isocoma sp. in full bloom. We looked at quite a few plants but didn’t find any beetles on them. There were also good numbers of Ericameria nauseosa plants as well (host for Crossidius coralinus), but they weren’t quite yet in bloom yet and the only thing I found on them was a mating pair of Agrilus walsinghami. Moved on quickly to the next spot!

Lancaster, California
We met up with Ron Alten and traveled to a classic “Crossidius” collecting site (up to four species have been taken there). We’d stopped at a couple of places on the way there but not found anything—either the host plants were not yet blooming or no beetles were found, so we had the feeling that we might be a week or two early. We had to drive into the habitat a ways before we started seeing host plants—in this case Ericameria nauseosa—but eventually we found a nice large area with the plants in full bloom. It didn’t take long before we found Crossidius coralinus (populations in this area are assigned to subspecies ascendens) on the blossoms. We worked the area for a couple of hours in the heat (97°F) and got a sufficient series for study with some individuals in ethanol for DNA analysis. Males exhibit quite a bit of variability in the degree of development of the elytral markings (thin to moderately expanded sutural marking), while females were quite consistently fully expanded. Males also outnumbered females by 3:1, and all of the individuals I collected were perfect and not damaged—both suggesting that the species is just beginning to emerge. Perhaps that is why we did not find individuals of the other species (mojavensis, suturalis, and testaceus). What I did find, however, was a small trachyderine cerambycid that none of us recognized! It was on the flowers of E. nauseosa—just like C. coralinus—and at first I thought it might be a small, aberrant C. coralinus, but the elytra are completely blue-black and the size was significantly smaller than the smallest C. coralinus male that we saw. I scanned BugGuide and didn’t find anything that matched, so this will have to remain a mystery for now. [Edit: I later determined this to be a heavily marked C. discoideus blandus. In the field I couldn’t see the orange laterals on the elytra.]

Crossidius coralinus ascendens (male) on flowers of Ericameria nauseosa.
Crossidius coralinus ascendens (female) on flowers of Ericameria nauseosa.
Crossidius coralinus ascendens (mating pair) on flowers of Ericameria nauseosa.

Day 7 – Santa Catalina Mountains, Mt. Lemmon, Arizona
We decided we were just a bit to early for things in California and decided to come back to Arizona where we’d been having better success. I wanted to take another shot at Acmaeodera resplendens and had been told that Oracle Ridge Trail was a good locality for them, though maybe a bit late. We began seeing them soon after getting out of the car—unmistakable by their brilliant metallic green to copper color. They were not numerous, so I had to work for them and walked the trail about 2 miles out collecting them off a variety of flowers. The majority were on Bahia dissecta, and I also found occasional individuals of them and other species of Acmaeodera (amabilis, amplicollis, decipiens, and rubronotata) on flowers of Heliomeris longifolia, Heterotheca fulcrata, Hymenothrix wrightii, Ageratina herbarea, Achillea millefolium, sweeping, Cirsium sp., and prob. Viguiera dentata. One other beetle I found was a Megacyllene sp. sitting on a plant under a stand of Robinia neomexicana [Edit: this is M. snowi snowi—another new one for my collection!].

View from Oracle Ridge Trail @ 1 mile north of the trailhead.
Acmaeodera resplendens on flower of Heliomeris longifolia.
Acmaeodera resplendens on flower of Heliomeris longifolia.

Scenic Overlook, Santa Catalina Mountains, Arizona
A quick stop on the way back down the mountain at a spot where we’d seen Gutierrezia microcephala and Heterotheca subaxillaris blooming along the sides of the road. There wasn’t much going on—a couple of Acmaeodera amplicollis and one A. rubronotata on the flowers of H. subaxillaris, one Enoclerus sp. on Solidago velutina, one A. solitaria on Baccharis sarothroides, and another swept from grasses. I did see Peucetia viridans (green lynx spider) feeding on a very bristly tachinid fly.

A clearwing moth (family Sesiidae).
Peucetia viridans (green lynx spider) feeding on a hairy tachinid fly.

Day 8 – Santa Rita Mountains, Montosa Canyon, Arizona (halfway up)
We decided to visit Montosa Canyon to take another shot at Deltaspis tumacacorii and also see if maybe we could find more Acmaeodera resplendens. We didn’t see many flowers along the way up the canyon until about the halfway point. When we did start seeing them we made a quick stop to see what might be on them. I collected some of the more common Acmaeodera (rubronotata, decipiens, and amplicollis) off a few different yellow composite flowers, but we quickly decided to take a look at the higher elevations.

Panoramic view from halfway up the canyon.
Apyrrothrix araxes (dull firetip skipper) on flowers of Baccharis salicifolia.
The larvae of these large skippers feed on oaks.

