Featured Guest Photo – Dromica kolbei

Dromica kolbei? - Kruger National Park, South Africa. Copyright © Joe Warfel 2011.

Shortly after I returned from Brazil, this stunning photo was sent to me by Joe Warfel, who himself had just returned from a trip to South Africa.  Joe had seen the beetle at Punda Maria camp in the northern part of Kruger National Park, had deduced that it represented a species in the genus Dromica, and included the following notes about its behavior:

It did not fly, only ran, ran, ran, ran…. you get the picture.  But stopped briefly now and then to deposit eggs in the  soil.  My best guess from my limited tiger beetle references is Dromica sp.  Any help for identification you may give would be appreciated.

Although I have not collected this genus myself, I recognized it instantly as a member of such based on specimens and images I have seen.  Carabidae of the World contains fine images of a number of species in this genus, of which Dromica kolbei (W. Horn, 1897) seems to be a pretty good match.  However, more than 170 species are currently included in the genus, and while a modern revision is in progress (Schüle and Werner 2001; Schüle 2004, 2007), the bulk of the genus still remains to be treated.  As a result, this really should be considered as just a provisional ID.

Dromica is a strictly sub-Saharan African genus of tiger beetles whose included species are denizens of dry lands – savannahs, grasslands, open woodlands, and semideserts, and are generally absent in the moister, more heavily forested areas of western Africa.  Like a number of other tiger beetle genera, they have given up the power of flight to capitalize on their fast running capabilities.  This flightlessness and the strict association of adults with often short rainy seasons has led to both spatial and temporal isolation of numerous, localized populations of restricted geographical range.  This has no doubt contributed to the diversification of the genus across the mosaic of suitable habitats covering central, eastern, and southern Africa.  Schüle and Werner (2001) suggest that a good number of new species may still await discovery in the more remote or yet inaccessible areas of the countries of occurrence.  I had hoped to encounter these beetles (and also Manticora, or the giant African tiger beetles) during my visit to South Africa in 1999, but luck was not with me in this regard (although I did collect several fine specimens of the handsome Ophryodera rufomarginata (Boheman) and also a few species in the genera Cicindina and Lophyra).

My sincere thanks to Joe Warfel for allowing me to post his excellent photograph.  I featured photographs by Joe in an earlier post (A Tiger Beetle Aggregation), and his other photos can be seen at EighthEyePhotography (you must see this striking harvestman from Trinidad!).

REFERENCES:

Schüle, P. 2004. Revision of the genus Dromica. Part II.  The “elegantula-group” (Coleoptera: Cicindelidae). Folia Heyrovskana 12(1):1–60.

Schüle, P. 2007. Revision of the genus Dromica. Part IV.  Species closely related to Dromica albivittis (Coleoptera: Cicindelidae). African Invertebrates 48(2):233–244.

Schüle, P. and K. Werner. 2001. Revision of the genus Dromica Dejean, 1826. Part I: the stutzeri-group (Coleoptera: Cicindelidae). Entomologia Africana 6(2):21–45.

Copyright © Ted C. MacRae 2011 (text)

Answers to ID Challenge #5 – Artrópodes em casca de árvore morta

Dead tree in Campinas, Brazil

After checking into my hotel in Campinas, Brazil I couldn’t wait to start exploring the grounds to see what insect life I might be able to find.  Almost immediately, I encountered this dead tree in back of the hotel.  To a beetle collector, a dead tree is an irresistible draw – especially one that is still standing and with loosely hanging bark, as in this one.  I approached the tree, gave it a look up and down the trunk to see if any beetles or other insects might be found on the outer surface of the bark, and when none were seen began carefully peeling sections of the bark away from the trunk.  Out from beneath the first section darted a small, black lizard – it reminded me in general form of our North American fence lizards (genus Sceloporus), but honestly it darted so fast up the trunk that I didn’t get a good look at it (much less even the chance to attempt a photograph).  Peeling the bark further away from the wood revealed a goodly number of what I took to be beetle larvae, although they were unlike anything I’d ever seen before.  They were fairly good-sized – about 25mm in length, and although there are a number of beetle families whose larvae may be encountered under the bark of dead trees, there aren’t many with larvae of this size.

