Diversity in Tiger Beetle Larval Burrows

One of the fascinating aspects of tiger beetle study is their often high degree of fidelity for specific habitats.  Some species prefer wet habitats, while others frequent the drier uplands.  Some like sand while others need clay.  Differences in salinity, vegetational cover, and even slope dictate what species might be expected to occur in a given habitat, thus, the diversity of tiger beetle species one encounters is directly proportional to the diversity of habitats explored.  Unfortunately, tiger beetles can be rather ephemeral in their occurrence as adults.  Despite a life cycle that requires at least one year (and may take 2-3 years or even more), adults are often present for only for a few short weeks.  Even during the time that adults are present, they often hide if conditions aren’t right (too cold, too hot, too wet, too early, too late, etc., etc.  Add to that their marvelous evasive capabilities, and it’s a wonder I ever see or catch any at all!).  The study of tiger beetles is not, however, entirely dependent upon the adults.  The presence of larval burrows in an area is also useful information, and through understanding of the species that might occur in an area and their habitat preferences, it is possible to identify – at least tentatively – the species that might be living in them.

Cicindela lengi? (sandy tiger beetle) - Sioux Co., Nebraska

To the uninitiated, tiger beetle burrows might seem nothing more than a simple hole in the ground – anything could have made it.  However, with experience one becomes able to distinguish tiger beetle larval burrows almost instantly from burrows made by other ground-burrowing organisms.  The most common type of burrow is recognized by a combination of characters – almost perfectly circular except for a slight flattening on one side that gives the burrow a faint D-shape, and with the edge smoothly beveled.  This is your classic tiger beetle burrow and, for most U.S. species of Cicindela and related genera, averages ~5-6mm in diameter for 3rd instar larvae (tiger beetle burrows are most often observed at 3rd instar, since it is this final instar in which the larva spends the majority of its time and the burrow becomes most noticable).  The above burrow is one such burrow, found at Monroe Canyon in northwestern Nebraska last September.  While a number of species are known from the area, there are only a few that make their burrows in deep dry sands such as those that occur at this site.  We can eliminate Cicindela formosa (big sand tiger beetle) for reasons discussed below, and we can also dismiss Cicindela limbata (sand blow tiger beetle) because the habitat is not the barren, wind-shaped sand blow habitat that the species prefers.  This leaves two possibilities – Cicindela scutellaris (festive tiger beetle), a common and widespread inhabitant of sand habitats throughout the Great Plains, and Cicindela lengi (sandy tiger beetle), a much more localized resident of sand habitats with more western distribution.  The burrow likely represents the latter, since adults of this species have been found with greater frequency than C. scutellaris on the very fine-grained sands that occur in this part of Nebraska.  My confidence in this ID is bolstered by the fact that a larva I collected in the area from just such a burrow successfully finished its development and emerged a few months later as an adult C. lengi.

Cicindela pulchra pulchra (beautiful tiger beetle) - Fall River Co., South Dakota

Sometimes size alone is enough to indicate the species responsible for a burrow.  The above burrow was encountered last September in southwestern South Dakota on a clay/shale embankment in sage/shortgrass prairie.  A number of tiger beetle species fond of clay were observed at the site, including the two generalist species Cicindela tranquebarica (oblique-lined tiger beetle) and Cicindela purpurea audubonii (Audubon’s tiger beetle).  However, at ~8 mm in diameter the burrow is too large to have been made by either of these species.  The only tiger beetle in the area capable of making a burrow this size is Cicindela pulchra (beautiful tiger beetle), and in fact this burrow was found at one of several sites recently discovered by Matt Brust for this species in South Dakota.  Note again the classic shape – slightly flattened along the bottom side (the flattening accommodates the mandibles of the larval head – tiger beetle larvae always orient themselves in one position when sitting at the burrow entrance).

Cylindera celeripes (swift tiger beetle) - Woodward Co., Oklahoma

Just as large size was diagnostic for the previous burrow, the small size of the above burrow was also diagnostic.  This burrow, found at Alabaster Caverns in northwestern Oklahoma in October, 2009, measured only 3-4mm in diameter and can only have been made by Cylindera celeripes (swift tiger beetle).  This provisional ID was suggested by the fact that adults of the species had been observed abundantly in the lichen-encrusted clay exposures of this shortgrass prairie the previous June.  This photo, in fact, represents the first-ever discovery of the larval burrow of this species, and the identity of the species was confirmed when the larva collected from this and neighboring burrows and placed in rearing containers in the lab later emerged as adults.  I have found very similar-sized burrows in bottomland forest habitats in southeastern Missouri where the closely related species Cylindera cursitans has been seen.  The burrows are identical in size and shape, but the drastic difference in habitat is enough to distinguish the species that made them.

Cicindela formosa formosa (big sand tiger beetle) - Sioux Co., Nebraska

Not all tiger beetles utilize the simple hole-in-the-ground style of burrow, but rather incorporate some rather unique engineering features that make specific identification much easier.  This burrow can only be made by Cicindela formosa (big sand tiger beetle), a common resident of a variety of dry sand habitats throughout the Great Plains and eastern U.S.  The burrow entrance is on the large size for U.S. Cicindela (~6mm in diameter), and rather than opening flush on the ground it is directed horizontally and opens into a pit that is excavated to one side and underneath the burrow entrance.  No other U.S. tiger beetle makes a burrow quite like this (although I have noted Cicindela limbalis (common claybank tiger beetle) burrows on steep clay banks with a similar but much less distinct excavation on their lower side).  The pit apparently functions as a trap for potential prey, and since I have most often encountered burrows of this species in areas with some slope, I suspect the pit may help the larva capture its prey by preventing the prey from tumbling down the slope at the first sign of trouble.