Santa Rita Mountains, Montosa Canyon, Arizona (entrance to Whipple Observatory)
The road was gated past the km-13 point—Jeff took the roadsides, and I took a ridgetop trail off to the south for a little over a mile. The panoramic views were spectacular, and at the southern terminus I stood at the edge amidst gale-force winds admiring the landscape! Acmaeodera were diverse and abundant, though not quite as abundant as yesterday on Mt. Lemmon or a few days ago on Kitt Peak. However, I did get another nice series of A. resplendens, along with decipiens, rubronotata, amplicollis, and amabilis. Host flowers were collected for most of these: Hymenothrix wrightii, Erigeron neomexicanus, Linum neomexicanum, Heliomeris multiflora, Verbesina enselioides, Heliopsis parvifolia, Heterotheca subaxillaris, and Machaeranthera tanacetifolia. I also collected a small series of A. decipiens perching on grass stems and a very cool-looking wasp—black with a bright orange thorax and whitish abdominal apex [edit: I believe this is the scoliid Psorthaspis portiae].

Vista from the southern terminus of the ridgetop trail off Mt. Hopkins Rd at km 13.
Acmaeodera decipiens on flower of Machaeranthera tanacetifolia.
A curious assemblage of bees on this Heliomeris longifolia flower. They were not active, just sitting. [Edit: these are Dufourea sp. (short-faced bees, family Halictidae)].

Santa Rita Mountains, lower Montosa Canyon, Arizona
We stopped at a spot near the bottom of the canyon on the way out to see what was going on at the lower elevations. The answer—not much! There were a variety of woody shrubs and other plants in bloom, but the area seemed rather “wilty”. I think this area has a lot of potential, we just didn’t hit it at the right time. I did take an impressively huge tarantula hawk, just because.

Fouquieria splendens (ocotillo) resembles cacti but is not a true cactus. Distantly related to persimmons, blueberries and acacias, it is now placed in its own family (Fouquieriaceae).
Apiomerus flaviventris (yellow-bellied bee assassin) with prey.
The prey is a soldier beetle (likely Chauliognathus lewisi).
Robber flies not only mate tail-to-tail, they fly coupled in the position also. This tandem of giant robber flies (possibly Promachus nigrialbus), flew by me and landed in the bushes. The male (right) tried to take flight again and pulled the female’s hind legs off her perch. She stood firm, however, forcing the male to grab a nearby branch with just his from and middle legs and leave his hind legs dangling also. Note that the female is also feeding on a honey bee—so much natural history going on here!

Flats below Montosa Canyon, Arizona
I was a bit disappointed at not finding any beetles at what seemed would be the last collecting stop of the trip. But on our way out we saw a patch of Isocoma tenuisecta in bloom in the lowlands some distance west of the entrance to the canyon—just what we were looking for! Jeff and I each quickly found Crossidius suturalis individuals on flowers of the plants and continued searching up and down along the roadway. We didn’t find any more for awhile but when I got back to the area where I started I spotted another one sitting on a plant on the other side of the barbed-wire fence. I extended my net handle to the max, maneuvered it in position, took an assertive swipe, and got it. Just as I was putting it into the bottle, I saw another one take flight from a plant right beside me. I hadn’t closed the bottle yet but didn’t want the other one to get away, so I capped my thumb over the opening, awkwardly wielded my net into position one-handed, chased after it and took a swing and got it, too! (More often than not these situations end up with me losing both specimens!). There was also a good amount of Hymenothrix wislizeni along the roadside, off the flowers of which I collected several Acmaeodera gibbula, A. disjuncta, and A. rubronotata. This is probably the last collecting locality of the trip, so I’m happy to end up having success with this subspecies of Crossidius (C. suturalis intermedius), which we havn’t found in large numbers on this trip. Just after leaving the site, we saw a bobcat on the side of the road—my first one! Unusual to see one in the middle of the day—it was a small one, must’ve been quite hungry!

Santa Rita Mountains from the highway.

Phoenix, Arizona (epilogue)
Bill Warner was kind enough to host Jeff and I for our last night in Arizona prior to returning home tomorrow. What an amazing collection he has built, and his use of flight-intercept traps in recent years has turned up even more amazing beetles. I was happy to also meet Andrew Johnston and Evan Waite, who joined us for dinner.

Bill Warner, an icon among Arizona beetle collectors, sits amidst newly collected I material waiting to be processed.
Sunset in Phoenix!

©️ Ted C. MacRae 2019

Hairy milkweed beetle

Across the Great Plains of North America, sand dune fields dot the landscape along rivers flowing east out of the Rocky Mountains. Formed by repeated periods of drought and the action of prevailing south/southwest winds on alluvium exposed by uplifting over the past several million years, many of these dunes boast unique assemblages of plants and animals adapted to their harsh, xeric conditions. Some are no longer active, while others remain active to this day. Among the latter is Beaver Dunes in the panhandle of northwestern Oklahoma.