Coleopteran larva (Tenebrionidae?) under bark of dead tree.

Despite their odd appearance, their basic gestalt suggested to me that they might be something in the family Tenebrionidae (darkling beetles).  Sadly, the state of beetle larval taxonomy is far from complete, especially in the tropics, and given the extraordinary diversity of the order as a whole I knew it could be difficult to impossible to identify them.  This task was further complicated by the fact that I did not collect any voucher specimens.¹

¹ Insect collecting permits are required in Brazil and are exceedingly difficult to obtain.  Although enforcement is lax, a few unlucky foreigners have been caught and suffered tremendous inconvenience at the hands of notoriously unsympathetic authorities.  This being a business trip, I had no desire to tempt fate for the sake of a few larvae in a group I don’t even study.

Despite a millipede-like appearance, six legs and loose cluster of ocelli indicate its true identity.

After consulting all of the print and online resources at my disposal and failing to find a convincing match at even the family level, I began to second guess not only whether these were tenebrionids, but larvae or even beetles.  I’m not aware of any tenebrionids with larviform adult females, but such are common in the Lampyroidea.  That didn’t seem to fit, however, as the latter tend to be much more flattened and armored in appearance, and the round head is really unlike the elongate and narrow head so often seen in that group.  The actually began to wonder if it was even a beetle – most xylophagous beetle larvae are light-colored and rarely so heavily sclerotized, and the antennae are unlike the typical 3-segmented antennae seen with most xylophagous beetle larvae.  In fact, the antennae and the shape of the head actually reminded me of a millipede, but the obvious presence of six legs (and no more) made this untenable (even though 1st instar millipedes are hexapod, the large size of these individuals precludes them from being 1st instar anything).  Eventually, I could only conclude that they were coleopteran – possibly a larviform adult, but more likely larval.  As a last resort I sent photos to Antonio Santos-Silva, a coleopterist at the University of São Paulo.  Although he specializes in Cerambycidae, I reasoned this might be a fairly common species since I had found good numbers on a single tree in an urban area near São Paulo, and as such it might be something he would recognize.  Antonio quickly replied saying that he agreed it was the larva of a species of Tenebrionidae, with an appearance similar to the larvae of Goniodera ampliata (a member of the Lagriinae, formerly considered a separate family).  I’ve not been able to find photos of the larva of Goniadera or related genera, but these do bear a striking (if more glabrous) resemblance to these presumed tenebrionid larvae from Australia.  Until a more convincing opinion is forthcoming, Tenebrionidae seems to be the consensus.

Polyxenid millipedes and two types of Collembola (several Poduromorpha and one Entomobryomorpha)

Three tiny adult coleopterans (family?) surround a large larval coleopteran

Although nobody zeroed in on Tenebrionidae for this challenge (#5 in the ID Challenge series), I must say that I enjoyed the diversity of opinion about what it might represent.  Moreover, congratulations to those who ‘took nothing for granted’ and noted the presence of several other organisms in the photo – this is where the big points were to be earned, and several participants successfully ID’d what I take to be a number of poduromorph collembolans, a single entomobryomorph collembolan, a central cluster of polyxenid millipedes, and several indistinct but clearly coleopteran adults (see super crops above).  David Hubble got the most correct answers to earn 15 points and the win in this inaugural post of BitB Challenge Season #2, while Dave and Troy Bartlett earned 13 and 10 points, respectively, to complete the podium.  Seven other participants got in on the fun and earned some points – I hope you’ll join the fun next time, too!

Copyright © Ted C. MacRae 2011

Recent literature – The Coleopterists Bulletin

Volume 64, No. 4 - December 2010

I returned to the office this week after spending two weeks in Brazil to find the December 2010 of The Coleopterists Bulletin in my inbox.  I don’t think there is another journal that I look forward to more eagerly than this one (with the possible exception of CICINDELA) – with each issue, I know that regardless of whether it contains any papers in my priority groups of interest (jewel beetles, longhorned beetles, and tiger beetles), it will nevertheless contain well-written articles presenting results of high-quality research on nothing but beetles – pure elytral ecstasy!  This latest issue, however, is a real keeper.  Gracing the cover is a stunning image of the South African jewel beetle species Julodis viridipes, photographed by my good friend and world jewel beetle expert Chuck Bellamy, and inside are three tiger beetle papers and two longhorned beetle papers – it’s almost as if the issue were produced just for me.

