Anatomy of a Tiger Beetle Larva

My first experience looking for tiger beetles in Florida had gone well.  Despite its small size and urban surroundings, the narrow strip of coastal scrub and saltwater marsh along the intracoastal waterway behind my sister-in-law’s condominium boasted a robust population of what I took to be a single tiger beetle species.  The specimens I collected and photographs I took would later reveal that two co-occurring and closely related species were present: Ellipsoptera marginata (Margined Tiger Beetle) and E. hamata lacerata (Gulf Beach Tiger Beetle).  I had spent close to two hours under the August sun observing and photographing the beetles before I decided that I had given the preserve a thorough enough look.

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As I was heading back, I noticed a little bit of high ground alongside a red mangrove thicket and went over to give it a look.  As I approached I saw something I hadn’t yet seen that day – tiger beetle larval burrows.  Larval burrows, especially larger ones such as these were, are unmistakeable – almost perfectly circular (slightly cut out on one edge) and smoothly beveled around the perimeter.  There were a number of burrows clustered on the small bit of high, dry ground, and my first thought was that their inhabitants represented the same (what I thought was a single) species that I had encountered so commonly that day as adults.  I then reasoned, however, that more likely they represented another species whose adults are active later in the season – perhaps one of the so-called “spring/fall” species whose larvae typically reach maturity during the heat of summer.  The size of the burrows (~5mm dia) suggested they were inhabited by 3rd instar larvae (the final instar before pupation), in which case it may be possible to rear a few to adulthood – if I could get at them. I tried fishing (Pearson and Vogler 2001) a few holes with a grass blade but didn’t get any bites, so I decided to watch for awhile and see if any of the larvae, believing the danger of my approach had passed, would reappear at the tops of their burrows.  Waiting for tiger beetle larvae to appear is a crap shoot – maybe they’re active, and maybe they’re not, and crouching in the stifling summer air of a coastal marsh in Florida is not an easy thing to do for very long.  Fortunately my wait was short, as within a few minutes I saw one re-appear at the top its burrow.  I slowly got out my knife and moved to place the tip on the soil about 1″ from the burrow at a 45° angle for an attempted tunnel block (Pearson and Vogler 2001), but it spooked and dropped back down into its burrow before I could get then knife in place.  No matter, I knew it was in there now and that it would likely reappear if I could muster the patience.  I positioned the knife and waited – crouched under the baking Florida sun, until when it did re-appear I plunged the knife into the soil with authority.  It was a good jab – I had blocked its retreat without injuring it, and a quick flip of the knife popped out the soil plug and exposed the startled larva, flipping vigorously in a vain attempt to escape before settling down amidst its unfamiliar, exposed surroundings.

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For those of you who have never seen a tiger beetle larva, they are among the most other-wordly creatures one can imagine.  The large, heavily sclerotized head bears two long, sickle-shaped, upward-pointing mandibles and up to three pair of highly-acute eyes whose arrangement on each side conveys the image of a “face” with congenital birth defects.  The top of the head is flattened to lie flush with the surrounding soil as the larva sits at the top of its burrow, and huge, powerful mandibular muscles fill the cranial cavity.  The remainder of the body – long, narrow, and cylindrical – hangs from the head at a 90° angle down into the burrow and is unremarkably grub-like, save for a curious hump on the dorsal side of the 5th abdominal segment.  Close examination of the hump reveals an intricate pattern of forward-facing hooks and spines that function in anchoring the larva against the side of its burrow to prevent struggling prey from dislodging it.  The life of a tiger beetle larva is a life of waiting – unlike the adults who run down their prey, the larvae sit in their burrows and wait for prey to come to within lunge’s reach.  While the eyes of most grub-like insects detect little more than light and dark, those of tiger beetle larvae are densely packed with photoreceptors that permit detailed focusing and depth perception for detecting whether potential prey has ventured close enough to their burrow (Pearson et al. 2006).  When that happens, they strike with lightning speed, plunge their mandibles into their prey, and drag it down into the depths of their burrow where it is summarily dispatched with a few bites of their powerful mandibles.  Larvae consume they prey in a manner similar to that of adults in that they chew but don’t swallow their prey. Rather, they secrete digestive secretions containing proteolytic enzymes that begin digesting the prey extra-orally as they chew.  The resulting bolus is masticated and its liquid components sucked out until nothing but a dry wad of indigestable chitin remains, which is spat out of the burrow (Pearson and Vogler 2001).

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Looking at this strange insect, it occurred to me that I had not yet attempted macrophotographs of a tiger beetle larva out of its burrow, and this would be a good opportunity to get more practice with my Canon MP-E 65 mm macro lens – a lens with incredible magnification capabilities, but one that is also a bit of a temperamental beast to use hand-held in the field.  The subject was unusually cooperative, perhaps too stunned by its sudden predicament to know what to do, and as I took the photographs I focused in particular on characters of the head and dorsal hump (often useful in identifying tiger beetle larvae, at least to genus).  Time was growing short once I finished taking photographs, so I placed the larva in a vial and returned the following day to extract a chunk of native soil to place in a rearing container, managing to collect two more larvae as well (unfortunately, one became instant “prey” for the other.  Note to self: when placing multiple tiger beetle larvae in a container of soil, seal the artificial burrows into which you place each one!).  I paid little further attention to the photographs, other than to transfer them onto my computer and add metadata upon my return to St. Louis.  I didn’t know what species the larvae represented, but I assumed they were something in the genus Cicindela or one of its several former subgenera.  However, had I studied the photos and considered the locality and habitat, I would have realized that my assumption was incorrect¹.  That realization would come in surprise fashion two months later when the two adults emerged within a few days of each other…

¹ Ten points to whoever can use this information to arrive at an identification before my next post 🙂

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Photo details:
All photos: Canon EOS 50D, manual mode, ISO-100, 1/250 sec, MT-24EX flash w/ diffuser caps.
Photo 1: Canon 100mm macro lens, f/22, 1/4 power flash (photo slightly cropped).
Photos 2-4: Canon MP-E 65mm 1-5X macro lens f/16, 1/8 power flash.