Cicindela formosa 3rd instar larvae - Sioux Co., Nebraska

This is a different burrow by the same species, also at Monroe Canyon last September, that shows a 3rd instar larva sitting at the burrow entrance.  The sickle-shaped mandibles are resting against the slightly flattened lower edge of the burrow entrance, while the round pronotum fills the rest of the entrance profile.  The upper pair of eyes can be seen above the mandibles, but the lower pair (between the upper pair and the mandibles) are not visible in this photo due to the downward-facing angle of the burrow entrance.  I waited for quite some time with camera in position in hopes that I could photograph the larva, and when it did return to the burrow entrance I had time enough to fire off just a couple of shots before it retreated once again to safety in the depths of its burrow.

Cicindela fulgida fulgida (crimson salt flat tiger beetle) - Sioux Co., Nebraska

This unusual-looking burrow was found in a dry clay saline creek bed in the Badlands of northwestern Nebraska last September.  The turret structure is unique, but the nearly perfectly round and smoothly beveled burrow entrance identify it, nevertheless, as that of a tiger beetle larva.  These burrows can only be made by Cicindela fulgida (crimson salt flat tiger beetle).  There are several other saline-tolerant tiger beetles species in Nebraska, but most such as Ellipsoptera nevadica knausii (Knaus’ tiger beetle), Eunota togata (cloaked tiger beetle), and Habroscelimorpha circumpicta johnsonii (Johnson’s tiger beetle) require much more moisture than was found in this bone-dry creek bad.  I’ve found two other much more widely distributed clay-associated species – Cicindela tranquebarica and Cicindela purpurea audubonii – at this and other sites where I’ve seen C. fulgida; however, the larvae of those species do not utilize this unique turret-shaped structure for their burrows.  The turret is thought to have a cooling function for the larva during the heat of summer by raising it above the hottest layer of air against the white salt-encrusted ground and by aiding in the dissipation of heat from the larval burrow.  I wanted to photograph the larva sitting at the burrow entrance and spent quite a bit of time stalking out this and nearby burrows for a chance to do so.  Alas, however, on this day the larvae had greater patience than I!

Cicindela tranquebarica kirbyi (Kirby's tiger beetle) adult & larval burrows - Sioux Co., Nebraska

The above burrow entrances were photographed in September 2008 at the same dry saline creek bed in Sioux Co., Nebraska.  I mentioned above that Cicindela tranquebarica kirbyi and Cicindela purpurea audubonii both occurred commonly at this site along with Cicindela fulgida; however, these burrows likely represent the former.  That species seems to be found more consistently in high saline environments than the latter, which in this case probably have their larval burrows in the more normal clay soil further away from the creek bed.  During that 2008 trip, I did collect larvae from burrows like these in several similar, high saline habitats in Nebraska, South Dakota, and Oklahoma, and in each case adults of C. tranquebarica kirbyi were what emerged.  I have also reared this species from larvae collected on clay banks and wet sand habitats – in all cases, the burrows are a tad larger than those I have seen for other species in the genus that I have reared, such as Cicindela limbalis and Cicindela repanda (common shore tiger beetle) – logical since adults of C. tranquebarica tend to be a little more robust than these other species (but smaller than Cicindela pulchra and Cicindela obsoleta vulturina (prairie tiger beetle)).  In the above photo, I believe the the upper-right burrow is that of a larva, while the the lower-left one is that of an emerged adult – note the not-perfectly-circular opening and more ragged edge to the burrow.  In fact, the latter burrow looks very much like the adult emergence burrow that I saw at this very location last September, in which the still unemerged adult was seen sitting!  Granular chunks of soil can be seen scattered about the latter burrow, but I believe these were actually tossed by the larva rather than the adult as a result of burrow excavation – the amount of soil an adult would need to remove to re-open its burrow for emergence would probably be far less than what can be seen in this photo.  I did not search the surrounding grasslands for larval burrows, but if I had done so, it is likely that I would have found similar burrows that belonged to the larvae of Cicindela purpurea audubonii – the only other tiger beetle that we have seen in this inhospitable place!

Copyright © Ted C. MacRae 2011

Flash solutions for the beautiful tiger beetle

Canon MP-E 65mm f/2.8 1-5X macro lens (f/13), Canon MT-24EX flash w/ concave diffuser.

Recently I’ve been trying some different lighting and flash diffusion techniques with the Cicindela pulchra adults I brought back from South Dakota (see previous post).  While the beetles themselves are certainly among the most spectacular tiger beetles I’ve ever seen, I’ve been less than impressed with the photographs that I’ve managed to take of them.  Two factors have been largely responsible for this: 1) the smooth, shiny integument of the beetle reflecting the flash to create strong specular highlights, and 2) the colors, though brilliant, are also dark and difficult to bring out without further exacerbating the specular highlights.  Normally, the Sto-Fen+Puffer diffuser combination that I use does a pretty good job at diffusing the flash, but it just can’t handle these beetles.  To deal with this problem, I finally got around to trying out the do-it-yourself concave diffuser that Kurt at Up Close with Nature has been using with stunning results (similar to the tracing paper diffuser used so famously by Alex Wild at Myrmecos).  Photo 1 above and 2-3 below were taken with this diffuser on my Canon MP-E 65mm f/2.8 1-5X Macro Lens, and I’m rather pleased with these initial attempts.  I do need to figure out a better way to attach the diffuser to my Canon MT-24EX Macro Twin Lite Flash.

