tiger beetles

North America’s largest tiger beetle (I think!)

/ Ted C. MacRae / 12 Comments

The generous among us might call it serendipity, while the rest of us would just call it luck.  By whatever name, I had it in spades in June last year when I made my first visit to the Glass Mountains of northwestern Oklahoma.  My original plan was to go to Tallgrass Prairie Preserve in eastern Oklahoma at the end of what proved to be a resoundingly successful day at Alabaster Caverns State Park.  However, soaking rains moved into the area and continued rumbling eastward across the plains towards the preserve, forcing a quick change of itinerary.  I decided to wait it out in the state’s western reaches, a “hunch” telling me that the red clay/gypsum hills in nearby Major County might be a fruitful place for hunting tiger beetles.  Sunny skies the next morning were a good sign, and my hunch was rewarded later that day when I discovered a previously unknown (and fortunately robust) population of the rare Cylindera celeripes (swift tiger beetle), making not just the day but the entire trip more successful than I could have ever imagined.  Icing on the cake came when I found decent numbers of the more secure but nevertheless uncommon Dromochorus pruinina (frosted dromo tiger beetle) also in the area.

Another find I made that day that I’ve mentioned on occasion but not talked about at length was a single, rather large tiger beetle larva.  I found several burrows at the base of a talus slope at the edge of a small ravine where many adult C. celeripes were scurrying, and though I tried with many, out of only one did I manage to “fish” its occupant with a blade of grass.  I didn’t know which species it represented, but its large size and occurrence in clay soil brought to mind two species – Cicindela pulchra (beautiful tiger beetle) and C. obsoleta (large grassland tiger beetle).  Both of these Great Plains species reach their eastern limit of distribution in this part of Oklahoma (Pearson et al. 2006), and their status as the largest species of the genus (and its former subgenera) in North America seemed to make them the leading candidates for this enormous larva.  There was one other possibility – Amblycheila cylindriformis (Great Plains giant tiger beetle), another Great Plains species at its eastern limit in western Oklahoma and (as the common name implies) the largest tiger beetle in all of North America.  However, to consider that species seemed too much wishful thinking.  From my understanding, larvae of that elusive species reach an incredible 45 mm in length and dig burrows  on steep slopes or at the mouths of rodent burrows that extend vertically to depths of up to 1.5 m or more (Brust et al. 2005).  Surely I could not have so casually stumbled upon such a grand grub!

I placed the larva in a terrarium of native soil and brought it back with me, and for one year now I have waited – feeding it a regular diet of the fat noctuid caterpillars that we rear so abundantly in our lab.  For a full year, I’ve watched it nab caterpillar after caterpillar, disappearing mysteriously for days on end, and just as mysteriously reappearing at the top of its burrow.  I knew getting a closer look at it would help in my attempts to determine its identity, but every time I approached with a camera it dodged down into its burrow and beat my patience.  Sometimes I would see it sitting about a centimeter below the burrow entrance – just waiting for a caterpillar to crawl by but refusing to expose itself to the lens.  I gradually decided it was likely C. pulchra, as I had seen that species in similar habitat not too far north in Barber County, Kansas.  So strong was my suspicion that I even made another trip out to the Glass Mountains in October of last year, expecting to see the fall-active adults bejeweling the exposed flats below the red clay slopes, their wine-red elytra and purple-margined bodies all aglitter under the crisp, autumn sun.  No such sight was beheld, however – my hopes dashed by the season’s sudden cold and wet turn, and with the terrarium containing the larva by then tucked away in a cool incubator for a winter’s rest, it would be several months before I would see the larva once again sitting at the top of its burrow.

In late March I pulled the terrarium out of the incubator, and within a week the larva reopened its burrow.  I fed it a few times, and then one day I saw that it had dug a new, larger  burrow – measuring a full 10 mm in diameter!  This seemed extraordinarily large for any species of Cicindela, so I resolved once again to photograph it and determine its identity.  For days I stalked it, keeping the terrarium just outside my office door where I could keep an eye on it, yet every time I approached within two feet or so it would drop down out of sight.  I decided to stop feeding it – perhaps hunger would overwhelm its patience and prompt it to return to the top of its burrow more quickly after retreating.  That seemed to work, as one day the larva came back up after only a few minutes – and I was ready!  Already  in position, I flashed off multiple shots as soon as it reappeared, moving slowly and deliberately between shots to avoid spooking it again, and managed to get a nice series from varying distances.  As a testament to its enormous size, all of the photos shown here were taken with the standard 100mm macro lens (1X maximum) – not the 65mm 1-5X beast that I needed for these shots of the super-tiny C. celeripes.

Thinking that the larva likely represented C. pulchra, I compared the photos to this photo taken by Matt Brust of a 3rd-instar larva of C. pulchra and immediately noted the differently shaped pronotum of my larva and its distinctly projecting anterolateral angles.  Compare to C. pulchra, in which the angles are in line with the median part of the anterior margin – it is clearly not that species.  It isn’t C. obsoleta either, as that species has the anterolateral angles of the pronotum even less projecting than C. pulchra (Drew and Van Cleave 1962).  Apparently I needed to rethink my assumption that it belonged to Cicindela or its close relatives – none that occur in Oklahoma are simply large enough!  Tetracha virginica is large enough, but I knew it wasn’t that species since it lacked the white margined pronotum distinctive of species in that genus (as can be seen in this post on the larva of Tetracha florida).  That left only A. cylindriformis, distinguishable from all other tiger beetle genera occurring in Oklahoma by the second (lower) pair of eyes distinctly smaller than the first (Hamilton 1925, Drew and Van Cleave 1962, Pearson et al. 2006) – clearly seen in the third photo above.  Matt Brust has also photographed the larva of A. cylindriformis – it’s not a close shot of the head and pronotum, but in general aspect my larva seems to match it well enough.