REFERENCES:

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 2009.

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(My) Introduction to Florida Tiger Beetles

On the last day of July, I hopped into my stuffed-to-the-brim SUV and began the +1000-mile drive to St. Petersburg, Florida.  I had with me my camera and my collecting gear, but also my bike, some beach wear… and my family!  No, this was not a collecting trip, but a family vacation.  We would be staying at my sister-in-law’s condo with my niece and enjoying typical Florida vacation fare – beaches, Disney World, sightseeing.  I even brought my road bike along to enjoy some pancake-flat terrain (a rarity around St. Louis) for a nice change of pace.  Still, I can be rather single-minded when it comes to opportunities for bug collecting, and summer in Florida presents opportunities galore!  Virtually everything there is different, and while I have been to Florida a number of times, my visits have all been during spring and prior to my more recent interest in tiger beetles.  It would be a family vacation, but still I would find a way to sneak off a couple times and satisfy my compulsions, and while the girls talked about Disney World and the beach, visions of Cicindela abdominalis, C. highlandensis, C. scabrosa, Ellipsoptera gratiosa, E. hamata lacerata, E. hirtilabris, E. marginata, Habroscelimorpha dorsalis, H. severa, and H. striga danced through my head!

I wouldn’t have to go far to find my first Florida tiger beetles.  My sister-in-law’s condo is in Seminole (west side of St. Petersburg peninsula), and as my niece and her husband showed us around after our arrival, they pointed out the “wild area” off their back patio next to the intertidal waterway and suggested, innocently, that I could go bug collecting back there.  In such a developed urban/tourist area, I figured it must be highly disturbed, dominated by exotics, and offering little in the way of quality habitat for the serious bug collector.  However, that night, as a cacophony of nasal queenks from the area made it clear that a healthy population of what I presume to be green treefrogs were thriving in the area, I reconsidered my skepticism and decided to take a quick look a day later.  What I found was a small but high-quality strip of coastal scrub and saltwater marsh bordering the intracoastal waterway, with thickets of red mangrove (Rhizophora mangle) along the water’s edge, black mangrove (Avicennia germinans) “woodlands” in the high scrub areas, and moist, briney, barren ground in between (Photo 1).  The whole area couldn’t have been more than a hundred yards wide but extended along the length of the waterway, and a wooden sign as I entered the area indicated it was a bona fide, albeit private, nature preserve established as part of the condominium development.

Intertidal salt marsh, Tara Cay Sound Nature Preserve, Seminole, Florida

Intertidal salt marsh, Tara Cay Sound Nature Preserve, Seminole, Florida

Within minutes after entering the preserve I saw the first tiger beetle.  I didn’t know what it was, and my first stalking efforts were woefully inadequate.  I saw another one a few minutes later and got within net handle distance but muffed the swing.  A few feet further along the path and onto an exposure by the water revealed several individuals, one of which I was finally able to capture after several clumsy misses.  I hadn’t yet memorized key characters for all of the species I had the potential to encounter, but I could see in the hand that this individual almost certainly belonged to the genus Ellipsoptera (Ellipsed-winged Tiger Beetles) because of its resemblance to E. macra (Sandy Stream Tiger Beetle) and E. nevadica knausii (Knaus’ Tiger Beetle), which I had photographed earlier this year in Missouri and Oklahoma, respectively.  With one now in the hand as a studio backup, I began my efforts to obtain field photographs.  The beetles were extremely wary, with fast running and strong flight capabilities that made them very difficult to approach.  It was only their abundance and my dogged persistance that allowed me to finally get close enough to one (Photo 2) to fire off a sequence of frames.  I then spent some time collecting a voucher series – finally getting a beat on their behavior and able to capture them with a little more efficiency despite their ultra-wariness before concentrating on getting more photographs of different individuals (including those shown in Photos 3 and 4).

Ellipsoptera marginata - Margined Tiger Beetle

Ellipsoptera marginata - note "tooth" under right mandible of this male

Ellipsoptera marginata - Margined Tiger Beetle

Ellipsoptera marginata - another male, with a distinctly bronzed pronotum

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Ellipsoptera hamata lacerata - Gulf Beach Tiger Beetle (mate guarding)

I returned to the condo after a couple of hours – completely drenched (remember, it was August in Florida!). When I had the chance to consult my “bible” (Pearson et al. 2006), I learned there there were two possibilities: Ellipsoptera marginata (Margined Tiger Beetle), occurring along the Atlantic seaboard from Maine to the Gulf coast of Florida; and Ellipsoptera hamata lacerata (Gulf Beach Tiger Beetle), resembling and closely related to E. marginata and occurring along the Gulf Coast from Florida to Texas.  Both species inhabit coastal beaches, mud flats and salt marshes, and their distributions overlap along Florida’s Gulf Coast. While they are easily distinguished from other species in the genus by the distinctively diffuse middle band of the elytra, distinguishing between the two requires examination of the male right mandible (marginata bears a distinct tooth on the underside, hamata does not) or female elytral apices (marginata curiously bent down at a 90º angle, hamata not).  At this point, I didn’t know if I had one species or two – and if I did have two, did I have photographs of both (and would I be able to identify them)?