Canon MP-E 65mm f/2.8 1-5X macro lens (f/13), Canon MT-24EX flash w/ concave diffuser.

Canon MP-E 65mm f/2.8 1-5X macro lens (f/13), Canon MT-24EX flash w/ concave diffuser.

I’d have to say the lighting with this diffuser represents a considerable improvement over the Sto-Fen+Puffer diffusers using the same lens.  Compare especially Photo 3 above and 4 below – both taken with the MP-E 65mm lens at 1:1 and f/13 – Photo 3 was taken using the concave diffuser, while Photo 4 used the Sto-Fen+Puffer diffusers.

Canon MP-E 65mm f/2.8 1-5X macro lens (f/13), Canon MT-24EX flash w/ Sto-Fen+Puffer diffusers.

The problem with the concave diffuser is that it won’t work so well on my Canon EF 100mm f/2.8 Macro USM Lens due to its longer working distance.  I actually use this lens in the field as often as the MP-E 65mm lens, especially for the tiger beetles on which I focus (heh!) – not only do they rarely require more than 1:1 magnification, but they also rarely allow the ultra-close approach needed to use the 65mm lens.  The only solution is to find some way to get the flash heads closer to the subject to increase the apparent size of the light source, but so far I haven’t figured out a satisfactory way to do this.  Some photographers use the stalwart Canon Speedlite 580EX II Flash, equipped either with a do-it-yourself snoot diffuser or mounted on a bracket extender with a small softbox.  However, I am not a photographer, but rather an entomologist with a camera – I need to carry with me a net, vials, and in some cases a beating sheet and hatchet.  Both of the previously mentioned approaches for diffusing 100mm shots add far more bulk to the camera setup than I can accept.  I’ve been looking for extender brackets that will move the MT-24EX flash heads out closer to the subject to increase apparent light size and have found a few.  The Really Right Stuff B85-B Flash Bracket equipped with two FA-EX1 Flash Extenders and an extra Flash Mount looks like it would do the job quite well, but it is still bulkier (and vastly more expensive) than I would like.  The PhotoMed R2-C Dual Point Flash Bracket is a much less bulky and more reasonably priced option; however, the lack of any vertical adjustment capabilities is an insurmountable shortcoming.  Why Canon hasn’t themselves designed a lightweight, low-cost accessory for extending the MT-24EX flash heads out away from the lens is beyond me, and I’ve actually been toying with some ideas on how to do this myself using a couple of Kaiser Adjustable Flash Shoes.

Until I do figure out a solution, at least there is always the white box for any captive-held individuals (and yes, I have considered a small, collapsible white box to bring into the field – I’m not ready to resort to that just yet!):

Canon 100mm macro lens (f/16), Canon MT-24EX flash indirect in white box.

Photo Details: Canon 50D (ISO 100, 1/250 sec). Typical post-processing (levels, minor cropping, unsharp mask).

Copyright Ted C. MacRae 2010

North America’s most beautiful tiger beetle

Cicindela pulchra pulchra (beautiful tiger beetle) - Fall River Co., South Dakota.

Five years ago this month, I got my first glimpse of North America’s most beautiful tiger – Cicindela pulchra.  This is not just my opinion – its name, given to it by Father-of-North-American-entomology Thomas Say, literally translates to “beautiful tiger beetle.”  Like Amblycheila cylindriformis, it was a species that I longed to see in the field ever since receiving a single specimen from tiger beetle guru Ron Huber.  That specimen came from the Gypsum Hills of Barber Co. in south-central Kansas – a known “hot spot” for the species.  For years I stared at that spectacular specimen as it sat in my cabinet, and in September 2004 Chris Brown and I finally made our first attempt to see it in the field for ourselves.  Unfortunately, we arrived ahead of the fall rains that seem to trigger emergence of this species, and C. pulchra would not be among the few species that we saw on that trip.  I don’t handle defeat very well, so the very next fall I resolved to try again – this time waiting until early October and also enlisting the assistance of local entomologist “Beetle Bill” Smith for access to better sites than what are available along the roadsides.  That trip was a tremendous success and was detailed in one of my Nature Notes articles (MacRae 2006), but Chris, unfortunately, was unable to join me on that second attempt.  He couldn’t join me last year, either, for my search of the species in the nearby Cimarron Gypsum Hills of northwestern Oklahoma.  Good thing, however, as a turn of the weather left me just cold and wet (although I do remain convinced that the species will eventually be found on those red clay slopes that have so far produced such prizes as Cylindera celeripes, Dromochorus pruinina, and Amblycheila cylindriformis).

Matt Brust (L) and Chris Brown (R). Matt discovered this site for Cicindela pulchra in 2009.