All that is left is to actually succeed in rearing this larva to adulthood.  These beasts may require up to three to four years to develop (Brust et al. 2005), although this is likely influenced by latitude and prey abundance.  I suspect it was a second instar larva when I collected it, and that it dug its new burrow this spring after molting to the third (and final) instar.  Hopefully by keeping it in a nice, warm growth chamber and feeding it generously with fat caterpillars, I can minimize the time to pupation and perhaps see the adult sometime later this summer.  If/when that happens, you can be sure to see a follow up to this post.

Photo Details: Canon 50D (ISO 100, 1/250 sec, f/16-18), Canon 100mm macro lens, Canon MT-24EX flash (1/4 ratio) w/ Sto-Fen diffusers. Post-processing: contrast and unsharp mask (no cropping).

REFERENCES:

Brust, M. L., S. M. Spomer and W. W. Hoback.  2005. Tiger Beetles of Nebraska.  University of Nebraska at Kearney.  http://www.unk.edu (Version 5APR2005).

Drew, W. A. and H. W. Van Cleave.  1962. The tiger beetles of Oklahoma (Cicindelidae).  Proceedings of the Oklahoma Academy of Science 42:101–122.

Hamilton, C. C.  1925. Studies on the morphology, taxonomy, and ecology of the larvae of Holarctic tiger beetles (family Cicindelidae).  Proceedings of the U.S. National Museum 65 (Art. 17):1–87.

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.

Copyright © Ted C. MacRae 2010

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Swift Tiger Beetle – good news, bad news

/ Ted C. MacRae / 13 Comments

Sorry about the noisy video – it was shot in one of our walk-in growth chambers with fans going full-bore! Anyway, the video shows a couple of mating pairs of Cylindera celeripes (swift tiger beetle) that I brought back from the Loess Hills of extreme northwest Missouri this past weekend.  Watch carefully and you’ll see the first pair actively copulating before they break apart. After that I pan over to the second pair, which is not actively mating but have remained coupled as an example of behavior called “mate guarding” You might also notice a few very small green “bugs” in the container – these are 2nd-instar Lygus nymphs, which I placed in the container earlier in the day as prey – it was quite a sight to see the tiger beetles immediately begin chowing down on them! At the end of the video, I poke at the second mating pair with my forefinger to give an indication of their tiny size – this flightless species is one of the smallest in all of North America! Knowing how tiny the beetles are and how well they blend into their surroundings (you’ll have to imagine the 1-2 ft of plant growth that was surrounding them in the field), you can appreciate just how difficult these beetles are to detect in their native habitat.

There is a “good news, bad news” aspect to the story behind these beetles. This rare Great Plains species was unknown from Missouri until last year, when we (Chris Brown and I) discovered it in loess hilltop prairie remnants at Brickyard Hill, Star School Hill Prairie, and McCormack Loess Mounds Conservation Areas. Loess hilltop prairie is among Missouri’s most critically endangered natural community due to its restricted occurrence at the southern tip of the Loess Hills landform and more than a century of overgrazing and relentless encroachment by woody vegetation and invasive exotics. The sites where we found the beetle last year contain the highest quality loess hilltop prairie remnants in Missouri, so we are now taking a more thorough look at some of the smaller remnants that still exist in the area. The most promising of these are at Squaw Creek National Wildlife Refuge and on several privately owned lands near the known sites, and these were the sites that I searched this past weekend. Some of these sites looked promising (one in particular looked excellent), but thorough searching at each revealed no beetles. By the time I finished searching the last of them I had begun to wonder if: 1) my “search image” for the species had gotten rusty, or 2) my timing was a bit too early (last year’s populations were discovered in late June). To test this, we (daughter Madison and I) returned to one of the known sites (McCormack) where we had seen only two beetles last year (unable to capture either one). As we hiked along the ridge top leading to the spot where we saw them, I kept a close watch on the narrow trail in front of me. Nothing. However, as soon as I came upon “the spot” I saw one! I dropped to my knees and slapped my hands down on the ground, forming an “arena” between my two thumbs and forefingers, but the beetle ran over my hand too quickly and escaped. No matter – in less than a minute I saw another one and successfully trapped it under my fingers as it ran over my other hand. During the next 15-20 minutes I would see at least eight individuals along the narrow trail in this single spot. While it was gratifying to see more individuals at this site than we had seen last year, it also meant that the timing of my searches at the other sites was fine and that I was not suffering from a rusty search image. The beetle could still be at those sites where I had failed to find it, but if it is then it certainly does not occur in very high numbers. It also bothers me that at this site the beetle seems to be restricted to one isolated ridge, which appears not to have been burned in recent years (in contrast to the rest of the preserve, which seems to have been burned within the past year or so). I searched all the remaining ridge top again thoroughly after finding the beetles again this year, but no beetles were seen anywhere except this tiny spot where we have now seen beetles in successive years.