Fortunately, closer examination of the voucher series I collected revealed both species present, and even more fortunately I had managed to get photographs of both.  The individual in Photo 2 is a male, and the angle of the photograph clearly reveals a distinct tooth on the underside of the right mandible, identifying it as E. marginata.  Photo 3 is another male, and although the angle doesn’t afford a view of the mandibles, what can be seen is a bronze cast to the pronotum – in my voucher series, all of the E. marginata specimens have a bronze cast to the pronotum, while the E. hamata lacerata specimens exhibit an olive cast. This suggests that this individual also represents E. marginata. In Photo 4, no tooth can be seen on the male mandible, but the angle of the photo doesn’t necessarily make it visible were it to exist.  The male does, however, exhibit an olive cast on the pronotum, and the female elytral apices show no indication of being bent down as would be expected for E. marginata (see Photo 5 below of E. marginata photographed a few days later at another locality).  As a result, the individuals in Photo 4 can be identified as E. hamata lacerata.

Ellipsoptera marginata - female elytral apices showing curiously "bent tips"

Ellipsoptera marginata - female elytral apices showing curiously "bent tips"

Of the 19 vouchers that I collected, 15 represent E. marginata and 4 represent E. hamata lacerata.  This exemplifies the challenges of field identification and photography of insect species in unfamiliar places.  Had I been satisfied with photographing only one or two individuals of what I thought were all the same species, I would have missed one of the species.  The experience further exemplifies the importance of adequate voucher series, as it was only a result of close examination of that series that I realized two species were present.  Based on the numbers of individuals that I collected, as well as the photographs that I took, it appears that E. hamata lacerata was much less common at that location than E. marginata. This difference in population density between two similar, co-occurring species emphasizes the importance of sampling an adequate number of individuals at a given location before concluding what species are – and are not – present.

REFERENCES:

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 2009

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Cold and wet

Photo details: Canon 65mm 1-5X macro lens on Canon EOS 50D, manual mode, ISO-100, 1/250 sec, f/14, MT-24EX flash 1/8 power through diffuser caps.

Photo details: Canon 65mm 1-5X macro lens on Canon EOS 50D, manual mode, ISO-100, 1/250 sec, f/14, MT-24EX flash 1/8 power w/ diffuser caps

The annual fall tiger beetle collecting trip is over, and nothing could be more emblematic of the trip than this cold, wet Cicindela splendida (literally translating to its well-deserved common name “Splendid Tiger Beetle”), found in the waning light of the last day of the trip secreted under a rock in the dolomite glades at White River Balds Natural Area in southwestern Missouri.  Overcast skies and a cold, stiff breeze had already dashed my hopes of finding Cicindela pulchra (also translating to its equally well-deserved common name “Beautiful Tiger Beetle”) in the Red Hills of northwestern Oklahoma two days earlier, and these same unrelenting conditions thwarted my backup plans to find this species the next day in the Gypsym Hills of nearby Barber County, Kansas.  With one last day to spend in the field, I had worked my way back to the White River Hills in hopes of photographing Missouri’s disjunct population of the enormous Cicindela obsoleta vulturina (Prairie Tiger Beetle).  That would also not come to pass, as the sun’s efforts to burn through the thick cloud cover just weren’t quite enough.  By the end of the day, I had resorted to flipping rocks (and replacing them exactly, of course) in hopes of finding an individual or two still in their nighttime/cool weather roosts.  I had seen C. obsoleta on previous occasions at this very locality along this very trail, but this time none were found – my finds instead limited to the “smiling” scorpion that I featured a few days ago and this lone C. splendida – wet with condensation and torpid against the cold ground.  I hatched a last gasp plan to look for C. obsoleta again the following day before heading back to St. Louis, but I awoke the next morning to steady rain and knew the 2009 entomology field season was officially over.

Don’t let me leave you with the impression, however, that the trip was a failure.  While I didn’t find either of the two species that I had set as my top goals for the trip, I still saw enough new things to make the trip worthwhile (including one very significant find that I’ll discuss in an upcoming post).  In addition, I’ve barely made mention of my August trip to Florida – my Assasin ate post of a couple weeks ago was just one of many interesting finds encountered during that trip that remain to be shared.  The cold and rain that has settled over the middle of the country during the past week have brought an unwelcome end to my field work for the year, but that just leaves me with more time now to process and share photographs from the past season and reflect on the stories behind them.  It will be welcome diversion, as I also begin the arduous and seemingly endless task of mounting and labeling the specimens collected during the past season – success’ bitter reward!

Copyright © Ted C. MacRae 2009

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A face only a mother could love

Centruroides vittatus - striped bark scorpion

Centruroides vittatus - striped bark scorpion

I found this guy yesterday secreted under a rock in a limestone glade at White River Balds Natural Area in extreme southwest Missouri.  Centruroides vittatus¹ is the most common scorpion in the U.S., occurring naturally in southern Missouri, western Arkansas and western Louisiana, west through Texas, Oklahoma and much of Kansas to southeastern Colorado and eastern New Mexico, and south into northern Mexico (Shelley and Sissom 1985). The Missouri and Mississippi Rivers, respectively, seem to form natural northern and eastern distributional boundaries, with occurrences just to the north and east attributed to rafting or natural alterations of the river’s courses and those occurring far outside the natural range regarded as the result of human introductions. This includes not only states in the eastern U.S., but several countries in South America (Sissom and Lourenco 1987)!

¹ The generic name Centruroides is from the Greek words centr-, meaning “pointed,” ur, meaning “tail,” and –oides meaning “like” or “the form of” (the original genus name, Centrurus, was preoccupied by another animal, thus, Centruroides, or “like Centrurus“). Centruroides is often misspelled as “Centuroides” in non-primary literature.

If you squit your eyes, it looks like he’s “smiling”! Also, note the eight eyes (two dorsal and three each side laterally)! Also, I know I didn’t nail the focus on the dorsal ocelli – depth-of-field limitations prevented getting both the ocelli and the jaws. I should’ve gone f/16 but thought I’d nailed it. That’ll teach me not to bracket anyway!

Photo details: Canon 65mm 1-5X macro lens on Canon EOS 50D, manual mode, ISO-100, 1/250 sec, f/14, MT-24EX flash 1/4 power through diffuser caps. Photo slightly cropped and darkened.