Fortunately, while I was getting skunked in Oklahoma, Matt Brust was discovering new populations of the species further north in the southwestern corner of South Dakota.  These discoveries were prompted by the initial discovery of the species on Pierre Shale exposures at a single site near the Black Hills (Larsen and Willis 2008).  The soft, dark gray soils of the Pierre Shales are in distinct contrast to red clay exposures with which the species has been typically associated further south, and by scouting a broader area for similar exposures Matt was able to find the species at six new sites during late summer 2009 (Brust 2010).  He found them associated not only with the Pierre Shale but also the Mowry Shale formation (and suspects they may eventually be found on Belle Fourche Shale formations as well).  When I learned of these discoveries, I decided I just had to see them for myself.  I had enjoyed my Fall 2008 trip to northwestern Nebraska and southwestern South Dakota, and the thought of seeing these beetles while spending time in the field with Matt once again seemed the perfect basis for another trip to the area.  It didn’t take much convincing for Chris to agree, thus, C. pulchra became goal #1 of the 2010 Fall Tiger Beetle Trip™.

Habitat for Cicindela pulchra in Fall River Co., South Dakota. Adults and larval burrows are found in sparsely vegetated gray shale slopes and open flats beneath.

As we drove to the site that Matt had selected for us to explore, I felt nothing but optimism.  The skies were clear and the temperature was already nearing 70°F.  Matt, however, was hedging his bets – “I hope they’re still out, I’ve never seen them this late!”  Still, I wasn’t worried.  We were two weeks earlier in the season than the 2005 Kansas trip, and the weather was simply spectacular – it just had to be a good tiger beetle day!  My optimism was justified, as within minutes of arriving at the site we saw the first individual.  I collected this one alive as a backup for photographs in the studio should that be the last one we saw, but no such contingencies were necessary – we began seeing individual after individual as we trolled across the barren gray slopes.

Adult male Cicindela pulchra taking in the morning sun.

These beetles are simply a marvel to see in the field.  Brilliant dark red with metallic green, blue, and purple borders on the head, thorax, and elytra, this relatively large tiger beetle (certainly among the largest in the genus) can be confused with no other tiger beetle in North America.  Unlike adults of most other species, which exhibit color patterns resembling the texture and hue of the soil substrate on which they occur, C. pulchra adults are obvious and non-cryptic.  It apparently mimics the large, similarly colored velvet ants of the genus Dasymutilla with which they are sympatric – even exhibiting similar behavior when alarmed such as stridulating (creating vibrations by scraping body parts across one another) and giving off defense chemicals (Pearson 1988).  Adults are powerful fliers that can fly long distances when alarmed (Spomer et al. 2008), but in the still relatively cool morning air Chris and I had relatively (emphasis on relatively!) little trouble getting close enough to attempt those coveted field photographs.  This, however, was a double-edged sword – the same cool temperatures that allowed us to get close enough for photographs also caused to the beetles to assume the most non-photogenic poses as they sprawled torpidly on the ground, sometimes hugging it closely in an attempt to conserve heat until incident radiation from the sun warmed them sufficiently to go about the day’s activities.  Once this did happen, we found getting close enough for photographs nearly impossible.  In the 2+ hours that we chased after them, we took many shots but failed to get that “perfect” shot of a brilliant beetle standing tall and alert.

Adult male Cicindela pulchra hugs the ground during the cool morning hours.

Cicindela pulchra is a “spring/fall” species – i.e., sexually immature adults emerge during fall to feed, then return to their burrows to overwinter before emerging again in spring to mate and lay eggs.   Pearson et al. (2006) state the fall period lasts from July to September; however, as I observed in Kansas in 2005 adults can remain active well into October as long as suitable weather prevails.  Larvae hatch shortly after eggs are laid in the spring, but larval burrows can be seen during the entire season since they require 2-3 to complete development.  It was actually the presence of the large larval burrows (see photo below) that alerted Matt to the occurrence of the species at this site.  Several other tiger beetle species are also found here, e.g. C. purpurea (cow path tiger beetle) and C. tranquebarica (oblique-lined tiger beetle); however, these species – and hence their larval burrows – are considerably smaller than C. pulchra.  The only other species of Cicindela in North America that matches C. pulchra in size is C. obsoleta (large grassland tiger beetle), a southwestern species that is not known to range as far north as Nebraska and South Dakota, and the slightly smaller C. formosa (big sand tiger beetle) larval burrow has a distinctive “pitfall trap” with the burrow opening situated horizontally above it (it is also restricted to dry sand rather than clay habitats).  We saw several C. pulchra larval burrows during our visit but no active larvae, and none of my attempts to “fish” them out of their relatively shallow burrows met with success.  I could have tried digging them out, but that is a time-intensive activity, and I decided instead to bring a few live adults back in a terrarium of native soil and see if I could rear the species from egg.

Despite the presence of at least two other tiger beetle species at the site, this can only be that of a 3rd-instar Cicindela pulchra due to its large size.

I had tried persistently during the last hour we were there to get a good field photograph of an active adult beetle standing tall and alert, but the following is the closest I was able to achieve.  Leaving the site without that “perfect” shot was difficult – as Matt put it, we had “pulchra fever”!  Still, there were other tiger beetles – e.g. C. nebraskana (prairie long-lipped tiger beetle) and C. decemnotata (badlands tiger beetle) – that we wanted to find in the limited time we had to explore the region, so I prepared a terrarium for the live adults I was bringing back with me and chalked up our first big success of the trip as we headed towards the shortgrass prairie sitting atop the nearby Nebraska Pine Ridge.

