Sand Prairie Conservation Area

I have a love-hate affair with Missouri’s Southeast Lowlands (formally known as the Mississippi River Alluvial Basin, but simply called the “bootheel” by most folk in reference to the shape of its boundaries). Of the four main physiogeographic regions in the state, it is by far the most altered. Yes, the Ozark Highlands have been degraded by timber mismanagement, overgrazing, and fire suppression, yet many of its landscapes nevertheless remain relatively intact – just a few burn and chainsaw sessions away from resembling their presettlement condition. The northern Central Dissected Till Plains and western Osage Plains are more disturbed, their prairie landscapes having been largely converted to fields of corn, soybean, and wheat. Still, riparian corridors and prairie habitats ranging from narrow roadsides to sizeable relicts combine to provide at least a glimmer of the regions’ former floral and faunal diversity. The alterations these regions have experienced are significiant, yet they pale in comparison to the near-total, fence-row-to-fence-row conversion that has befallen the Southeast Lowlands. Its rich, deep soils of glacial loess, alluvial silt, and sandy loam originally supported vast cypress-tupelo swamps and wet bottomland forests – massively treed and dripping with biotic diversity. Exposed by relentless logging and an extensive system of drainage ditches and diversion canals, those same soils now support monotonous expanses of soybean, wheat, rice, and cotton. Giant plumes of dark smoke dot the unendingly flat landscape in late spring, as farmers burn wheat stubble in preparation for a double-crop of soybean (the need for which could be obviated by adopting more environmentally benign no-till drillers). Only a tiny fraction of the original swamp acres remain intact, preserved more by default due to their defiant undrainability than by human foresight, and wet bottomland forests now exist only as thin slivers hemmed in by levees along the Mississippi River to the east and the St. Francois River to the west. Solace is hard to find in these remaining tracts – hordes of mosquitoes and deer flies, desperate for blood to nourish their brood, descend upon anyone who dares to enter their realm, while impoverished locals leave behind waste of all manner in their daily quest for fish. The cultural history of the region parallels its natural history – nowhere in the state is the gap between wealth and poverty more evident, a testimony to its checkered history of race and labor relations.

Yet, despite its shortcomings, I am continually drawn to this region for my explorations. Driving down the southeastern escarpment of the Ozark Highlands into the Lowlands is like entering another world – a world of grits, fried catfish, and sweet tea, a world where it is odd not to wave to oncoming vehicles on gravel back roads, a world where character is judged by the subtleties of handshake, eye contact, and small talk. Again, its natural history follows suit, with many insects occurring here and nowhere else in Missouri – a distinctly Southern essence in an otherwise decidedly northern state. My recent discussion of Cicindela cursitans in the wet bottomland forests along the Mississippi River is just one example of the unique gems I have encountered in this region. Others include the rare and beautiful hibiscus jewel beetle (Agrilus concinnus), a sedge-mining jewel beetle (genus Taphrocerus) that is new to science (and, due to my sloth, still awaiting formal description), the striking Carolina tiger beetle (Tetracha carolina), and numerous other beetle species not previously recorded from the state. The small and scattered nature of the habitat remnants and often oppressive field conditions make insect study challenging here, but the opportunity for discovery makes this region irresistible.