Missouri’s few existing loess hilltop prairie remnants are not only small but highly disjunct, and the flightless nature of the beetle makes re-colonization of a remnant unlikely in the event of a localized extirpation. There is obviously much we still do not know about the impact of burning on the beetle and how best to devise management plans that consider both the habitat and the beetle. However, one thing is clear – both the habitat and the beetle are critically imperiled in Missouri, and the fate of both are in our hands, right here and right now! We’d better get this figured out quick if we’re going to save both, and there seems to be little room for error. For my part, in addition to pinpointing where our populations occur and precisely what habitats are supporting them, I am trying to develop an effective rearing technique for this never-before-reared species in the event that captive rearing becomes necessary for reintroduction or augmentation of native populations. The adults seem very delicate and do not travel well, but I have found that if I prepare a terrarium in the field for transporting the adults then they survive well – even when traveling for several days. The container measures 6 1/4” H x 8” L, and I’ve placed a chunk of native soil cut from the site where I found the beetles and kept intact. The debris on the soil surface is intact as well, but the plants growing in the soil have (obviously) been trimmed. I’ll collect eggs from these individuals and experiment with different methods that I’ve been working on for rearing the larvae to see which are the most efficient and effective.

Copyright © Ted C. MacRae 2010

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Where’s Ted now?

/ Ted C. MacRae / 5 Comments

…in the Loess Hills of northwest Missouri, looking for additional sites for Cylindera celeripes (swift tiger beetle). Recall that we (Chris Brown and I) finally found this rarely collected species last year in Missouri (after many years of looking) in high quality remnants of loess hilltop prairie (a critically endangered natural community in Missouri).  The beetle was found at Brickyard Hill, Star School Hill Prairie, and McCormack Loess Mounds Conservation Areas, which combined contain nearly half of the 50 or so acres of loess hilltop prairie still existing in Missouri. The remaining acres are located at Squaw Creek National Wildlife Refuge and on private lands – my sincere appreciation goes out to Squaw Creek Refuge staff and three private landowners, who have all graciously extended to me (and my able field assistant Madison) access to the loess hilltop prairie remnants under their stewardship to better characterize the beetle’s distribution in the area. The sites I am visiting have varied burn histories, ranging from recent to 6 years or more since the previous burn, thus, I am also hoping to better understand the possible impact of prescribed burns on the species’ occurrence in loess hilltop prairie remnants. The beetle needs these remnants to survive, and prescribed burning is an important tool for helping to restore this natural community after decades of shrinkage due to woody encroachment. The trick will be to design management plans that accomplish these restoration objectives while at the same time minimizing possible negative impacts of the burns on existing beetle populations.

Copyright © Ted C. MacRae 2010

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Are we loving our prairies/glades/woodlands to death?

/ Ted C. MacRae / 25 Comments

I had such high hopes for last weekend’s collecting trip – late May is boom time for insects across Missouri, we have had good moisture this spring, and I would be visiting some high-quality natural communities that I had not visited for a long time.  My stated goals (the jewel beetles, Agrilus impexus and A. frosti) were long shots – I knew that and would have been fine coming home without those species (which I did) had the the collecting been otherwise productive (which it was not).  Still, I’ve been doing this for a long time now, and I’ve learned to draw on my accumulated experience when things don’t go as planned to give myself the best shot at turning a bad collecting trip into a decent one when things don’t go as planned.  The itinerary with which I start is rarely the one that I actually follow, and this past weekend was a good example of such.

My first stop was Ha Ha Tonka State Park, one of Missouri’s premier parks, boasting high-quality chert, dolomite, and sandstone savanna interspersed with dolomite glades.  It is on these glades and savannas that I hoped to find Agrilus impexus, or failing that at least collect a nice diversity of other jewel beetles on the oaks and hickories of the savannas and surrounding woodlands.  However, it was with some reservation that I even came here after being told by my colleague at the Department of Natural Resources just 2 days before my trip that 75% of the park’s grasslands and woodlands had been burned within the past two years.  For an insect collector, this is never good news – in all my years of collecting insects, my experience in relatively recently-burned habitats has been consistent: collecting sucks!  I decided, however, to visit Ha Ha Tonka anyway because of the quality of the natural communities it contains, thinking perhaps I might be able to find pockets of unburned habitat supporting good insect populations.  This was not to be. I beat oak after oak in the savannas and woodlands – nothing!  I swept little bluestem and Indian grass in the glades – nothing!  The foliage was lush and green and the savanna and glade landscapes highly diverse – given the time of season the place should have been teeming with insect life, yet it almost seemed sterile. Were it not for a few Chrysobothris quadriimpressa jewel beetle adults that I found attracted to a recently wind-thrown black oak tree, I would not have seen any insects here at all.  It appeared my fears about park-wide depression of insect populations had been realized.  However, not one to waste a visit I decided to explore some of Ha Ha Tonka’s fascinating geological features.  Ha Ha Tonka contains one of Missouri’s best examples of karst geology, with complex structures formed from the collapse of a major cave system.  The Devil’s Promenade is one of the more spectacular examples of such, its horseshoe-shaped cliff representing the former interior walls of a now-collapsed cave.  As dusk approached, the day’s poor insect collecting caused me to abandon my plans to stay here and blacklight for nocturnal beetles.  Instead, I decided to break from the itinerary, drive further west and explore Lichen Glade Natural Area in the morning before heading to the Penn-Sylvania Prairie BioBlitz later that afternoon.