Friend and colleague Mathew Brust takes credit for two of the three tiger beetle papers.  In the first (Brust et al. 2010), he compares the efficiency of the three main methods for collecting tiger beetle larvae: fishing, stab-and-grab, and excavation. They found fishing to be the most efficient and least damaging of the three methods, an important finding for tiger beetle conservation where the availability of efficient, non-lethal survey methods is critical.  (What are “fishing” and “stab-and-grab” you ask?  Read the paper.)  In the second paper (Brust and Hoback 2010), Matt teams up with University of Nebraska entomologist Wyatt Hoback to provide new distributional records and larval descriptions of Nebraska’s tiger beetle, Cicindela nebraskana.  Ironically, this species is quite rare in it’s namesake state, and their findings give clues about the habitats in which it is most likely to occur (I had the good fortune to contribute a small amount of data).  In the third tiger beetle paper, Robert Richardson (2010) notes an overwhelming preponderance of left-superior mandibles in two species of Omus and speculates on the selective pressures that might operate on different tiger beetle clades to produce such a finding – a truly interesting read.

As for longhorned beetles, Sánchez-Martínez et al. (2010) report the utilization of living oaks by an apparently disjunct population of the marvelously beautiful Crioprosopus magnificus in central Mexico, complete with photographs of the larval workings and detailed emergence data.  (I am also reminded that I have a post on this very species that I need to put up).  Closer to home, Terence Schiefer and Patricia Newell (2010) independently recognized the existence of an undescribed subspecies of the red-edged saperda, Saperda lateralis, in the lower Mississippi Delta Region of the southeastern U.S., which together they describe as S. lateralis rileyi (named for Edward Riley, indefatigable collector of beetles, and collector of much of the type material).  Okay, I know what you’re thinking – “A new subspecies of S. lateralis? No way!”  Well, I was skeptical also when I first saw the title – several untenable and no longer recognized subspecies have already been described for this broadly distributed and variable species.  However, after noting the nature of the diagnostic characters, seeing the photographs, and studying their partially allopatric distributions, I was convinced.

In addition to the above papers, there were also a number of interesting book reviews in the issue, including The Chiasognathinae of the Andes, reviewed by M. J. Paulsen, A Field Guide of the Dynastidae Family of the South of South Americano access, reviewed by Ronald M. Young, and three book reviews by our beloved Art Evans: Illustrated Identification Guide to Adults and Larvae of Northeastern North American Ground Beetles (Coleoptera: Carabidae) [including tiger beetles]; The African Dung Beetle Genera; and Weevils of South Carolina (Coleoptera: Nemonychidae, Attelabidae, Brentidae, Ithyceridae, Curculionidae)no access.

REFERENCES:

Brust, M. L. and W. W. Hoback. 2010. Larval description and new Nebraska distribution records for Nebraska’s tiger beetle, Cicindela nebraskana Casey (Coleoptera: Carabidae). The Coleopterists Bulletin 64(4):341-346.

Brust, M. L., W. W. Hoback, and J. J. Johnson. 2010. Fishing for tigers: A method for collecting tiger beetle larvae holds useful applications for biology and conservation. The Coleopterists Bulletin 64(4):313-318.

Richardson, R. K. 2010. Mandibular chirality in tiger beetles (Carabidae: Cicindelinae). The Coleopterists Bulletin 64(4):386-387.

Sánchez-Martínez, G., O. Moreno-Rico, and M. E. Siqueiros-Delgado. 2010. Crioprosopus magnificus Leconte (Coleoptera: Cerambycidae) in Aguascalientes, Mexico: Biological observations and geographical distribution. The Coleopterists Bulletin 64(4):319-328.

Schiefer, T. L. and P. Newell. 2010. A distinctive new subspecies of Saperda lateralis F. (Coleoptera: Cerambycidae) from the southeastern United States. The Coleopterists Bulletin 64(4):329-336.

Copyright © Ted C. MacRae 2011

Brazil Bugs #3 – Gorgulho Enorme!