Prior to this season, I had already visited most of the publicly-owned examples of swamp and forest found in the Southeast Lowlands. One natural community, however, that I had not yet seen happened to be one of Missouri’s rarest and most endangered – the sand prairie (I suppose you’ve surmised this by now from the photos). While conducting our recent survey for Cicindela cursitans, I took the opportunity to explore a recently acquired example called Sand Prairie Conservation Area. Geologically, sand prairies lie on our state’s youngest landscape, arising during the relatively recent Pleistocene glacial melts. Tremendous volumes of water from the melting glaciers scoured through loose sands and gravels deposited earlier during the Cretaceous and Tertiary periods by the present day Ohio River (the Mississippi River, much smaller at that time, actually drained northward into Hudson Bay!). After the last of these glacial melts formally ended the “ice age” (only 10,000 years ago), two long sandy ridges were all that remained of the original sand plain. Water drains quickly through the sandy soil of these ridges, which lie some 10 to 20 feet above the surrounding land, creating dry growing conditions favorable for prairie and savanna habitats where only drought-tolerant plants can survive. Dr. Walter Schroeder has conservatively estimated that 60 square miles of sand prairie were present in the Southeast Lowlands at the time of the original land surveys. Because settlement was already occurring at that time, a substantial amount of sand prairie had already likely been converted to agriculture, urban centers, and travel routes to staging areas for access across the swamps. Considering the conversion that might have already taken place, it is possible that as much as 150 to 175 square miles of sand prairie occupied the sand ridges. Sandy areas with higher organic soil content and supporting tallgrasses would have been the first to be converted, since this organic content would have also made them the most suitable for agriculture. Those with lower organic content created drier conditions more suitable for shortgrasses and were the next to be converted. Today, less than 2,000 acres of sand prairie remain – not even 1% of the original amount, and these relicts likely represent the sandiest (and driest) examples of the original sand prairie.

Walking onto the site, I was immediately greeted by an otherworldly expanse of sand dunes, blows, and swales. Ever the entomologist, and with tiger beetles in the fore from hunting C. cursitans, I immediately thought of two dry sand associated species that I have seen in the sand woodlands of nearby Crowley’s Ridge – Cicindela formosa (big sand tiger beetle) and Cicindela scutellaris (festive tiger beetle). These are both so-called “spring-fall” species – i.e., adults are active primarily during spring and fall, so I thought it might be a little late (my first visit was in late June) to see either one. It wasn’t long, however, before I scared up a C. formosa (pictured – but unfortunately facing the setting sun) on one of the dunes. I also encountered one individual of another dry sand associated species, Cicindela lepida (a white “summer” species aptly named ‘ghost tiger beetle’) but was not able to photograph it (I have to say this – I’m a patient man, but photographing tiger beetles is hard. Actually, stalking them until you can get close enough to photograph them is hard. Stalking them until you can get even closer to photograph them with a ‘point and shoot’ – hoping and praying they settle into a pose with the sun on their back because you can’t use the blindingly dinky little built-in flash – just about breaks every last fiber of patience I have within my soul!). Though the site represents a new county record for both species, this is not unexpected, since we have recorded each at multiple dry sand sites near big rivers throughout the state. The occurrence of C. scutellaris at this site, on the other hand, would be significant, and though I did not find it on these two summer visits, I will certainly return this fall to have another look. Cicindela scutellaris has been recorded from just three widely separated locations in the state. Individuals from the two northern Missouri sites are assignable to the more northerly and laterally maculate subspecies C. scutellaris lecontei, but those from the Crowley’s Ridge population (some 20 miles to the west) show an intergrade of characters between C. s. lecontei and the more southerly all-green and immmaculate subspecies C. scutellaris unicolor. I should mention that I believe the classic definition of subspecies (i.e., allopatric populations in which gene flow has been interrupted by geographic barriers) has been grossly misapplied in Cicindelidae taxonomy, with many “subspecies” actually representing nothing more than distinctive extremes of clinal variation. Nevertheless, I am anxious to see if C. scutellaris does occur at Sand Prairie, and if so does it exhibit even more of the “unicolor” influence than does the Crowley’s Ridge population?