REFERENCES:

Shelley, R. M. and W. D. Sissom. 1985. Distributions of the scorpions Centruroides vittatus (Say) and Centruroides hentzi (Banks) in the United States and Mexico (Scorpiones, Buthidae). The Journal of Arachnology 23:100–110.

Sissom, W. D. and W. R . Lourenço. 1987. The genus Centruroides in South America (Scorpiones, Buthidae). The Journal of Arachnology 15:11–28.

 

Copyright © Ted C. MacRae 2009

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Do you know what time it is?

I’ll give you a few hints:

  • It’s fall.
  • I haven’t collected bugs for a few weeks.
  • A new crop of tiger beetles has emerged from their burrows.

What time is it?

IT’S TIME FOR THE ANNUAL FALL TIGER BEETLE TRIP!

 I don’t think there is any trip during the year that I look forward to more than this one. Hunting for insects is fun no matter what, but it is particularly enjoyable when the sweltering days of summer give way to the cool days of fall – crisp air, pungent, earthy aromas, shadows long and sharp, and skies so blue above a golden, tawny, morphing landscape. How I adore fall, and how I thrill at any chance to travel across the fall landscape chasing after gorgeous tiger beetles that have spent the spring and summer as larvae, hidden in their unseen burrows, growing fat on the few hapless insects that chanced too close to their burrow, until the rains of late summer and early fall trigger their transformation to adulthood – glittering jewels that emerge out into the autumn world for a brief session of dining and play before winter forces them back into their burrows for the long wait to spring.

This year’s edition is somewhat abbreviated – little more than a long weekend due to a combination of job and family responsibilities. Still, five days is a little better than four (and a lot better than none) and is long enough for me to play a hunch that I’ve had ever since I returned from the Red Hills of northwestern Oklahoma this past June. You’ll recall that I had some rather amazing luck on that trip, discovering a robust population of the very rare Cylindera celeripes (Swift Tiger Beetle) and a slight western range extension of the seldom seen Dromochorus pruinina (Frosted Dromo Tiger Beetle). While I was exploring that landscape, the habitat reminded me of another tiger beetle – Cicindela pulchra (Beautiful Tiger Beetle), a glorious species – brilliant purple with glassy wine-red elytra – that I had seen in 2005 in the nearby Gypsum Hills of south-central Kansas. I don’t know if that species occurs in the Red Hills of northwestern Oklahoma or not, but my impression is that the area has not been very well explored. If a species as rare as C. celeripes can be found there, perhaps C. pulchra will occur there as well. The enormous tiger beetle larvae that I saw in their burrows in the Gloss Mountains during June gives me further reason to believe there may now be some impressive adult activity in the area.

Should I not succeed in finding C. pulchra, it will nevertheless be a glorious, though frenetic trip. On Friday I’ll drive 525 miles from St. Louis to the Gloss Mountains, where I’ll explore during the early part of Saturday and then finish the day at Alabaster Caverns State Park. Sunday’s itinerary depends upon whether I succeed at finding C. pulchra in the Gloss Mountains – if I do, I’ll head on over to Salt Plains National Wildlife Refuge to photograph some of the fall tiger beetles that were not out during my June trip (hopefully including Eunota togata globicollis, or Alkali Tiger Beetle). If I don’t find C. pulchra in the Gloss Mountains, I’ll explore the Red Hills of Barber Co. Kansas, where I’ve seen this insect in the past and attempt to find and photograph it there, then move on to Salt Plains on Monday. I’m really hoping my C. pulchra hunch plays out, because if it does that gives me an extra day to shoot back east to my beloved White River Hills in southwestern Missouri and photograph its small, disjunct population of Cicindela obsoleta vulturina (Prairie Tiger Beetle) – the largest member of the genus in North America. Regardless of how events play out, I’ll need to blast back to St. Louis on Tuesday, work a couple of days, then leave town again for my niece’s wedding in New York (congratulations Shannon and Tamer).

While I’m gone, you can click on the interactive map to see where I’m going, or you can catch up on several newly issued Blog Carnivals (I’ve been a busy submitter this past month):

  • Circus of the Spineless. With discussion that is restricted to the 95% of life forms that do NOT have vertebrae, CotS #42 is up at Quiche Moraine.  For my part, I have proposed a replacement name for a rather ‘ubiquitous’ species of tiger beetle.
  • Berry Go Round.  After a brief summer vacation, botanical discussions resume with BGR #20 at Further Thoughts.  My contributions cover zygomorphic flowers with oily rewards, a very ungentianlike gentian, and plant-insect relationships.
  • Carnival of Evolution.  From Darwin to Drift to Deleterious Mutation, find it all at CoE #16 hosted by Pleiotropy.  I’ve added a little ‘perspective’ to the discovery of new species.

No longer just a contributor, at the end of this month I will host my first Blog Carnival in the form of Berry Go Round #21.  I know,  it’s strange that a bug dude is jumping into the Carnival hosting pool with a botanical carnival, but duty calls!  Submissions are due to me by Oct. 27, with a scheduled issue date of Oct. 30.  If you’ve never contributed to a Blog Carnival before, it’s a great way to get exposure for your blog and possibly find other blogs of interest.