Devils Promenade, Ha Ha Tonka State Park

Lichen Glade Natural Area is a small area owned by The Nature Conservancy that boasts a high-quality sandstone glade surrounded by post oak/black jack oak forest.  My first visit to the area more than 20 years ago was during May, and it was one of the most productive collecting trips I’ve had with a number of Agrilus spp. (including A. frosti) beaten from post oak (Quercus stellata) along the woodland edge.  I didn’t visit again until fall of 2002, when Chris Brown, Rich Thoma and I found claybank tiger beetles (Cicindela limbalis) sunning on the exposed sandstone outcrops, and I made one more visit the following May to beat more insects off of post oak.  The Lichen Glade that I returned to this past weekend was a very different place from when I last visited – the surrounding woodlands had been extensively opened (I would guess within the past few years based on the size of the post oak resprouts), and fire had been used throughout the area.  Anticipation turned to frustration when no amount of beating of the woodland vegetation and sweeping of the glade vegetation turned up beetles in any appreciable numbers (or any insects for that matter) and two hours worth of effort yielded not a single buprestid beetle!

Sandstone glade community, Lichen Glade Natural Area

With resignation, I headed on over to Penn-Sylvania Prairie, where during the introduction to the BioBlitz I learned that nearly half of the 160-acre prairie was burned last December and all of it had been burned within the past few years.  I knew what I was going to find – nothing!  Okay, I shouldn’t say nothing, as there actually were some beetles present.  However, the numbers and diversity were low, with all of the species encountered representing common, widespread species.  Moreover, it was not just beetles – all of the invertebrate group leaders (which included experts on snails, ants, butterflies, and bees) reported low overall abundance and diversity in their groups of interest.  Only the vascular plants – the metric by which the value of prescribed burning is always assessed – showed high diversity, with 300 species of mostly native prairie plants recorded for the site.  It was a fun event, with probably ~75 attendees and a delicious pot luck dinner that evening; however, it would have been more enjoyable had there actually been a nice diversity of insects present to document for the preserve.

My comments may make it seem that I am against the use of prescribed burning.  This is not true – I understand the critical role that fire as a management technique plays in restoring and maintaining examples of Missouri’s historically fire-mediated landscape. Without fire and other processes to mimic natural disturbance factors, most of Missouri’s historical grasslands and woodlands suffer relentless encroachment by woody vegetation. However, the modern landscape is very different from the historical landscape, where fires of unpredictable scale, intensity, and frequency operated within a vastly larger scale to create a shifting mosaic of natural communities in various stages of ecological succession. Such processes cannot be recreated on today’s severely fragmented landscape, where the precious few remaining tracts of native habitat are relatively to extremely small and more often than not separated from each other by vast expanses of homogeneous and “inhospitable” habitat (e.g., agricultural, urbanized, or severely degraded lands).  It is in that context that I have great concerns about how aggressively fire has been used in recent years on our state’s natural areas and the impact this is having on insect populations – specialist and generalist alike.  Fire proponents will point to published studies that show little to no effect by the use of fire for managing small, isolated remnants on specialist insects (see review in Henderson 2010).  However, there are an equal number of studies that suggest such concerns are well-founded (see review in Panzer 2002). A consistent limitation in all of the studies that have been conducted is the lack of very large and long un-burned remnants.  Prescribed burning has been adopted so rapidly and pervasively that there just aren’t any significant un-burned remnants left to properly include as controls in such studies.  As a result, the insect fauna present at a given site at the start of such a study is already skewed towards those species that successfully recolonized the area post-burn.  At a minimum, the data to this point are inconclusive, and certainly the potential for impacts has not been given the consideration it warrants in designing fire-management plans for our own state’s prairies and glades. Furthermore, as rapidly and aggressively as fire has been adopted on our few, small, widely disjuct remnants, the opportunity for proper investigation of those potential effects may be gone.  A particularly egregious example of the lack of consideration being given to prairie invertebrates in designing fire management plans is shown in these photos of Iowa’s Sylvan Runkel State Preserve before and after a late May burn and the impact of that burn on a resident population of Nevada buck moths (Hemileuca nevadensis).