I’ve mentioned previously my weakness as a botanist, a fact I found especially annoying as I explored this new area and found myself unfamiliar with much of the flora that I encountered. I’ve taken photographs and will, over time, attempt to identify them. Still, some plants are unmistakeable, such as this clasping milkweed (Asclepias amplexicaulis, also known as sand milkweed) – unfortunately well past bloom. Asclepias is a favorite plant genus of mine (I’ve made it a personal goal to locate all 16 of Missouri’s native Asclepias), so you can imagine my delight when I encountered numerous robust green milkweed (Asclepias viridiflora) plants in full bloom. As I approached one of these plants, I noticed the unmistakeable form and color of a milkweed beetle (genus Tetraopes). It didn’t have the look of the common milkweed beetle (Tetraopes tetrophthalmus), which is widespread and abundant throughout Missouri on common milkweed (Ascelpias syriaca), and as soon as I looked more closely, I recognized it to be the much less common Tetraopes quinquemaculatus. Additional individuals were found not only on A. viridiflora, but also on A. amplexicaulis. The latter is also a suspected host (the larvae are root borers in living plants) in other parts of the species’ range, but in Missouri I’ve found this species associated only with butterfly weed (Asclepias tuberosus). These observations suggest not only that A. viridiflora may also be utlized as a host, but that three species of milkweed are serving as such in this part of the state – unusual for a genus of beetles in which most species exhibit a preference for a single milkweed species in any given area. More questions to answer!

Amazingly, there were no publicly owned representatives of this community type in Missouri until just recently, when the Missouri Conservation Department acquired Sand Prairie CA through the efforts of the Southeastern Sand Ridge Conservation Opportunity Area, a consortium of private and public agencies dedicated to the conservation and restoration of sand prairies in the Mississippi River Alluvial Basin. Restoration efforts are now underway to promote species that historically occupied native sand prairies on the Sikeston Sand Ridge. Fire is one such management tool, although there seems to be some debate about the role of fire in the history of this natural community. Some have argued that the Southeast Lowland sand prairies are an anthropogenic landscape, created by Native Americans who regularly cleared and burned the land after arriving in the Mississippi River Alluvial Plain. Had it not been for such intervention, the sand ridges communities would have remained sand woodlands and forests, dominated by hickories and oaks. Several lines of evidence – convincingly summarized by Allison Vaughn in “The Origin of Sand Prairies” (June 2008 issue of Perennis, Newsletter of the S.E. Missouri Native Plant Society) – suggest a more natural origin. These include the presence of rare sand prairie endemics that do not occur in the sand woodlands of nearby Crowley’s Ridge and the fact that the remaining sand prairie relicts have not succeeded back to sand woodland despite 150 years of post-settlement fire suppression. Perhaps the truth lies somewhere in between, with the driest prairies remaining open regardless of fire, while those with somewhat higher organic content in their soils supported shifting mosaics of prairie, savanna, and woodland as fire events (whether natural or anthropogenic) flashed across different areas. Regardless of their history, the sand prairies of the Southeast Lowlands are truly unique communities that deserve protection. Restoration efforts are well underway at Sand Prairie CA, as evidenced by the charred grass clump next to eastern prickly pear (Opuntia humifusa) in the above photo. There is still more work to do, however, as illustrated by this attractively scenic, yet unfortunately exotic Persian silktree (Albizia julibrissin) still remaining on the parcel – emblematic of Man’s pervasive alterations in even the most unique of landscapes.

For further reading on the sand prairies of the Southeast Lowlands, I recommend the excellent article, “A Prairie in the Swamp”, by A. J. Hendershott and this blog entry by the ever-eloquent author of Ozark Highlands of Missouri. In the meantime, so as not to disappoint the botanists who may stumble upon this silly post, I leave you with a few photographs of some of the wildflowers I saw during my visits. I consider the plant in the first photograph to be camphorweed (Heterotheca sp., either camporum or subaxillaris), frequntly associated with sandy soils in southern Missouri (especially the Southeast Lowlands). My colleague James informs me the second plant is plains puccoon (Lithospermum caroliniense), another sandy soil associate found primarily in the Lowlands and distinguished from the much more common L. canescens by its robustness and rougher pubescence. Both of these species were common near the perimeter of the barren sand areas and nearby. The third plant appears to be spotted beebalm (Monarda punctata) (my thanks to michael for the ID). It was confined, as far as I could tell, to a small area in a swale (moister?) away from the barren sand. This plant, a clump-forming perennial that prefers prairies and open sandy soils, is apparently not common in Missouri, having been found primarily in a few eastern counties adjacent to the Mississippi River.