Copyright © Ted C. MacRae 2009

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North America’s most beautiful longhorned beetle

I’ve written a few posts in recent weeks highlighting some of the more interesting finds encountered during two visits this past July to the White River Hills region of extreme southwestern Missouri. It’s a land of extremes, with deeply dissected layers of limestone/dolomite bedrock supporting xeric glades, dry woodlands and riparian watercourses. The hilltop glades (“balds”), in particular, feature prominently in the region’s natural and cultural history and are the most extensive system of such habitat in Missouri. They support a number of plants and animals more characteristic of the grasslands of the south-central U.S., such as the recently featured Megaphasma denticrus and Microstylus morosum, North America’s longest insect and largest robber fly, respectively. Sadly, the glades in this region are much reduced in size and quality compared to their pre-settlement occurrence, primarily due to overgrazing and suppression of fire. These anthropogenic forces have combined to reduce overall vegetational diversity and accelerate encroachment by woody species (chiefly eastern red-cedar, Juniperus virginiana). Nevertheless, there still remain several high quality glade remnants in the area, and the public agencies charged with their conservation are increasingly utilizing mechanical removal of woody growth, controlled burns, and managed grazing in an effort to simulate the natural forces that mediated this landscape for thousands of years.

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Chute Ridge Glade, Roaring River State Park, Barry Co., Missouri

My reason for returning to the White River Hills this year was simple—find and photograph the magnificent longhorned beetle, Plinthocoelium suaveolens (family Cerambycidae). This species, occurring across the southern U.S. from Florida and Georgia west to New Mexico and Arizona, is truly one of North America’s most beautiful longhorned beetles due to its large size, brilliant iridescent green coloration, and super-elongate wildly-contrasting orange and black legs.  Until recently, this species was known in Missouri only from sporadic records across the southern part of the state (MacRae 1994). I knew of its association with gum bumelia (Sideroxylon lanuginosum [= Bumelia lanuginosa], also called gum bully and woolly buckthorn), which was first noted by Missouri’s first State Entomologist, C. V. Riley (1880) and later discussed in detail by Linsley and Hurd (1959) and Turnbow and Hovore (1979); however, my repeated searches over the years whenever I encoutered this plant came up empty.  A few years ago, Chris Brown and I were conducting a survey of tiger beetles in the White River Hills and noted the relatively common occurrence of bumelia on these glades.  Bumelia, like P. suaveolens, is one of only a few North American representatives of a largely tropical group, and it is one of the few woody species naturally adapted to the xeric conditions found on these glades.  Recalling the association of P. suaveolens with this plant, and also recalling that adults could be attracted to fermenting baits of the type described by Champlain and Knull (1932), we placed fermenting bait traps on several glades in the area and succeeded in trapping a number of individuals during the month of July.  When I began searching the bumelia trees at these glades, I found adults perching on the lower trunks of several trees. It was the first time I’d seen live individuals of this species in Missouri.  At the time I was not a photographer, and that experience became one of the many moments that I would later look back upon and think, “If only I’d taken a picture of that!”  Thus, at the end of June this year, having successfully found Cylindera celeripes in Missouri on the first day of a planned 3-week search, my attention immediately turned to the new goal of finding P. suaveolens and photographing it on its host plant.

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Sideroxylon lanuginosum (gum bumelia) at Blackjack Knob, Taney Co., Missouri

I knew this wouldn’t be easy—the beetles were not abundant when I had last observed them, and those that I did find were quite wary to my approach.  Getting within striking distance with a net was one thing; doing so with a camera and macro lens would be another thing entirely.  In my first trip to the area (early July), I went to Chute Ridge Glade, a magnificently restored glade in Roaring River State Park where I had seen the greatest number of individuals before.  I was full of optimism on that first day as I zigzagged across the rough terrain from one bumelia tree to the next, but my optimism began to wane as I cautiously approached each tree and saw nothing.  Within an hour, I’d looked at every bumelia tree I could find on the glade and not even seen a beetle, much less attempted a photograph.  It would take a 2-hour drive along twisting back roads to reach the other sizeable glade complex where I had seen beetles before (Blackjack Knob in Taney County), and another hour of searching on several dozen trees would again yield nothing.  By now I was feeling rather frustrated—the day’s oppressive heat and humidity had taken its toll, and my 4.5-hour drive from St. Louis was looling like it would be for naught.  I had noted that the bumelia flowers were almost but not quite open yet—perhaps it was too early in the season still?  

Plinthocoelium suaveolens larval frass pile at the base of living Sideroxylon lanuginosa

Plinthocoelium suaveolens larval frass pile at trunk base of living Sideroxylon lanuginosum

The remnant glades at Blackjack Knob are more extensive than those at Chute Ridge, so many more trees still awaited examination—if I could only muster the energy!  I trudged back to the truck, guzzled a nice, cold Powerade, and started off in another direction.  I looked at a number of trees and still had seen no sign of the beetle, but on one particular tree I noticed an enormous pile of sawdust on the ground at the base of the tree.  I looked at it more closely and saw that it had the rough, granular texture so characteristic of longhorned beetle larvae that like to keep their galleries clean, and its bright, moist  color suggested that it was being ejected by a larva tunneling through living wood.  I looked up into the tree above the pile to find where it was coming from but could find no ejection hole.  I checked the base of the trunk itself and still couldn’t find anything.  Then I started poking into the pile and felt a root.  Further poking revealed a soft spot on the root, and I immediately knew that I had found a P. suaveolens larval gallery—no other cerambycid species is known to bore in roots of living Sideroxylon, especially one as large as this based on the size of the frass pile.  I hurried back to the truck and grabbed my hatchet, returned to the tree, and scraped away the soil above the root to find an obvious ejection hole a few inches away from the base of the trunk.  I started chipped into the root at the ejection hole and found a large, clean gallery extending down the center of the root away from the trunk.  About 18” away from the trunk I found it—a large, creamy-white cerambycid larva.