Here in Missouri, as in Iowa, it’s a problem of scale – the landscape is too fragmented and remnants too disjunct to manage based strictly on floristic response.   Populations of generalist insect species will recover, and even specialist species may be able to overcome such management practices if they are widely distributed and sufficiently mobile. But what about conservative species with low vagility, such as the swift tiger beetle (Cylindera celeripes) and our disjunct population of the frosted dromo tiger beetle (Dromochorus pruinina), flightless species restricted in Missouri to the few tiny remnants of loess hilltop prairie in northwestern Missouri and a single 2.5-mile stretch of roadside habitat in west-central Missouri?  Until directly relevant data, gathered here in Missouri, are forthcoming to suggest otherwise, I believe the most judicious use of fire possible should be practiced in restoring and maintaining our grasslands and woodlands.  In-season burns may have been a part of the historical landscape, but their use today has great potential to result in local extirpations and should be used only after the most careful consideration.  Leaving un-burned refugia within remnant habitats to accelerate recovery would also be prudent – yet many land managers disregard this practice because of its logistical difficulties. This is especially true in small parcels, yet it is precisely these remnants that have the most to gain from their use (or lose from not doing so!).  In the historical landscape, every burn was a patch burn – no matter what its size, there were always adjacent or proximal unburned habitat from which recolonization could occur.  Elk and bison, too, were integral components of the presettlement prairie landscape – their roamings caused intermittent, localized disturbances that were likely not only crucial to the tiger beetles that I study but may also have contributed to vegetational diversity through patch succession.  Techniques that mimic these natural disturbance factors include mowing, haying, and managed grazing.  They can be utilized to mimic those disturbances as well as delay woody encroachment, and their use in land management should be considered for their ecological value rather than deprioritized because of their relatively greater complexity and cost to implement. Mechanical removal and selective use of herbicides offer additional tools for addressing woody encroachment while minimizing potential impacts to insect populations. An effective management program that considers all of the flora and fauna of a remnant may not be possible unless all of these management tools are utilized, or at least properly considered. As my good friend James Trager said in a recent email (quoting Andrew Williams), habitat restoration “cannot rest on any single management practice, nor practicing it too extensively.”

REFERENCES:

Henderson, R. A.  2010. Influence of Patch Size, Isolation, and Fire History on Hopper (Homoptera: Auchenorrhyncha) Communities of Eight Wisconsin Prairie Remnants.  Wisconsin Department of Natural Resources, Research Report 189, 22 pp.

Panzer, R. 2002. Compatibility of prescribed burning with the conservation of insects in small, isolated prairie reserves. Conservation Biology, 16(5):1296-1307.

Copyright © Ted C. MacRae 2010

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Cylindera celeripes Larva Revealed

/ Ted C. MacRae / 22 Comments

In a recent post, I provided the first ever glimpse of the previously unknown larva of Cylindera celeripes, or swift tiger beetle.  This little-known flightless species is among the tiniest in North America (adults measure only 8 or 9 mm in length), and so far nobody has succeeded in rearing the species in the lab, or even finding its larva.  As the photographs in that post showed, I am reasonably close to accomplishing that first goal, having successfully obtained a number of eggs from field-collected adults placed in a terrarium of native soil. I fed the subsequent larvae a diet of small rootworm larvae and Lygus nymphs before putting them to sleep for the winter in a cold incubator, and the larvae resumed activity when I pulled them out of the incubator 2 months ago. Since then, they have feasted heavily on small noctuid larvae that we rear in our lab, and now most of the dozen or so larvae have sealed their burrows – I presume for pupation before (hopefully) emerging as adults in the next few weeks.

Cylindera celeripes 3rd instar larva - USA: Oklahoma, Woodward Co.

There is more to the story, however.  I had brought the adults back home in June 2009 from a population I found at Alabaster Caverns State Park in northwestern Oklahoma.  This was a reasonably robust population – news enough for a species that has not been seen in good numbers for many years now, and my discovery of equally healthy populations at several other locations in the general area gives new hope for the long-term prospects of a species that some regard as a potential candidate for listing as an endangered species. It also gave me hope that I might be able to find the larva were I to return to the area in the fall.  I also had a hunch that Cicindela pulchra (beautiful tiger beetle) could be found in the area, based on some very large larvae I found during that June trip, so in early October I made a quick return to northwestern Oklahoma to search for these two species.  While it was too cold and wet to have any hope of finding Cicindela pulchra adults (I still think the species is there), it did not prevent me from realizing my other goal.  May I present one of the first ever field-collected larvae of Cylindera celeripes!

Cylindera celeripes 3rd instar larva - closeup of hump on 5th abdominal segment with hooks to aid in securing the larva in its burrow

I found the larvae at Alabaster Caverns where I had found the adults earlier in June, and although the larval burrows were very small (only 2 to 3 mm in diameter), I knew what they were immediately when I saw them.  As I had observed for the adults, burrows tended to be near the edges of barren patches of soil in proximity to vegetation and not out in the middle of the barren areas.  This makes sense, considering where it would be more likely for prey to be encountered.  Because the weather was cold and gray, I didn’t see (or expect to see) larvae actively sitting at the tops of their burrows, so I began “fishing” to see if I could yank a few from their burrows.  I fished quite a few burrows for the first half hour or so, but none of my attempts were successful.  I began wondering if the larvae were even active at all or if they had already entered hibernation for the upcoming winter.  While I was fishing, I noticed that the burrows all seemed rather shallow – only about 6” or so (most tiger beetles, having larger larvae, dig burrows that are much deeper). This gave me an idea.  I went back to the truck and retrieved a small spade that I carry in case… well, I’d never actually used it before.  Anyway, I inserted a grass stem into a burrow and sunk the spade into the ground right next to it, making sure I got the spade at least as deep as the grass blade.  I then removed the spade and sunk it into the ground on the other side of the burrow, then pried until the entire chunk of soil came up intact.  With the bottom of the soil chunk exposed, I used my knife to carefully remove slivers of soil until I found the end of the grass stem that I had inserted into the burrow.  Carefully removing the soil in this area revealed the larva in a side chamber at the bottom of the burrow.  Success!  I took many photos of that larva right then and there, and over the next hour or so collected several more larvae, all but one of which I presumed were 3rd instars.  I packed each larva in its own small vial of native soil for the trip home, and although I have been attempting to rear them out for confirmation of their identity, there is little doubt that they do indeed represent this species.