Cicindela cursitans in Missouri

For several weeks now my colleagues and I have been immersed in surveys for selected tiger beetle species in Missouri. We placed 50 pitfall traps in western Missouri and 75 in the southeastern lowlands in mid-June and have been checking them weekly for several weeks now. It’s been a frenetic schedule for all of us – working regular jobs all week and covering two different parts of the state during weekends. Add to that spouses, children, and the desire to watch television coverage of two little sporting events called Wimbledon and the Tour de France, and you have the makings of a severe case of sleep deprivation.

Even with such a focused, dedicated effort success is not assured. Our previous work over the past several years has generated copious data on the more common, widespread species of tiger beetles occurring in the state. The distributions and habitats of these species are well documented now – the low hanging fruit has been picked. We’re now focusing on those few, rare species that I talked about in a previous post – the critically imperiled Cicindela circumpicta johnsonii, found in the equally critically imperiled saline spring habitats of central Missouri; Cicindela pruinina in western Missouri, normally associated with grasslands habitats further west; and the enigmatic Cicindela cursitans, until last year known in Missouri from just a single specimen collected somewhere “nr. Portageville” in the Mississippi lowlands of extreme southeast Missouri.

While our survey efforts are still ongoing for the season, I’m happy to report that robust populations of Cicindela cursitans have been located at several spots along the Mississippi River. Many dozens of individuals were observed at two locations in Mississippi County, and another new population was located further south in New Madrid County. Combined with the sites discovered last year, this gives five confirmed sites for the species within the state. All of these sites share similar features – bottomland forest immediately adjacent to the Mississippi River, with an open understory dominated by poison ivy (Toxicodendron radicans) and trumpet creeper (Campsis radicans) (a ‘radical’ understory to say the least) on a ridge and swale topography of sandy loam soil. The beetles favor the relatively drier, more openly vegetated ridges but avoid areas of excessive sand. None were seen in the wetter sand beach areas leading down to the water’s edge, nor were any observed on the relatively sand-free soils found further away from the river. Unusual for tiger beetles, adults were never found in open sunny areas, being entirely restricted to forest habitats where they darted through the open understory from one poison ivy plant to another. This is in distinct contrast to the wet meadow habitats reported by Brust et al. (2005) for populations of this species in Nebraska. Their small size and rapid running capabilities made them quite difficult to capture or even to notice at first – appearing more like ants or small spiders.

Of equal interest are the sites where the species was not observed, which include sites along the St. Francois River (western side of the Mississippi lowlands) and along the Arkansas border in between the two river systems. All of these sites offered similar bottomland forest, open understory, and ridge and swale soil topography, but they differ from the sites along the Mississippi River where the species was observed in that the soils are a heavy clay and contain no sand. It’s difficult to say conclusively that the species does not occur in these habitats, but the abundance with which we have observed it in the Mississippi River habitats is strongly suggestive.

The Mississippi Lowlands of Missouri, once a vast assemblage of bald cypress (Taxodium distichum) and tupelo gum (Nyssa aquatica) swamps and mixed deciduous bottomland forest, have been almost completely drained, cleared, and converted to agriculture. Only small remnants of natural forest and swamp remain amongst the fields of soybean, wheat, corn, rice and cotton. Despite this, the ribbons of forest that occupy the narrow corridor between the Mississippi River and the levees that confine it seem to offer much potential habitat for Cicindela cursitans. Combined with their confirmed occurrence and abundance at several sites within this habitat, it appears that this species is secure within the state and will not require any special conservation measures to assure its continued presence. In celebration, I share with you some photographs of the adults, taken in their natural habitat at one of the Mississippi County sites, along with a few additional photos of some other tiger beetle species I observed on the wet sand beaches closer to the river’s edge. These latter three species are common in Missouri along the Mississippi and Missouri Rivers: Cicindela repanda (bronzed tiger beetle), Cicindela cuprascens (coppery tiger beetle), and Cicindela hirticollis shelfordi (hairy-necked tiger beetle). After taking pictures of these latter three species along the river bank amidst puzzled looks from a few of the locals, I had an amusing conversation with one of the more ‘colorful’ of them, who had come to the baffling conclusion that I could only have been taking pictures of rocks. I cleared up the confusion and showed him a few of the beetles, and we both returned to doing what we both love – drinking beer and looking for beetles (respectively, that is!).