I chased this adult female for some time trying to get a closer photograph, but warming temperatures made this impossible.

In addition to attempting to rear the species,  bringing live adults back with me also gave me more opportunity to photograph them.  In addition to the native crumbly shale soil that I used to fill the terrarium, I placed in it one of the nicely colored, presumably volcanic, rocks that littered the slopes on which the beetles occurred.  The dark color of the rock makes a nice backdrop to really show off the extraordinary colors of this species – especially the bright white labrum and mandibles of the particularly impressive male in the following photograph.  The beetles are all now sound asleep for the winter in a 10°C incubator.  Hopefully, when I move the terrarium back into warm temperatures next spring they will re-emerge, mate, and lay eggs (hmm, photographs of a mating pair would be really nice!).

The all-white labrum and mandibles of this male Cicindela pulchra are displayed nicely in this terrarium photograph.

Photo Details:
Insects: Canon 50D w/ 100mm macro lens (ISO 100, 1/250 sec, f/16), Canon MT-24EX flash w/ Sto-Fen + GFPuffer diffusers.
Habitat: Canon 50D w/ 17-85mm wide-angle lens (ISO 100, 1/160 sec, f/11), natural light.
Matt and Chris: Canon 50D w/ 17-85mm wide-angle lens (ISO 100, 1/250 sec, f/11), natural light.
All photos w/ typical post-processing (levels, minor cropping, unsharp mask).


Brust, M. L.  2010. New distribution records for Cicindela pulchra pulchra Say in South Dakota and notes on habitat use and natural history.  Cicindela 42:1–10.

Larsen, K. J. and H. L. Willis.  2008. Range extension into South Dakota for Cicindela pulchra (Coleoptera: Carabidae).  The Coleopterists Bulletin 62(4):480.

MacRae, T. C. 2006. Beetle bits: The “beautiful tiger beetle”. Nature Notes, Journal of the Webster Groves Nature Study Society 78(4):9–12.

Pearson, D. L.  1988. Biology of tiger beetles.  Annual Review of Entomology 33:123–147.

Pearson, D. L., C. B. Knisley and C. J. Kazilek. 2006. A Field Guide to the Tiger Beetles of the United States and Canada. Oxford University Press, New York, 227 pp.

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

Copyright © Ted C. MacRae 2010

Sweet Sixteen!

The 7th Annual Fall Tiger Beetle Trip™ is officially over – Chris and I rolled back into town a little after midnight last night. It was an amazing trip – perfect weather, unparalleled scenery, and a record-breaking 16 species of tiger beetles seen in 13 localities across four states. Not only does this beat my previous trip record of 13 species, but we did it with only five days in the field. At the time of my previous update, we had visited several locations in the South Dakota Badlands and Nebraska Pine Ridge and found ten different tiger beetle species, including Cicindela pulchra (beautiful tiger beetle) – our top priority for the trip – C. nebraskana (prairie long-lipped tiger beetle), and C. lengi (blowout tiger beetle). Our plan for the next day was to visit the Badlands of Wyoming to look for C. decemnotata (Badlands tiger beetle – appropriately) and the Yampa River Valley of northwestern Colorado to look for C. scutellaris yampae and C. formosa gibsoni, all three of which we managed to find (though with caveats – stay tuned). Our originally planned final field day was to take us back into Wyoming to look for C. longilabris (boreal long-lipped tiger beetle) in the mountains east of Laramie and the Nebraska Sand Hills to look for the delicate little C. limbata before heading back home. However, we were finally paid a visit by “the skunk” and did not see any of these species (although our sighting of C. limbata (common claybank tiger beetle) in Wyoming did officially break the old trip record). Not wanting to end the trip on a disappointing day, we delayed our departure for home yesterday and visited two more sites at the eastern edge of the Nebraska Sand Hills (sites M and N in the above map) – a clay bank site where we saw a robust population of C. denverensis (to augment the single individual we had seen earlier in the trip) and several C. splendida (splendid tiger beetle), and another sand dune/blowout system where we at last succeeded in finding C. limbata.

The day after the end of the Annual Fall Tiger Beetle Trip™ is usually a somewhat depressing day for me. Not only is the trip over, but likely so is the entire insect collecting season. I know I need the down time to process the specimens and knowledge acquired during the season, but the field work itself remains my favorite aspect of this pursuit. Nevertheless, the experiences from this trip will fuel my memories for years to come, and in the next weeks I’ll share some of the stories that unfolded. Until then, I leave you with this portrait of C. pulchra – looking rather annoyed with me for my persistent efforts to take his photograph.

Cicindela pulchra - the ''beautiful'' tiger beetle

Copyright © Ted C. MacRae 2010

2 days, 6 localities, 10 species…

Here’s an updated itinerary for the 7th Annual Fall Tiger Beetle Trip that fellow cicindelophile Chris Brown and I are in the midst of. We’ve spent the past two days visiting six localities in Nebraska and South Dakota. So far, we’ve found a total of 10 species – including every species we had hoped to see at this point in the trip. The list so far (in chronological order) is:

  • Cicindela (s. str.) tranquebarica kirbyi – ho hum, we’ll see this in several places.
  • Cicindela (s. str.) purpurea audubonii – über common Great Plains species, although the black form is always a treat to see.
  • Cicindela (s. str.) pulchra – YEAH! Seen in good numbers at one of the new South Dakota localities discovered in 2008 by Matt Brust (our personal chaperone for the day). Marvelous field photographs.
  • Cicindela (s. str.) fulgida – Only one seen, but Chris got a nice series of field photographs (I’ve seen good numbers of this species from my previous trips to this area in 2008 and in Oklahoma last year).
  • Cicindela (s. str.) nebraskana – Another “A list” species for the trip, but we’ve only seen one so far.
  • Cicindela (Cicindelidia) punctulata punctulata – also known as Cicindela ubiquita.
  • Cicindela (s. str.) scutellaris scutellaris – even though this is a common Great Plains species in any sandy area, I never tire of its dazzling red elytra and blue/green head and pronotum.
  • Cicindela (s. str.) lengi – The third species on our “A list” that we’ve seen, with some real nice field photographs from Monroe Canyon.
  • Cicindela (s. str.) formosa generosa – another common Great Plains species.
  • Cicindela (s. str.) denverensis – I didn’t expect to see this one on the trip (just a single individual at Monroe Canyon), but I’ll take it!

Tomorrow we’ll hit a Wyoming location where Cicindela (s. str.) decemnotata is known to hang out – a species I’ve not yet seen, either alive or preserved. Most sources regard this species as closely related to C. denverensis, but Matt thinks it is actually more closely related to C. fulgida due to similarity in form and shine but green instead of purple. Afterwards, in a major addition to our planned itinerary (hence the updated Google Map), we’ll go into northwestern Colorado to look for two very cool subspecies of the otherwise widespread species – C. formosa gibsoni and C. scutellaris yampae. If we’re lucky we’ll also see the delicate little sand lover, Cicindela (s. str.) limbata, but if we don’t see it there then we should see it the next day when we finish out the trip back in the Nebraska Sand Hills just east of Alliance. But before that, we’ll veer back up into Wyoming and look around in the high elevations east of Laramie in hopes of finding Cicindela (s. str.) longilabris laurentii. That one may be a stretch, but if we are successful then we have the potential to see a total of 15 species – that would be a trip high for me (literally and figuratively).

Copyright © Ted C. MacRae 2010

Megacyllene comanchei revisited

On my recent trip, I reported finding the rarely collected Megacyllene comanchei at several localities in northwestern Nebraska and southwestern South Dakota. These are significant findings, since they not only represent new records for both states, but also an impressive 700-mile northern extension to the known range of the species (on top of a previously reported northern range extension from Texas into Kansas). The intriguing part of the situation is that these new records fall within the southern portion of the known distribution of M. angulifera, its closest relative, which has been recorded from several northern Great Plains states and provinces (although it has not yet been recorded specifically from Nebraska).

Upon reading about these findings, a friend and fellow student of Cerambycidae has expressed doubts to me about the distinctiveness of M. comanchei versus M. angulifera, regarding the slight color differences upon which it was based as insufficiently distinctive. In its original description (Rice & Morris, 1992, J. Kans. Entomol. Soc. 65:200-202), M. comanchei was distinguished from M. angulifera by a specific combination of characters, i.e., the premedian and sutural segment of the postmedian elytral bands are white, while the remaining pubescent bands are yellow. They also noted the subapical and apical bands often coalesce along the elytral suture and lateral margins. I do not have material of M. angulifera for direct comparison, but my specimens (two of which are pictured in this post) seem distinct enough from this specimen of M. angulifera pictured on Larry Bezark’s impressive website, A Photographic Catalogue of the CERAMBYCIDAE of the New World. In contrast, the pattern of coloration seen in this photo of the holotype of M. comanchei (also from that fine site) seems to agree well with my material.

Whether these color differences are significant remains to be seen. Neither species has been commonly collected, therefore large series of material have not been available for good comparative studies. However, there do seem to be significant differences in reported host plants and adult biology. Adults of M. angulifera are usually found in the fall on flowers of Solidago (goldenrod), and it was recently discovered that the larvae utilize root crowns of Dalea candida (family Fabaceae) for development (Blodgett et al. 2005). All but one of the remaining Nearctic species of Megacyllene for whom larval hosts are known also utilize fabaceous plants. These include M. robiniae and M. snowi (Robinia), M. decora (Amorpha), and M. antennata (Prosopis). A notable exception is M. caryae, which breeds in a variety of deciduous plant genera but most often Carya and is also unique in that adults occur during early spring instead of fall. Megacyllene comanchei, on the other hand, was discovered by examining the dead root crowns of Heterotheca sp. (family Asteraceae), with a few crawling on the ground in areas where such plants were growing. This is not proof that it serves as a larval host, but the repeated association of adults at the base of dead stems of this plant is highly suggestive. Additionally, no adults were encountered on goldenrod or other flowers, as is common among other members of this genus. The apparent utilization of a non-fabaceous larval host and behavioral difference exhibited by the adults seem to support its status as separate species. Again, the specimens that I collected agree not only morphologically with M. comanchei, but also behaviorally in that all of them (six specimens at three localities) were found crawling on the ground in shortgrass prairie rather than on flowers. I did note Heterotheca growing at one of the locations but did not find adults on the crowns of the few plants I inspected. I do not recall seeing any Dalea, but I wasn’t looking specifically for that plant either. Nor did I not see any goldenrod, which I would have certainly noticed had it been present.