Cylindera celeripes 3rd instar larva - that grotesquely beautiful head!

The photographs I’m showing here are not those first field photographs that I took when I first discovered the larvae.  Looking at those photographs after I returned home, I was dissatisfied with the amount of soil and debris that covered the larvae – especially their grotesquely unique head and pronotum.  Instead, I removed one of the larvae from its rearing tube and gave it a “bath” – brushing it with a fine camel-hair brush in a shallow dish of water – to clean it up for the photographs shown here.  After the photo shoot, I sacrificed this larva for the collection – it will be the basis for a formal description of the larva of this species (along with examples of the 1st and 2nd instars that I had sacrificed from my rearing, not yet confident that I would succeed in getting any of the others to 3rd instar).  The only thing I am waiting on before preparing that description is to see whether I actually succeed in rearing this species from egg to adult – stay tuned!

Photo Details: Canon 50D (ISO 100, 1/250 sec, f/13-16). Canon MP-E 65mm 1-5X macro lens, MT-24EX flash (1/8 power) w/ Sto-Fen diffusers. Typical post-processing (levels, unsharp mask).

Copyright © Ted C. MacRae 2010

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Hello World!

/ Ted C. MacRae / 28 Comments

These are two of the Cylindera celeripes (swift tiger beetle) larvae that I’m rearing.  Note: nobody has ever reared this species before!  Nobody has ever even seen its larvae (before now, that is).

These larvae hatched from eggs that were laid by adults I brought back from northwestern Oklahoma last summer.  I placed the adults in a small terrarium of native soil – at first just to see if I could keep them alive, and then to see if I could get them to lay eggs.  The adults lived for about 4 weeks, and a short time later larval burrows started appearing in the soil.  I fed them once or twice a week by placing 2nd instar corn rootworm larvae in the open burrows or dumping Lygus nymphs into the terrarium and letting them catch them naturally.  I wasn’t sure this was working, because as the summer progressed I saw fewer and fewer open burrows.  By October, there were no open burrows, and I feared none had survived.  Nevertheless, I placed the terrarium in a cool (10°C, or 50°F) incubator for the winter and pulled it back out in late March.  Within one week ten larvae had reopened their burrows – I believe all but one of them are 3rd instars, which is the last instar before pupation, and since they have awoken they have fed voraciously on 3rd instar fall armyworm larvae, which I dangle above their burrow.  I love watching them snatch the armyworm from my forceps and drag the hapless prey down into their burrow.  I’ve already preserved examples of the three larval instars and will describe it shortly (although truth be told, the 2nd and 3rd instars are from larvae I found in the field – but that is a post for another day).  However, I’m keeping my fingers crossed for the piece de resistance – successfully rearing the species from egg to adult!

Photo Details: Canon MP-E 65mm 1-5X macro lens on Canon 50D, ISO 100, 1/250 sec, f/13-16, MT-24EX flash 1/8 power w/ Sto-Fen diffusers.

Copyright © Ted C. MacRae 2010

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A Tiger Beetle Aggregation

/ Ted C. MacRae / 17 Comments

Not long ago, I received an interesting series of photographs from Joe Warfel, a nature photographer and macro specialist based in Massachussetts.  Joe traveled to Arizona last July, where he photographed an aggregation of Cicindela (Cicindelidia) sedecimpuntata (Western Red-bellied Tiger Beetle¹) near a small pool in the bottom of a dry creek bed at night.  Joe estimates that there may have been as many as 200 to 300 beetles per square meter in the aggregation, most of which were just “hanging out” and with only occasional individuals mating or feeding on moths that had been attracted to his headlamps.

¹ Found in the Sonoran and Chihuahuan Deserts of the southwestern U.S. and south through Mexico to Costa Rica. U.S. and northern Mexican populations are assigned to the nominate subspecies, while more southern populations are classified into four additional subspecies (Erwin and Pearson 2008).

Western Red-bellied Tiger Beetles are among the first tiger beetles to appear prior to the summer monsoons in the Sonoran Desert.  The species is famous for its daytime aggregations of as many as several thousand individuals, which congregate along the drying waterways and prey upon stranded tadpoles and other aquatic organisms (Pearson et al. 2006).  Joe noted that he has seen these aggregations many times before during the daytime at small pools and mudflats, with beetles usually mating and feeding frantically.  However, the aggregation shown in these photographs differs from those daytime aggregations by the relative inactivity of the beetles and the fact that they were congregated on dry ground rather than the moist areas that they frequent during the daytime.  In these respects, it seems to more resemble a communal nocturnal roost such as has been reported for several species of Odontocheila in South America.  In those cases, up to 70 beetles have been found resting on the foliage of low shrubs, apparently as an adaptation to avoid predation by multiplying chemical defense effectiveness as well as awareness of approaching enemies (Pearson and Vogler 2001 and references therein).  Cicindela sedecimpunctata is primarily a diurnal species (i.e., it is active during the daytime), though individuals are often attracted to lights at night, and adults of most diurnal species have been reported spending the night protected in burrows or under detritus and vegetation.  I am not aware of communal nocturnal roosts as a reported behavior for C. sedecimpunctata or any other North American tiger beetle species.