Plinthocoelium suavelones larva in root of living Sideroxylon lanuginosa

Plinthocoelium suaveolens larva in root of living Sideroxylon lanuginosum

Plinthocoelium suavelones larva in root of living Sideroxylon lanuginosa

Plinthocoelium suaveolens larva in root of living Sideroxylon lanuginosum

Finding a P. suaveolens larva was gratifying, but it wasn’t what I had come here to do, which was photograph the adult. After placing the larva live in a vial for preservation later on (dropping into scalding water to “fix” the proteins and prevent discoloration when stored in 70% ethanol), I continued searching the trees for adults.  I found one tree on which the flowers were just barely beginning to open and collected a few of the pedestrian species of scarabs that are attracted to bumelia flowers in droves when fully open (e.g. Cotinis nitidus and Trigonopeltastes delta)—for the record.  There was still no sign of adult Plinthocoelium, and I was on the verge of calling it a day when I approached another tree and saw it!  I froze, then slowly geared up with the camera and started stalking slowly towards it.  It was not in a very convenient location, down low on the trunk and partially screened by foreground vegetation.  I got close enough to start attempting some shots—not ideally composed, but just to ensure that I had something before I tried to get any closer.  After the third shot, however, it became alarmed and started to flee, and I had no choice but to capture it for a “studio backup.”  That taste of success gave me the motivation to resume my search, but no additional beetles were seen before a dropping sun put an end to the day.

Plinthocoelium suaveolens on lower trunk of living Sideroxylon lanuginosum

Plinthocoelium suaveolens on lower trunk of living Sideroxylon lanuginosum

Not entirely satisfied with the shots that I’d gotten, I returned to Blackjack Knob the following day and also searched some of the extensive habitat at nearby Hercules Glades Wilderness.  I wouldn’t see another beetle the entire day, although encountering a nice series of Cicindela rufiventris (red-bellied tiger beetle) was some consolation for suffering the day’s oppressive heat and humidity.  I still had the live beetle, so I placed my hopes on getting better photographs of the beetle in confinement after returning home.  That would not come to pass—the beetle refused to sit obligingly on the stick I placed in the large screen cage, and instead clung to the cage itself.  For days I watched it, giving it honey-water for sustenance and waiting for an opportunity to photograph it on the stick on which it refused to sit.  It became clear to me that studio photographs, at least in the manner I was attempting, would not be possible.  Not entirely satisfied with having seen only a single beetle on my trip, and thinking that I may have been too early based on the flowering phenology of the bumelia host trees, I did what any dedicated entomologist would do—I made a second trip to the area two weeks later!

I didn’t mess with Chute Ridge Glade this time, instead making a beeline for Blackjack Knob right away.  Unfortunately, the weather was uncooperatively drizzley (I would have preferred hot and humid to rain!).  Nevertheless, daughter Madison and I made our way to the glades and began inspecting the trees that I had just examined two weeks earlier.  I noted immediately that the bumelias were now in full flower, and it wasn’t long before I saw the first adult flying into these flowers.  Exciting for sure, and this was a good sign to see an active adult despite the drizzly weather, but the situation of the beetle on a high branch left no possibility for photographs (and only with a rather acrobatic swing of my fully extended net handle amidst a jumble of dead branches was I able to capture it).  This same scenario would replay several times over the next two hours before rain finally drove us back to the car.  In total, we saw half a dozen active adults, but in each case they were seen flying to flowers on high branches and could not be photographed.  Despite that disappointment, I’ll never forget the spectacularity of seeing these beetles in flight—shimmering green and bold orange, with legs and antennae spread wide in all directions.  I was also fortunate to find another tree with a fresh frass pile at its base indicating an active larva.  This time, I cut the tree some inches above the ground and extracted the trunk base and root intact for transplanting into a large soil box upon my return home.  The appearance of new frass on the soil surface afterwards confirmed that I had gotten the root containing the larva and that it had survived the extraction and transplanting.  Hopefully I will be able to successfully rear this individual to adulthood.

Despite the rain, we then went back to Hercules Glades Wilderness to see if luck would follow suite there as it had at Blackjack Knob.  It didn’t, as rain continued to doggedly pursue us, but the day was not a total loss as daughter and I got in a nice 7-mile hike through some of Missouri’s most ruggedly scenic terrain and were rewarded with the sighting of a western pygmy rattlesnake.  The next day was sunny, much to our delight, and I considered going back to Blackjack Knob where we had seen a good number of adults the previous day.  In the end, I decided I’d played that card and rather than continue trying for photographs I’d rather see if the beetle could be found at another glade complex further to the east at Long Bald Glade Natural Area in Caney Mountain Conservation Area.  Things didn’t look promising, as I found bumelia trees occurring only sporadically across the main glade complex—with no sign of the beetles.  Nevertheless, we enjoyed the day and spent a bit of time chasing after some enormous robber flies that later proved to be Microstylum morosum, a new record for Missouri and a significant northeastern range extension.  I thought that would be the highlight of the day, but as we were heading back to the car I spotted a small glade relict on the other side of the road.  It was overgrown and encroached, apparently not receiving the same management attention as the glades in the main complex. Regardless, I went over to check it out and immediately spotted several bumelia trees amongst the red-cedars, and within minutes I saw a beetle—low on the trunk of a very small bumelia tree!  Once again I froze, then slowly geared up with the camera and began my ultra-cautious approach (remember, this was only my second photo chance after a combined four days in the field).  Like last time, I took one shot while still some distance away, then moved in for closer attempts.  Unlike last time, there was no bothersome vegetation cluttering the view, and when I moved in for closeups the beetle turned around, crawled up the trunk a short distance, and then paused.  I snapped off a small series of shots while it sat there, and then suddenly it became alarmed and flew away.  Though still not perfect, these photographs were better than the previous ones I had obtained (check out the pronotal armature in the last photo!), and the finding of this species at Long Bald Glades also represented a new county record.

Plinthocoelium suaveolens on trunk of living Sideroxylon lanuginosum

Plinthocoelium suaveolens on trunk of living Sideroxylon lanuginosum

Plinthocoelium suaveolens on trunk of living Sideroxylon lanuginosum

Plinthocoelium suaveolens on trunk of living Sideroxylon lanuginosum

Missouri populations are assignable to the nominotypical subspecies (southeastern U.S.), which is distinguished from subspecies plicatum (Texas, New Mexico, Arizona, and northern Mexico) by the bronze or cupreous tints and weak transverse rugae on the pronotum (Linsley 1964).  The distributional ranges of the two subspecies intermingle in northeastern Texas.