So, for now, I’m inclined to continue calling these M. comanchei, and I’m also inclined to consider it distinct from M. angulifera. I do agree, however, that a critical examination of the distinctiveness of these two species might be warranted, and it may be worthwhile to pull together as much of the existing material of these two species as possible. I am interested in hearing other opinions about this situation.

Rattled in the Black Hills

My first day in the Black Hills of South Dakota was spent at McNenny State Fish Hatchery near Spearfish – on the north side of the Black Hills. I went to this place on the advice of my esteemed colleagues in Nebraska, who suggested that I might be able to find several interesting tiger beetle species there: the closely related trio of beauties C. denverensis (green claybank tiger beetle), C. limbalis (common claybank tiger beetle), and C. splendida (splendid tiger beetle) in the red clay eroded banks; C. fulgida (crimson saltflat tiger beetle) and C. tranquebarica kirbyi (oblique-lined tiger beetle) around the lakes; and – again, if I’m lucky – intergrades between the prairie and boreal long-lipped tiger beetles (C. longilabris x nebraskana) along a trail through the shortgrass prairie east of the hatchery. For the first time since Saturday, I awoke to baby blue skies which filled me with an optimism and anticipation that made the 3-hour drive from Chadron, Nebraska to Spearfish, South Dakota seem interminable.

What my esteemed colleagues failed to include on that list of species I might encounter was Crotalus viridis – the prairie rattlesnake! Now folks, I’ve seen a number of rattlesnakes before – mostly in Texas – but I’ve never heard this sound in real life, much less heard it coming from a rattlesnake poised to strike. I encountered this fellow in the eroded red clay slopes above the lake, and even though I wasn’t too terribly close it gave me quite a start (my bravery in taking this photo is vastly exaggerated by the twin miracles of telephoto and cropping!). I walked a little more cautiously afterwards but gradually let my guard down over time. About an hour later, I was startled again by another rattler – I had come within 2 feet of it before it started rattling. I nearly jumped out of my skin, and once I got my heart stuffed back down my throat I noticed several dark juveniles coiled up with her. They slunk away, and I tiptoed back to the car having had my fill of the red clay slopes for the time being.

I did manage some success on the slopes before the rattlers drove me away – not with the claybank and splendid tigers that I had hoped to find, which were largely missing in action save for two individuals of C. limbalis that I spotted amongst the annoyingly similar appearing and ridiculously numerous C. purpurea audubonii (clay path tiger beetle). Success instead came in the form of this cerambycid beetle – Megacyllene comanchei. Recently described from Texas, nothing more was published about this species until I recorded a northern range extension into south-central Kansas (MacRae & Rice 2007). Its occurrence in the Black Hills is not only a new state record for South Dakota but also represents an incredible 700-mile northern range extension – on top of the previous one! Actually, Matt and I each found one individual a few days ago in Sioux County, Nebraska (also a new state record) – I had thought of this species at the time but decided I must be wrong and that I should wait until I got back before making an identification. But the capture of these three additional individuals even further north renewed my suspicions, and consultion of my databases shows good agreement with this species – note the white rather than yellow antemedian elytral band and medial portion of the postmedian elytral band, along with the medial and lateral coelescence of the apical and subapical bands, which distinguish this species from the closely related M. angulifera. The records from this trip show that M. comanchei is much more widely distributed than previously thought. Curiously, all five of the individuals I’ve seen (so far!) were crawling on the ground – an unusual habit for Megacyllene, which are normally found on flowers of goldenrod. The type series was associated with plants in the genus Heterotheca, which I did note growing in the area.

After escaping the snake slopes, I began surveying the lake margins to look for potential tiger beetle habitat. I was especially interested in C. fulgida – Matt and I had seen a single individual along a dry salt creek in Sioux County. The lakeshore around the upper lake was completely surrounded by thick vegetation – no tiger beetles there, but when I arrived at the lower lake I found some small areas of open ground along one side. They didn’t look very extensive, and my initial search of the area showed no activity. Closer inspection, however, showed the presence of larval burrows, and when I grabbed my fishing gear (the nearest grass stem) I promptly managed to extract a couple of larvae. Okay, so there are tiger beetles here, but which one I don’t know – probably C. tranquebarica kirbyi, which we had seen rather commonly at the same dry salt creek in Sioux County. Although the sun would not set for another two hours, it was quite cool already. I wondered if maybe the adults had already started digging in for the night and began looking for evidence of adult burrows. I looked carefully along the edge of the grassline when I saw movement – it was the back end of an adult C. fulgida kicking dirt out as it excavated its burrow. Success! I dug it out, took a few photos (one shown here) and started looking for similar appearing burrows. I not only found several more C. fulgida in their burrows, but also several C. tranquebarica kirbyi. The larvae I collected may or may not represent one of these species – there are other species associated with alkaline habitats that active at other times during the season. I collected a few more larvae, filled a container with soil from the spot – cutting out a section of salt-encrusted surface to place on top, placed all of the collected larvae in it, and watched them immediately start digging new burrows with their shovel-like heads. More babies to take care of!