The second night at the hotel on the outskirts of Campinas (São Paulo, Brazil), I found this enormous weevil laying on the ground underneath some windows.  It was dead but completely relaxed and in perfect shape.  I wondered if it had been attracted to lights in the window the previous evening and flown there as its “last hurrah.”  This beast of a weevil – measuring a good 30mm from the tip of the snout to the apex of the elytra – immediately brought to my mind giant palm weevils of the genus Rhynchophorus (Coleoptera: Curculionidae).  I looked around and saw palm trees everywhere, so I figured that must be what it was.  I picked up one of the dead fronds from under a nearby palm tree and placed the weevil on the stem for a few photographs.  Given it’s freshness, it was quite easy to place in a lifelike pose, and I placed the stick in various positions to get a few different backgrounds.  As I was shooting, I noticed another, smaller, weevil on the ground near where I had found the first one – also dead and fresh.  Considering that Rhynchophorus often reaches pest status on palms, this further confirmed the ID in my mind.

Once back in the hotel room I smugly entered “giant palm weevil” into Google to bring up some images and confirm my ID (and also make sure my photos were at least on par with published images).  When the images came up, I saw immediately that my weevil was not a giant palm weevil at all.  While the South American species (R. palmarum) is black, the pronotum and elytra are much smoother and distinctly striate.  I don’t know if this weevil is a member of the same subfamily (Rhychophorinae) as the giant palm weevils, but it certainly not a member of that genus.  Still, considering its large size and distinctive appearance, I remained confident that an ID would come quickly.  Sadly, this has not proven to be the case – no search term I’ve tried involving the term “giant black weevil Brazil” or its many variations brings up anything remotely similar.  So, after spending much more time trying to identify this beetle than I should have, I leave it in the hands of my readers in the hopes that one of you will recognize it and provide a name.  Only when we have a name will it be possible to know something more about how this beetle lives (and if my placing it on a dead palm frond was a truly artificial setup).

Photo Details: Canon 50D w/ 100mm macro lens (ISO 100, 1/200 sec, f/16-18), Canon MT-24EX flash w/ oversized DIY concave diffuser. Typical post-processing (levels, minor cropping, unsharp mask).

Update 01/24/11: Thanks to Henry Hespenheide (via Facebook) and Art Evans, who clued me in to the genus Homalinotus (subfamily Cholinae).  I found a key to the 21 known species of this exclusively Neotropical genus (Vaurie 1973) and consider this individual to represent H. coriaceus – broca do cacho do coqueiro (black coconut bunch weevil).  This species breeds in the stems of old leaves of several native palms; however, it has adapted to the introduced coconut palm and apparently become a limiting factor in the commercial production of coconuts.  Thus, I can breathe easy now knowing that my photographs of this individual on the stem of a dead palm frond represent an entirely natural setting.

REFERENCE:

Vaurie, P.  1973.  The weevil genera Homalinotus and Ozopherus of the Neotropical Cholinae (Coleoptera, Curculionidae).  Bulletin of the American Museum of Natural History 152(1):1–49.

Copyright © Ted C. MacRae 2011

Brazil Bugs #2

A few more photographs from this past week in Campinas, Brazil.  It rained during the afternoon but stopped by the time I arrived back at the hotel, allowing me to stroll the lavishly landscaped grounds during the mild evening hours.  There is a pink-flowered shrub forming a hedge row in back of the hotel that is highly attractive to many types of insects.  The identity of the shrub remains a mystery to me, and most of the insects I’m finding on it I can recognize only to family – I’m hoping the hotel staff will be able to name the former and that the readers of this blog might be able to provide IDs for the latter.

Calycopis sp. poss. origo (Lepidoptera: Lycaenidae). ID by Dave Hubble and Chris Grinter.

It took a bit of effort to find an unobstructed view of this hairstreak butterfly (family Lycaenidae) as it visited the flowers within the shrub.  Every time I tried to move foliage out of the way to get a good view, the butterfly became alarmed and flew to another part of the hedge row.  My antics drew the attention of a hotel worker, who was apparently interested enough in what I was doing to act as a spotter whenever the butterfly flew to help me relocate it.  Eventually I got a few shots that I was happy with, including the above.