Photo details:
All photos: Canon 100mm macro lens on Canon EOS 50D
Photo 1 (Chute Ridge Glade): normal mode, ISO-400, 1/250 sec, f/16, natural light.
Photo 2 (Sideroxylon lanuginosum): landscape mode, ISO-100, 1/160 sec, f/6.3, natural light.
Photos 3 (P. suaveolens larval frass pile), 6—8 (P. suaveolens adult): manual mode, ISO-100, 1/250 sec, f/9-11, MT-24EX flash 1/2 power through diffuser caps (photo 7 slightly cropped).
Photos 4—5 (P. suaveolens larva): manual mode, ISO-100, 1/60 sec, f/14 (closeup f/25), MT-24EX flash 1/2 power through diffuser caps.

REFERENCES:

Champlain, A. B. and J. N. Knull.  1932.  Fermenting bait traps for trapping Elateridae and Cerambycidae (Coleop.).  Entomological News 43(10):253–257.

Linsley, E. G. 1964.  The Cerambycidae of North America. Part V. Taxonomy and classification of the subfamily Cerambycinae, tribes Callichromini through Ancylocerini.  University of California Publicatons in Entomology, 22:1—197, 60 figs., 1 pl.

Linsley, E. G. and P. D. Hurd, Jr.  1959.  The larval habits of Plinthocoelium suaveolens plicatum (LeConte).  Bulletin of the Southern California Academy of Sciences 58(1):27–33.

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

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

Riley, C. V.  1880.  Food habits of the longicorn beetles or wood borers.  The American Entomologist 3(10):237–239.

Turnbow, R. H. Jr. and F. T. Hovore.  1979.  Notes on Cerambycidae from the southeastern U. S.  Entomological News 90(5):219–229.

Copyright © Ted C. MacRae 2009

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Assassin ate

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I came upon this interesting scene last month while hiking through Allen David Broussard Catfish Creek Preserve State Park, which preserves some of the highest quality remnants of sand scrub habitat on the Lake Wales Ridge of central Florida. The spider seems to be Peucetia viridans (green lynx spider), widespread across the southern U.S. and distinguished by its bright transparent green color with red spots and black spines (Emerton 1961). These largest of North American lynx spiders hunt diurnally on low shrubs with an agility excelled only by the jumping spiders (Salticidae) and aggressively attack their insect prey. In this case, the prey is one of the so-called “bee assassins” of the genus Apiomerus (Hemiptera: Reduviidae). The common and generic names of these insects both derive from their habit of preying upon bees, not only on flowers but also by ambushing them at nest entrances, although other insects are preyed upon as well. Ironically, this particular assassin himself got ate.

An interesting situation was uncovered while I tried to determine which species of Apiomerus was represented by the prey. By virtue of its pale ventrals with the front and hind margins black, it keys to A. spissipes in a literature-based key to Florida Reduviidae (Bierle et al. 2002) – one of two species considered widely distributed across the eastern U.S. In reality, however, it appears that this individual represents another species named almost 30 years ago but which remains officially undescribed. As explained in this BugGuide post by Daniel Swanson, the genus was revised by Berkeley grad student Sigurd Leopold Szerlip in partial fulfillment of the requirements for a Ph.D., who proposed a number of taxonomic acts including the description of 19 new species. Among these were eastern U.S. populations to which the name A. spissipes had been applied, with those in Florida being described as the new species “A. floridensis“. However, dissertations do not meet the criteria of publication according to Article 8 of the International Code of Zoological Nomenclature (ICZN 1999), and none of the dissertation was formally published. Thus, “A. floridensis” remains an invalid, unpublished name.  This is a most unfortunate situation, as Swanson considers the dissertation to be well done.  It is not only names, but important information about life histories and detailed genitalic studies that remain unavailable to the scientific community as well.  What are the nomenclatural impacts of this work remaining unpublished?  Is this as much a failure by the advising professor as by Szerlip himself?  What ethical considerations would need to be addressed in order for it to be published in absentia, or is this even possible?

Photo details: Canon 100mm macro lens on Canon EOS 50D (manual mode), ISO-100, 1/250 sec, f/13, MT-24EX flash 1/2 power through diffuser caps.

REFERENCES:

Bierle, S., E. Dunn, S. Frederick, S. Garrett, J. Harbison, D. Hoel, B. Ley and S. Weihman. 2002. A literature-based key to Reduviidae (Heteroptera) of Florida (assassin bugs, and thread-legged bugs). Unpublished manuscript, University of Florida, Department of Entomology and Nematolgy, Insect Classification ENY 4161/6166, 18 pp.

Emerton, J. H. 1961. The Common Spiders of the United States. Dover Publications, Inc., N.Y., xx + 227 pp.

International Commission on Zoological Nomenclature [ICZN]. 1999. International Code of Zoological Nomenclature, 4th Edition. The International Trust for Zoological Nomenclature, c/o Natural History Museum, London. xxix + 306 pp.

Szerlip, S. L. 1980. Biosystematic revision of the genus Apiomerus (Hemiptera: Reduviidae) in North and Central America. Unpublished Ph.D. thesis, University of California, Berkley, CA.