It is a bit ironic to think of tiger beetles – voracious predators that they are – as prey, but they must have many of their own predators to deal with since most species employ multiple antipredator mechanisms. In addition to the communal roosting behavior seen in these photos, a second antipredator characteristic exhibited by this species can be seen in their bright orange abdomen.  The abdomen is fully exposed only during flight, seemingly implying a “flash coloration” function for the bright color that disappears upon landing, momentarily confusing potential predators.  However, Pearson (1985) experimentally determined that orange abdomens in tiger beetles actually have an aposematic function in protecting them from predation against robber flies.  Most tiger beetle species with an orange abdomen also release a combination of benzaldehyde and cyanide² when captured (any tiger beetle collector is familiar with the characteristic “fruity” smell of a tiger beetle releasing benzaldehyde).  Pearson painted the abdomen of paper tiger beetles models either orange or black and endowed them with or without a drop of fresh benzaldehyde.  When presented on a tether to robber flies in the field, orange-abdomened models with benzaldehyde triggered significantly fewer attacks from robber flies than any other combination.  Interestingly however, vertebrate predators (lizards and birds) were not deterred by the defense chemicals or by the orange abdomen, perhaps explaining why only some and not all tiger beetle species produce defense chemicals and have bright orange abdomens (Pearson and Vogler 2001).

² Tiger beetles, thus, join millipedes as being among the few invertebrates that are capable of producing cyanide.

My sincere thanks to Joe Warfel for allowing me to use his photographs. More of his work can be seen at Eighth-Eye Photography.  Joe also recently had several images published in American Scientist magazine (November/December 2009 issue) for an article on harvestmen.  Check out the jaws on that juvenile!

REFERENCES:

Erwin, T. L. and D. L. Pearson. 2008. A Treatise on the Western Hemisphere Caraboidea (Coleoptera). Their classification, distributions, and ways of life. Volume II (Carabidae–Nebriiformes 2–Cicindelitae). Pensoft Series Faunistica 84. Pensoft Publishers, Sofia, 400 pp.

Pearson, D. L.  1985.  The function of multiple anti-predator mechanisms in adult tiger beetles (Coleoptera: Cicindelidae).  Ecological Entomology 10:65–72.

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.

Pearson, D. L. and A. P. Vogler.  2001. Tiger Beetles: The Evolution, Ecology, and Diversity of the Cicindelids.  Cornell University Press, Ithaca, New York, 333 pp.

Copyright © Ted C. MacRae 2010

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Alkali Tiger Beetle

/ Ted C. MacRae / 4 Comments

Eunota togata globicollis - Salt Plains NWR, Oklahoma

I haven’t written much about my early October trip to Oklahoma, where I had hoped to confirm a hunch that the gorgeous Cicindela pulchra (Beautiful Tiger Beetle) would be found in the red clay/gypsum hill habitats of Woodward and Major Counties (the same place where I had found the much rarer Cylindera celeripes the previous June).  Unfortunately, a sudden cold snap and overcast skies conspired against me for the duration of that short, 5-day trip, reducing tiger beetle activity to near zero and sending me back to Missouri with little to show for my efforts — save a scorpion, a torpid Cicindela splendida, and some very beautiful ladie’s-tresses orchids in peak bloom.  I did have one moderately successful day, however, when I returned to Salt Plains National Wildlife Refuge in north-central Oklahoma, a place where I observed seven species of tiger beetles during my June trip.  An eighth species that I did not see on that trip, but which I had observed in previous years, was my goal this time, and despite the cold temperatures and cloudy skies I was fortunate to find several individuals of Eunota togata globicollis.  Occurring primarily on saline flats in the central and southern Great Plain, this subspecies was called the Alkali Tiger Beetle¹ by Erwin and Pearson (2008), who reserved for the nominate form (found in salt marshes and tidal flats along the Gulf Coast) the more descriptive name White-cloaked Tiger Beetle².  A third subspecies, E. togata fascinans (Salt Flat Tiger Beetle) is restricted to salt flats in central New Mexico and west Texas (Pearson et al. 2006) (you may remember this subspecies from my habitat partitioning post last month).

¹ In reality, I have come to consider the term ‘alkali’ as a bit of a misnomer, as it is saline soils specifically — not just those with high pH (alkaline) — that the species is fond of. Moreover, there are many species of tiger beetles in addition to this one that are associated with saline soils.