A flesh fly (Diptera: Sarcophagidae).

I presume this to be a type of flesh fly (family Sarcophagidae) based on the stout bristles and color pattern that seems typical for the family.  I like the striking contrast in coloration between the fly and the flower.  There are a few fly bloggers who I’m hoping might be able to give a better identification.

A potter/mason wasp? (Hymenoptera: Vespidae).

This appears to me to be some kind of potter or mason wasp (family Vespidae, subfamily Eumeninae) – it was a bit smallish at only about 12mm in length.  I hope one of the knowledgeable wasp bloggers out there (ahem… Eric?) can at least confirm this level of identification and perhaps the tribe or genus as well. 

Azya orbigera (Coleoptera: Coccinellidae). ID by Tucker Lancaster.

Every ladybird beetle (family Coccinellidae) I’ve ever seen is some variation of black and red/orange/yellow and has a smooth, glabrous appearance.  This beetle is cobalt blue with a dense pubescence over the dorsal surface, but it still seems to me to be some type of ladybird beetle.  It was a tiny little thing, so I suppose it could be one of the multitude of small beetle families with which I am unfamiliar.

Quedas sp.? (Hemiptera: Cicadidae).

This cast cicada exuvium was not on the shrub, but on a nearby tree at about eye level.  I really wish I could have seen the cicada that emerged from it, because this is certainly the biggest cicada exuvium I have ever seen.  I was about to simply label it “family Cicadidae” but seem to recall that cicada higher classification is in a bit of flux these days.  At any rate, given its great size I wonder if it might represent one of the giant cicadas in the genus Quesada.

I still have many more insect photographs from the past week and will certainly increase that number over the next week as well.  Stay tuned!

Copyright © Ted C. MacRae 2011

Calm waters, frenzied beetles

North Fork River - Ozark Co., Missouri

The North Fork River in south-central Missouri, like most Ozark rivers and streams, flows clear and cold over gravelled bottoms. Sustained year-round by the numerous seeps and springs that result from the region’s unique Karst geology, it meanders through a mix of forest and woodland alongside massive bluffs of half-a-billion-year-old dolomite. While small rapids can be found where gravel bars approach the bluffs, for the most part the shallow waters course lazily and idyllically south toward the White River in northern Arkansas.

Dineutus sp. (poss. discolor, per Brady Richards)

Lazy waters are the domain of whirligig beetles (family Gyrinidae).  We encountered this ‘raft’ of beetles in a sheltered pool near the shore of the North Fork River while hiking the Ozark Trail last October.  These frenzied little beetles live almost exclusively on the surface of the water, where they feed on organisms or scavenge debris in their famously and erratically conspicuous aggregations.  Such behavior might make them seem vulnerable to predation, but in actuality the reverse is true.  Beetles in rafts benefit from the increased number of eyes that can better scan the environment for potential threats than can individual beetles (Vulinec and Miller 1989), and the larger the raft the more efficiently this occurs.  There is also evidence that the appearance of the rafts themselves is a signal to warn potential predators (primarily fish) of the noxious chemicals produced in the beetles’ paired pygidial glands (Ivarsson et al. 1996), despite the decidedly non-aposematic coloration of the beetles themselves.

Photo Details: Canon 50D w/ 17-85mm zoom lens, natural light. Photo 1 – 17mm, ISO 100, 1/25 sec, f/5.6; photo 2 – 85mm, ISO 500, 1/160 sec, f/5.6. Typical post-processing (levels, minor cropping, unsharp mask).

REFERENCES:

Ivarsson, P., B.-I. Henrikson and J. A. E. Stenson.  1996.  Volatile substances in the pygidial secretion of gyrinid beetles (Coleoptera: Gyrinidae).  Chemoecology 7(4):191–193.

Vulinec, K. and M. C. Miller. 1989. Aggregation and Predator Avoidance in Whirligig Beetles (Coleoptera: Gyrinidae). Journal of the New York Entomological Society 97(4):438–447.