Copyright © Ted C. MacRae 2009

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North America’s largest robber fly

Female Microstylum morosum perched on fragrant sumac (Rhus aromatica) bush

Female Microstylum morosum perched on fragrant sumac (Rhus aromatica)

A few days ago, I featured Promachus hinei, one of the so-called “giant robber flies” and a common inhabitant of the glades and grasslands that dot Missouri’s largely forested landscape. That individual was seen at Long Bald Glade Natural Area in Caney Mountain Conservation Area, one of the many limestone glades that are a prominent feature of extreme southwestern Missouri’s White River Hills, as it snacked on a small carpenter bee (Ceratina sp.) and posed obligingly for a series of super close-up photographs. Promachus and its congeners are impressively large; however, I would see an even larger robber fly that day. I didn’t know what it was at the time, but I knew that never before had I seen such a magnificent fly, with its large, shimmering, emerald eyes, streamlined body almost devoid of setae (hairs), and ludicrously large size. These monsters were actually quite common at the glade, so I failed to appreciate the significance of what I was seeing as I chased one after another – more intent on securing photographs than specimens. This was not an easy task – they were extremely wary, rarely allowing me to approach within 12 feet no matter how cautiously and slowly I moved. Not one to back down from such a challenge (remember, I stalk tiger beetles), I persisted, traversing the rough, rock-strewn terrain amidst clumps of big bluestem (Andropogon gerardii), Indian grass (Sorghastrum nutans), and fragrant sumac (Rhus aromatica) until, at last, I got within striking distance of the impressive female shown in these photos. Taking flight before I felt assured of a good shot, I followed her repeated long, loping escape flights until I was able to get another few shots and she disappeared for good.

Same individual as in previous photo after flying to another perch.

Same individual as in previous photo after flying to another perch.

It didn’t take long after I returned home to figure out what I had seen, as there is really nothing that can be mistaken for Microstylum morosum, North America’s largest robber fly (Back 1909)¹. At 35–40 mm of length, this individual didn’t quite match the astounding 50-mm upper body length for the species (that’s 2 inches, folks!). Nonetheless, it was an impressive beast indeed! It is not surprising that North America’s largest robber fly should be a species of Microstylum, as it is this same genus that contains the world’s largest robber fly – the aptly named M. magnum from Madagascar, with a body length of 60 mm and an almost preposterous wingspan of up to 84 mm (that’s over 3 inches folks!). I don’t know if any flies exist that are larger than this, but certainly none can be more imposing.  While I’m happy with the photos that I did obtain, I must confess some disappointment that I wasn’t able to get more than these basic lateral profile shots.  Of the several photographs of this species that can be found on the web, this female, photographed by Greg Lavaty of Houston, Texas, is (in my humble opinion) certainly the most stunning.

¹ Puzzled by the use of the prefix “micro” in the genus name – hardly seeming appropriate for such an enormous fly – I asked Eric Fisher (retired, California Department of Food and Agriculture) about the name’s derivation, to which he replied, “The name refers to the quite small ‘stylus’ of the antenna apex; Macquart specifically mentions this character in his 1838 original description of the genus. (This is not a very helpful diagnostic character, as many asilids share this feature…).”

Even more significant than its size, however, was its very occurrence on this glade. Like Ospriocerus abdominalis, which I had seen just a few weeks earlier in the Loess Hills of extreme northwestern Missouri, M. morosum is a denizen of the Great Plains, and also like that species it has until now not been known from Missouri. That’s right – another new state record!  Unlike O. abdominalis, however, the Missouri occurrence of M. morosus represents a significant northeastern extension of its known range.  The species was long considered a Texas endemic until Beckemeyer and Charlton (2000) confirmed its occurrence in southeastern Arizona and documented significant range extensions into Oklahoma, Kansas, extreme southeastern Colorado, and extreme northeastern New Mexico.  Its eastern distributional limit was thought to occur along a north-south line from Douglas County, Kansas to Mayes County, Oklahoma to Brazoria County, Texas; however, Warriner (2004) documented its occurrence some 200 miles east of this line in the blackland prairies of southwestern Arkansas.  The occurrence of M. morosum in the White River Hills of Missouri represents yet another significant eastern extension of its known range – Long Bald Glade lies 185 miles NNE of the collection site in Arkansas and 155 miles ENE of the nearest known record in Mayes County, Oklahoma (Locust Grove), making it the easternmost known locality for this species.

As in Arkansas, where the collection site represents one of the highest quality blackland prairie remants in the state, Long Bald Glade represents a high quality remnant of the limestone glades that once occurrred much more extensively within Missouri’s White River Hills.  Like the blackland prairie of Arkansas, the limestone glades of the White River Hills have been dramatically reduced since EuroAmerican settlement due to land use conversion, and fire suppression and overgrazing of the remaining tracts have resulted in significant woody encroachment – chiefly by eastern red-cedar (Juniperus virginiana) – and loss of vegetational diversity. This has caused dramatic reductions in populations of the many Great Plains plant and animal species that are found here and nowhere else in the state.  Considering the overall distribution of M. morosum, it is unlikely that it occurs more extensively within Missouri than the White River Hills, emphasizing the importance of continued conservation and restoration activities in this unique part of Missouri.  However, since the White River Hills extend into northwestern Arkansas, M. morosum may occur in that part of Arkansas as well as the southwestern part of the state.

I thank Eric Fisher and Herschel Raney for confirming the identity of this species and its status as a new record for Missouri.

Photo details: Canon 100mm macro lens on Canon EOS 50D (manual mode), ISO-100, 1/250 sec, f/10-11, MT-24EX flash 1/2 power through diffuser caps.

REFERENCES:

Back, E. A. 1909. The robberflies of America, north of Mexico, belonging to the subfamilies Leptograstrinae and Dasypogoninae. Transactions of the American Entomological Society 35:137–400.

Beckemeyer, R. J. and R. E. Carlton.  2000.  Distribution of Microstylum morosum and M. galactoides (Diptera: Asilidae): significant extensions to previously reported ranges.  Entomological News 111(2):84–96.

Warriner, M. D.  2004.  First Arkansas record of the robber fly Microstylum morosum (Diptera: Asilidae).  The Southwestern Naturalist 49(1):83–84.

Copyright © Ted C. MacRae 2009

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