Glades of Jefferson County

We stood a moment to contemplate the sublime and beautiful scene before us, which was such an assembly of rocks and water—of hill and valley—of verdant woods and naked peaks—of native fertility and barren magnificence… – Henry Rowe Schoolcraft, 1818-1819

In the Ozark Border south of St. Louis, a series of natural openings punctuate the dry, rocky forests of Jefferson County. Commonly called “glades” or “cedar glades,” these islands of prairie in a sea of forest are home to plants and animals more commonly associated with the Great Plains region further to the west. Extending in a narrow arc from central Jefferson County east and south into northern Ste. Genevieve County, these glades occur most commonly on south and southwest-facing slopes below forested ridges and are characterized by thin soils and exposed dolomite bedrock of Ordovician age. Glades are, in fact, a common natural feature throughout much of the Ozark Highlands, an extraordinary plateau where the great eastern deciduous forest begins to yield to the western grasslands. A much more extensive system of dolomite glades occurs in the White River Hills of southwest Missouri, where they often extend up steep slopes and over the tops of knobs to form what Schoolcraft called “naked peaks” and are now called “balds” (and spawning the “Baldknobbers” of Branson fame). Additional glade complexes occur throughout the Ozark Highlands on different rock substrates – igneous glades abound in the St. Francois Mountains, sandstone glades dot the Lamotte landscape in Ste. Genevieve County and the northern and western Ozarks, limestone glades can be found in the northern Ozarks near Danville and Lake of the Ozarks, and chert glades occur in extreme southwest Missouri. These different glade systems share a common feature – shallow soils where tree establishment is limited due to summer moisture stress. They differ vegetationally, however, due to differences in hydrology and soil chemistry as a result of their different substrates. Floristically, dolomite glades exhibit a high degree of diversity relative to other glade types.

The term “glade” is derived from the Old English “glad,” meaning a shining place – perhaps the early settlers found their open landscapes a welcome respite after emerging from the confining vastness of the eastern deciduous forest. Whatever the meaning, the glades of Jefferson County hold a special place in my heart, for I “grew up,” entomologically speaking, in those glades. As a young entomologist, fresh out of school, I spent many a day scrambling through the glades and surrounding woodlands. It was here where my interest in beetles, especially woodboring beetles, was born and later grew into a passion. For eight years I visited these glades often – attracted by the extraordinary diversity of insects living within the glades and congregating around its edges. My earliest buprestid and cerambycid papers contain numerous records from “Victoria Glades” and “Valley View Glades” – the two best-preserved examples of the glades that once occurred extensively throughout the area (more on this later). My visits to these glades ended in 1990 when I moved to California, and although I moved back to the St. Louis area in 1995, the focus of my beetle research has more often taken me to places outside of Missouri. It had, in fact, been some 10 years since my last visit to these glades until last week, when I was able to once again spend some time in them.