Copyright © Ted C. MacRae 2011

Diversity in Tiger Beetle Larval Burrows

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Copyright © Ted C. MacRae 2011

Monroe Canyon epilogue – Audubon’s tiger beetle

Cicindela purpurea audubonii (Audubon's tiger beetle) - green morph

In my first post about Monroe Canyon in the Pine Ridge of northwestern Nebraska, I featured the sumptuous Cicindela lengi (blowout tiger beetle) – a target species for the trip and one of six tiger beetle species that Chris Brown and I would find at this quarter-mile long sandy roadside embankment.  Another species we found there was C. denverensis (green claybank tiger beetle), unexpected given its preference for clay soils, but like C. lengi also a target species and thus a welcome find.  We also saw some more common species – the nominotypical forms of C. formosa (big sand tiger beetle) and C. scutellaris (festive tiger beetle), both reliable residents of sand habitats throughout the Great Plains, as well as a few individuals of the often ubiquitous C. punctulata (punctured tiger beetle).  The most numerous of all, however, was C. purpurea audubonii (Audubon’s tiger beetle).  Despite being uncommon in other parts of its range (in fact, nominotypical populations are feared extirpated in some parts of the eastern U.S.), C. purpurea audubonii is one of the most commonly encountered tiger beetles in grassland habitats throughout the central Great Plains and Rocky Mountains.  This species belongs to a complex of several that are generally green in color and fond of clay soil habitats, such as C. denverensis, C. decemnotata (badlands tiger beetle), C. limbalis (common claybank tiger beetle), and C. splendida (splendid tiger beetle).  All of these species can co-occur with C. purpurea audubonii in the Great Plains, but the latter is distinguished by its faint purple tinge and distinct metallic purple border around the edge of the elytra and its reduced elytral markings consisting of a short, oblique middle line and a white rear tip at the edge of the elytra (Pearson et al. 2006).  All of these characters can be seen easily in classic pose in the above photograph.

Cicindela purpurea audubonii (Audubon's tiger beetle) - black morph

Despite its commonness, however, I actually did look forward to seeing this species – the reason being the occurrence of occasional all-black individuals in the population.  Tiger beetles as a whole are a variable lot – polytopism (geographically based variability) is the rule!  Despite this, for the most part individuals within a given population usually exhibit a fairly uniform appearance.  Occasionally, populations of some species – generally those at intergrade zones where different subspecies meet – will show variation on a continuum between two extremes.  The occurrence of two distinct morphs within a population, however, is rather unusual.  I lacked good field photographs of the all-black morph of C. purpurea audubonii (and also the green morph, for that matter), so I was pleased to encounter several individuals of the black morph while we were at Monroe Canyon.  As can be seen in the above photograph, black morph individuals are truly all-black, perhaps with a purple reflection but without a trace of green anywhere on the body.  They do retain the same pattern of reduced white markings exhibited by the green morphs.  As a result, these individuals can be confused with some other black species that also occur in the Great Plains, such as C. nebraskana (prairie long-lipped tiger beetle) and, at higher elevations, C. longilabris (boreal long-lipped tiger beetle).  These latter species were also targets for our trip, so we had to pay close attention to any black morphs that we saw to confirm their identity (C. purpurea audubonii black morphs are distinguished by their distinctly hairy frons).

Chris Brown waits patiently to photograph a burrowing wolf spider at the entrance of its burrow.

I’ve not been able to find any additional information about these black morph individuals and the possible causes for their occurrence.  Cicindela purpurea is a so-called “spring-fall” species, emerging in the fall as sexually immature adults and then spending the winter in burrows before re-emerging in the spring to mate and lay eggs.  As spring-fall species go, it is one of the earliest to appear in the spring and last to disappear in the fall.¹  This immediately brings to my mind a possible thermoregulatory function.  Low temperatures may be a challenge for the adults during early spring and late fall, and the black coloration could be an adaptation to maximize absorption of solar radiation for heat gain. This idea seems to be supported by the fact that the incidence of black morphs is greater at more northern latitudes and in the higher elevations of the western part of the subspecies’ range (as much as 20-40%), where overall lower temperatures would be expected to occur.

¹ Karl Werner even amusingly stated that this species “rather delights in chilly weather” (Acorn 2001).

REFERENCES:

Acorn, J.  2001. Tiger Beetles of Alberta: Killers on the Clay, Stalkers on the Sand. The University of Alberta Press, Edmonton, xix + 120 pp.

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 2011