Beaver Dunes, Oklahoma

Beaver Dunes State Park, Beaver Co., Oklahoma

As I explored the more vegetated areas around the perimeter of the dunes, I spotted the characteristically hairy, fleshy, opposite leaves of Ascelpias arenaria. Known also as “sand milkweed,” this plant is associated with sand dunes and other dry sandy soil sites throughout the central and southern Great Plains. I always give milkweeds a second look whenever I encounter them due to the association with them by longhorned beetles in the genus Tetraopes. It wasn’t long before I spotted the black antennae and red head of one of these beetles peering over one of the upper leaves from the other side.

Tetraopes pilosus on Asclepias arenaria

Tetraopes pilosus on Asclepias arenaria | Beaver Dunes State Park, Oklahoma

This was no ordinary Tetraopes, however. Its large size, dense covering of white pubescence, and association with sand milkweed told me immediately that this must be T. pilosus (the specific epithet meaning “hairy”). Like its host, this particular milkweed beetle is restricted to Quaternary sandhills in the central and southern Great Plains (Chemsak 1963), and also like its host the dense clothing of white pubescence is presumably an adaptation to prevent moisture loss and overheating in their xeric dune habitats (Farrell & Mitter 1998).

Tetraopes pilosus

Species of Tetraopes have the eyes completely divided by the antennal insertions—thus, “four eyes.”

Tetraopes is a highly specialized lineage distributed from Guatemala to Canada that feed as both larvae and adults exclusively on milkweed (Chemsak 1963). Larval feeding occurs in and around the roots of living plants, a habit exhibited by only a few other genera of Cerambycidae but unique in the subfamily Lamiinae (Linsley 1961). Milkweed plants are protected from most vertebrate and invertebrate herbivores by paralytic toxins, commonly termed cardiac glycosides or cardenolides. However, a few insects, Tetraopes being the most common and diverse, have not only evolved cardenolide insensitivity but also the ability to sequester these toxins for their own defense. Virtually all insects that feed on milkweed and their relatives have evolved aposematic coloration to advertise their unpalatability, and the bright red and black color schemes exhibited by milkweed beetles are no exception.

Species of the genus Tetraopes are characterized by the completely divided eyes.

Adult beetles, like the leaves of their hosts, are clothed in white pubescence.

As  noted by Mittler & Farrel (1998), variation in coloration among the different species of Tetraopes may be correlated with host chemistry. Milkweed species vary in toxicity, with more basal species expressing simpler cardenolides of lower toxicity and derived species possessing more complex and toxic analogs. Most species of Tetraopes are associated with a single species of milkweed, and it has been noted that adults of those affiliated with less toxic milkweeds on average are smaller, have less of their body surface brightly colored, and are quicker to take flight (Chemsak 1963, Farrell & Mitter 1998). Thus, there seems to be a direct correlation between the amount of protection afforded by their host plant and the degree to which the adults advertise their unpalatability and exhibit escape behaviors. Asclepias arenaria and related species are the most derived in the genus and contain the highest concentrations of cardenolides. In fact, they seem to be fed upon only by Tetraopes and monarchs while being generally free from other more oligophagous insect herbivores such as ctenuchine arctiid moths and chrysomelid beetles that feed on less derived species of milkweed (Farrell & Mitter 1998). Accordingly, T. pilosus is among the largest species in the genus and has the majority of its body surface red. Also, consistent with it being more highly protected than others in the genus, I noted virtually no attempted escape behavior as I photographed this lone adult.

Asclepias arenaria

Asclepias arenaria (sand milkweed) growing at the base of a dune.

In addition to metabolic insensitivity to cardenolides, adult Tetraopes also exhibit behavioral adaptations to avoid milkweed defenses (Doussard & Eisner 1987). The milky sap of milkweed is thick with latex that quickly dries to a sticky glue that can incapacitate the mouthparts of chewing insects that feed upon the sap-filled tissues. Adult Tetraopes, however, use their mandibles to cut through the leaf midrib about a quarter of the way back from the tip. This allows much of the sticky latex-filled sap to drain from the more distal tissues, on which the beetle then begins feeding at the tip. Leaves with chewed tips and cut midribs are telltale signs of feeding by adult Tetraopes.

REFERENCES:

Chemsak, J. A. 1963. Taxonomy and bionomics of the genus Tetraopes (Coleoptera: Cerambycidae). University of California Publications in Entomology 30(1):1–90, 9 plates.

Doussard, D. E. & T. Eisner. 1987. Vein-cutting behavior: insect counterploy to the latex defense of plants. Science 237:898–901 [abstract].

Farrell, B. D. & C. Mitter. 1998. The timing of insect/plant diversification: might Tetraopes (Coleoptera: Cerambycidae) and Asclepias (Asclepiadaceae) have co-evolved? Biological Journal of the Linnean Society 63: 553–577 [pdf].

Linsley, E.G. 1961. The Cerambycidae of North America. Part 1. Introduction. University of California Publications in Entomology 18:1–97, 35 plates.

Copyright © Ted C. MacRae 2013

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

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

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

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

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''

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