Ozark glades differ from the true cedar glades of the southeastern U.S. in that they are not a climax habitat – they depend upon periodic fires to prevent succession to forest. Some recent authors have suggested the term “xeric dolomite/limestone prairie” be used to distinguish the fire-dependent glades of the Ozarks from the edaphic climax cedar glades of the southeast (Baskin & Baskin 2000, Baskin et al. 2007). Fires have been largely suppressed throughout Missouri since European settlement, leading to encroachment upon the glades by eastern red-cedar (Juniperus virginiana). Pure stands of red-cedar have developed on many former glades, crowding out the herbaceous plants that depend upon full sun and leading to soil formation that supports further encroachment by additional woody plant species such as post oak (Quercus stellata), blackjack oak (Q. marilandica), flowering dogwood (Cornus florida), and fragrant sumac (Rhus aromatica) from the surrounding woodlands. Fire has returned to many of the Ozark glades situated on lands owned or managed by state and federal agencies such as the Missouri Department of Conservation, Missouri Department of Natural Resources, and U.S. Forest Service, as well as private conservation-minded organizations such as The Nature Conservancy. These agencies have begun adopting cedar removal and fire management techniques to bring back the pre-settlement look and diversity of the Ozark Glades. This is particularly true at Victoria Glades and Valley View Glades, the two largest and most pristine examples of the Jefferson County dolomite glade complex. Fires have been used to kill small red-cedars in the glades, as well as rejuvenate their herbaceous plant communities. Larger red-cedar trees are not killed outright by fire and must be removed by chainsaws. This above distant view of the TNC parcel at Victoria Glades shows many such burned red-cedars. The glades themselves are not the only habitat to benefit from this aggressive management – when I was doing my fieldwork here in the 1980’s the surrounding woodlands were a closed post oak forest bordered by fragrant sumac and with little or no understory in the interior. The photo at right now shows an open savanna with a rich understory of not only sumac and other shrubs, but also many herbaceous plants as well such as black-eyed susan (Rudbeckia hirta) and American feverfew (Parthenium integrifolium). Such open woodland more closely resembles what Schoolcraft saw across much of the Ozarks during his journey almost two centuries ago.

Victoria and Valley View Glades are dominated by little bluestem (Schizachyrium scoparium), Indian grass (Sorghastrum nutans), big bluestem (Andropogon gerardii) and prairie dropseed (Sporobolus heterolepis). A smaller but highly charismatic non-grass flora is also found on the glades – species such as Missouri evening primrose (Oenethera macrocarpa) (left), pale purple coneflower (Echinacea simulata) (pictured above and below), and prairie dock (Silphium terebinthinaceum) not only add beautiful color but also support both vertebrate and invertebrate wildlife. The Fremont’s leather flower (Clematis fremontii) is a true endemic, occurring only in this part of Missouri and entirely dependent upon these glades for its survival. Less well studied is the vast insect fauna associated with the glades. It is here where I first discovered the occurrence of Acmaeodera neglecta in Missouri. This small jewel beetle is similar to the broadly occurring A. tubulus but at the time was known only from Texas and surrounding states. In collecting what I thought were adults of A. tubulus on various flowers in the glades, I noticed that some of them were less shining, more strongly punctate, and exhibited elytral patterning that was often coalesced into longitudinal “C-shaped” markings rather than the scattered small spots typical of A. tubulus. These proved to be A. neglecta, which I have since found on many glade habitats throughout the Ozark Highlands. Both species can be seen in this photo feeding on a flower of hairy wild petunia (Ruellia humilis) – the lower individual is A. neglecta, while the upper individual and two inside the flower are A. tubulus. Another interesting insect-plant association I discovered at these glades was the strikingly beautiful Dicerca pugionata – another species of jewel beetle – and its host plant ninebark (Physocarpus opulifolius). Only a single Missouri occurrence had been reported for D. pugionata, despite the common occurrence of its host plant along rocky streams and rivers throughout the Ozark Highlands. This plant also grows at Victoria and Valley View Glades along the intermittent streams that drain the glades and in the moist toeslopes along the lower edges of the glades where water that has percolated through the rocks and down the slopes is forced to the surface by an impermeable layer of bedrock. Unlike the tall, robust, lush plants that can be found in more optimal streamside habitats with good moisture availability, the ninebark plants of Victoria and Valley View Glades are small and scraggly, usually with some dieback that results from suboptimal growing conditions. I surmise these plants have reduced capabilities for fending off attacks by insects, including D. pugionata, and as a result a healthy population of the insect thrives at these glades. Some might be inclined to call this beetle a pest, threatening the health of one of the glade’s plants. In reality, the insect finds refuge in these glades – unable to effectively colonize the vast reserves of healthy plants that grow along streams throughout the rest of the Ozarks, it strikes a tenuous balance with plants that are themselves on the edge of survival.