With tiger beetle success under my belt and the sun setting fast, I decided the day was done and packed up the car. As I was closing the hatch, I happened to look over and saw something of great interest – milkweed! I had, in fact, been looking for milkweed all day long in the hopes – faint, I thought – of encountering the newly described Tetraopes heutheri (Skillman 2007). Mirror Lakes, at the McNenny Fish Hatchery, is the type locality of this species, and although the type series was collected in August I held out hope that the adults might persist until September. These hopes faded quickly, however, as I located milkweed plant after milkweed plant on the shortgrass prairie above the eroded clay slopes – all completely senesced, with nary a sign of any milkweed beetles. The plant I’d just spotted – only a small sprout – was green, and on it were two milkweed beetles! I excitedly took some pictures, then started looking carefully about and found several more on additional small sprouts in the area. Apparently, the sprouts represented regrowth from late-season mowing of the roadside, as several full-sized, completely senescent plants were found in the adjacent unmowed area. My excitement at having “found” T. heutheri (because of their small size and occurrence at the type locality) was short lived – closer examination of the specimens after returning home showed them to be very small individuals of the more common Great Plains species T. femoratus. I did have some doubts when I found the beetles, since the milkweed species on which I found them is not the same species with which T. heutheri was associated (Asclepias verticillata, a small species with narrow, linear leaves).

Day 2 in the Black Hills was spent at nearby Boundary Gulch, just across the border in the northeast corner of Wyoming. This was another attempt to find the C. longilabris x nebraskana intergrades that eluded me at McNenney, and although I failed to find them at this location also, I did find five other species of tiger beetles, including several beautifully marked C. limbalis to go along with the two I found the previous day. After that it was some spurious collecting here and there – including larvae from two spots in the southern Black Hills – as I traveled back to Chadron, Nebraska for the night. On tap for tomorrow – Nebraska’s famed Sand Hills! The beautiful sandy tiger beetle (C. limbata) – vivid white and iridescent green to red – hopefully will be found among the super abundant festive (C. scutellaris) and big sand (C. formosa) tigers, and I’ll get another shot at seeing the C. lengi (blowout tiger beetle) that I missed a few days ago.

A hunting we will go!

Maps have been prepared. Relevant emails from my esteemed colleagues to the northwest have been read and re-read. Summary sheets on the distribution, biology, and biogeography of the many different species I hope to encounter are in hand. Google Earth images of each locality I plan to visit – annotated with potential species occurrences and pinpointing precise locations of their likely habitats – have been assembled into a Powerpoint presentation, and detailed driving directions from Point “A” to Point “B”… all the way to Point “X” (home!) have been determined. All of this has been printed out and organized into a 3-ring binder. Why the extraordinary attention to detail? Because…

It’s time for the annual fall tiger beetle trip!

View Larger Map

The annual fall tiger beetle trip started several years ago when I, along with my friend and colleague Chris, began studying Missouri’s tiger beetle fauna. At first it was a diversion – buprestids and cerambycids are pretty well played out by fall, but tiger beetles across much of the U.S. exhibit a unique spring/fall fauna that is quite distinct from the summer fauna. Chris and I would go to different parts of Missouri, documenting the species encountered to fill in distributional data gaps. It was on these trips that I discovered how much I truly love early fall collecting – the cool air, the crisp smells, the long sharp shadows, and a landscape of foliage ever so lightly tinged with shades of red and yellow while grasses morph into fields of gold. In recent years, I’ve begun adventuring beyond Missouri’s borders on these fall trips, allured by the diversity of species found in the Great Plains – species alien to Missouri in an equally alien landscape. First, it was Barber County, Kansas, with its red gypsum hills inhabited by the aptly named Cicindela pulchra (beautiful tiger beetle) – deep wine-red and iridescent purple flashing across the barren red clay. Then last year I got my first taste of the Sand Hills of Nebraska at their farthest eastern extent. I watched in amazement as Cicindela limbata (sandy tiger beetle) – vivid white and metallic green – danced across the surface of sand blows, undaunted by scouring 30 mph winds. It was on that trip that I decided a long weekend wasn’t cutting it – I needed to take a whole week and get myself into the heart of the Great Plains. The annual fall tiger beetle weekend has just become the annual fall tiger beetle week.

As the map above indicates, I’ve got a rather ambitious itinerary of locations that I’d like to visit – 22 in all. I leave tomorrow, and if I have planned properly (and have a little luck) I might be able to visit all of them in the 9 days I have set aside for the trip. My “trip bible” will be my constant companion, along with my already worn copy of the newly issued Tiger Beetles of South Dakota & Nebraska (Spomer et al. 2008), as I explore deep into the Sand Hills and experience for the first time ever the Black Hills of South Dakota. I’ll even sneak over into Colorado and Wyoming for a spot or two. Unfortunately, my faithful colleague isn’t able to join me. I tried to seduce him with visions of Cicindela limbata and C. lengi (blowout tiger beetle) in the numerous sand blows, C. fulgida (crimson saltflat tiger beetle) around countless alkaline lakes, C. longilabris (Boreal long-lipped tiger beetle) in the high pine forests, and C. nebraskana (prairie long-lipped tiger beetle) and (if we’re really really lucky) C. decemnotata (Badlands tiger beetle) just sneaking into the shortgrass prairies of the extreme northwestern corner of Nebraska. I reminded him of my (wanting) photographic skills and the images we would have to settle for if his talent and equipment didn’t accompany me. I almost had him, but in the end he muttered some lame excuse about his 15-month old baby and wife needing him (just kidding, Chris!).

The map above should be fully interactive, so give it a click and follow me along on this adventure. If you happen to be at any of the spots marked by a balloon and see a khaki-clad fellow – insect net in one hand, camera in the other – how’s about joining me for a bit of tiger beetle hunting.