² Okay, I might as well just get all this off my chest. Pearson et al. (2006) gave common names to each species of tiger beetle in the U.S., but not subspecies. I think most non-taxonomists probably consider this a good thing, although it is not without its problems (some species already had multiple common names applied to them, forcing choices that are sure not to please everyone). Erwin and Pearson (2008) took this further and came up with common names for all of the subspecies as well, and like any good taxonomist they steadfastly applied existing common names only to nominate forms. Eunota togata, however, is an example where the original common name would have been better applied to one of the non-nominate subspecies. The species epithet togata means “cloaked” (being derived from the Latin word toga — a reference to the broad white band running along the elytral margins). Each of the two non-nominate forms are distinguished by the white band being more broadly expanded (indeed, almost entirely covering the elytra in subspecies fascinans), yet it is the nominate subspecies — the least “cloaked” of the three — that retains the original common name. A silly argument I suppose, but if we start applying the “prinicple of priority” to common names in the same manner as scientific names, then what have we gained? Of course, I am of the opinion that most insect groups are too diverse and their taxonomy still too unstable to warrant a rigid system of “official” common names. Is it really any easier to learn White-cloaked Tiger Beetle than Eunota togata? How about Mount Ashland Night-stalking Tiger Beetle instead of Omus cazieri? And this is not even considering what happens when category-level shifts occur. For example, the genus Tetracha was formerly called the Big-headed Tiger Beetles; however, its former subgenera were recently elevated to genus level. Erwin and Pearson, accordingly, applied the common name to the entire subtribe containing Tetracha and its relatives and applied a new common name, Metallic Tiger Beetles, to the new, more limited concept of Tetracha. Thus, in an ironic case of common name instability despite no change in scientific name, the Virginia Big-headed Tiger beetle (Tetracha virginica) became the Virginia Metallic Tiger Beetle. Are your eyes bugging yet? Common names may be appropriate for higher vertebrates, but can they really be used effectively for beetles and other insect groups where the increasing use of molecular tools is sure to result in additional, perhaps radical, shifts in taxonomy? There — I said it, and I feel a lot better!

This species is restricted to saline flats in the central/southern Great Plains.

Of the eight tiger beetle species that I’ve now observed at Salt Plains NWR, half of them (Cicindela fulgida, C. nevadica knausii, E. togata globicollis, and Habroscelimorpha circumpicta johnsonii) are true saline habitat specialists.  One of the other four species (Cicindela tranquebarica kirbyi) is also fond of saline habitats but also occurs commonly on dry, sandy soils as well, and two show a high affiinity for nearly any moist (Cicindela repanda) or moist to dry (Cicindela punctulata) soils with little regard for salinity.  Only Cicindela formosa, a denizen of dry, deep sands seems a little out of its element on the moist, salty mud at Salt Plains NWR — perhaps the few individuals I’ve observed here are incidental visitors, mistaking the white, barren expanses of salt-encrusted soil for the dry sand the species prefers during disperal searches.  This again brings up the question of habitat partitioning for competition avoidance among tiger beetle species sharing the same habitat.  Eunota togata globicollis is active during the spring and fall and, thus, temporally isolated from C. nevadica knausii and H. circumpicta johnsonii (both summer-active species).  The other saline specialist at Salt Plains NWR (C. fulgida) is active during the same seasons as E. togata globicollis; however, in my observations that species prefers the sparsely-vegetated zone at the edge of the saline flats, while E. togata globicollis prefers to stay out in the more open areas.  These observations mirror those of Melius (2010) for E. togata fascinans and the other seven species he noted in the Laguna del Perro area of New Mexico, and of Willis (1967), who recorded as many as 11 sympatric tiger beetle species in saline habitats in the central U.S.

Saline flats at Salt Plains NWR are home to eight species of tiger beetles.

Microhabitat selection and seasonal occurrence are not the only isolating mechanisms that can minimize interspecific competition among the different tiger beetle species at Salt Plains NWR.  Cicindela tranquebarica kirbyi is also a spring/fall species and doesn’t appear to display a preference for open versus vegetated areas, potentially allowing it to compete directly with both E. togata globicollis and C. fulgida.  However, C. tranquebarica kirbyi is a decidely larger species, while the other two are smaller, and correlated with such differences in overall size is the size of their mandibles.  Mandibular size directly correlated to prey size in a number of tiger beetle species (Pearson and Mury 1979), thus providing another mechanism for avoiding competition between these three co-occurring species. 

Photo details:
Beetles: Canon 100mm macro lens w/ 68mm Kenco extension tubes on Canon EOS 50D (manual mode), ISO 100, 1/250 sec, f/18-20, MT-24EX flash 1/4 power w/ Sto-Fen diffusers.
Landscapes: Canon 17-85mm zoom lens (22mm) on Canon EOS 50D (landscape mode), ISO 100, 1/100 sec, f/10, natural light.

REFERENCES:

Erwin, T. L. and D. L. Pearson. 2008. A Treatise on the Western Hemisphere Caraboidea (Coleoptera). Their classification, distributions, and ways of life. Volume II (Carabidae-Nebriiformes 2-Cicindelitae). Pensoft Series Faunistica 84. Pensoft Publishers, Sofia, 400 pp.

Melius, D. A. 2009. Post-monsoonal Cicindela of the Laguna del Perro region of New Mexico. CICINDELA 41(4):81-89.

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.

Pearson, D. L. and E. J. Mury. 1979. Character divergence and convergence among tiger beetles (Coleoptera: Cicindelidae). Ecology 60:557–566.

Willis, H. L.  1967.  Bionomics and zoogeography of tiger beetles of saline habitats in the central United States (Coleoptera: Cicindelidae).  The University of Kansas Science Bulletin 47(5):145-313.

Copyright © Ted C. MacRae 2010

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