Despite the success in moving Victoria and Valley View Glades closer to their pre-settlement character, the integrity of these areas continues to be challenged. Poachers take anything of real or perceived value, and ATV enthusiasts view the open spaces as nothing more than tarmac. Pale purple coneflower occurs abundantly on these Jefferson County glades (but sparingly in other habitats – primarily rocky roadsides), where they provide a stunning floral display during June and sustain innumerable insect pollinators. Plants in the genus Echinacea also have perceived medicinal value, as herbalists believe their roots contain an effective blood purifier and antibiotic. There are no conclusive human clinical trials to date that fully substantiate this purported immune stimulating effect (McKeown 1999). Nevertheless, demand for herbal use has skyrocketed in recent decades, prompting widespread illegal harvesting of several coneflower species throughout their collective range across the Great Plains and Ozark Highlands. I witnessed massive removals of this plant from both Victoria and Valley View Glades during the 1980’s, but the pictures I took this year suggest that such illegal harvests have been suppressed and that the populations at both sites are recovering nicely.

The same cannot be said for the practice of rock flipping. This was a problem I witnessed back in the 1980’s, and I saw fresh evidence of its continued occurrence at both sites. The thin soils and sloping terrain leave successive layers of dolomite bedrock exposed, the edges of which shatter from repeated freeze-thaw cycles to create rows of loose, flat rocks along the bedrock strata. Lizards, snakes, tarantulas, and scorpions find refuge under these loose rocks, only to be ripped from their homes by flippers and transferred to a dark, cold terrarium to endure a slow, lingering death. As if poaching the glade’s fauna and watching them slowly die isn’t bad enough, the flippers add insult to injury by not even bothering to replace the rock in its original position after stealing its inhabitant, amounting to habitat destruction three times greater than the area of the rock itself. Firstly, the habitat under the rock is destroyed by sudden exposure of the diverse and formerly sheltered microfauna to deadly sunlight. Next, the habitat onto which the rock is flipped is also destroyed, as the plants growing there begin a slow, smothering death. Lastly, the upper surface of the rock, sometimes colonized by mosses and lichens that might have required decades or longer to grow, usually ends up against the ground – its white, sterile underside becoming the new upper surface. Rock flipper scars take years to heal, and nearly all of the flat, loose rocks seen in the more accessible areas of the glades exhibit scars of varying ages next to them. If a scar is fresh (first photo), I generally return to the rock to its original position – the former inhabitants cannot be brought back, but at least the original habitats are saved and can recover quickly. However, if a scar is too old (2nd photo) it is best to leave the rock in its new position – replacing it only prolongs the time required for recovery.

Even more damaging is ATV use. Herbaceous plants and thin soils are no match for the aggressive tread of ATV tires, and it doesn’t take too many passes over an area before the delicate plants are killed and loose soils ripped apart. I witnessed this become a big problem particularly on Victoria Glades during the 1980’s – actually finding myself once in a face-to-face confrontation with an ATV’er. Fortunately, he turned tail and ran, and it appears (for now) that such abuses have stopped, as I saw no evidence of more recent tracks during this visit. But the scars of those tracks laid down more than two decades ago still remain painfully visible. I expect several more decades will pass before they are healed completely.

My return to Victoria and Valley View Glades was a homecoming of sorts, and I was genuinely pleased to see the progress that has been made in managing these areas while revisiting the sites where my love affair with beetles was first kindled. Sadly, however, the larger glade complex of Jefferson County continues to deteriorate. Restoration acreage aside, red-cedar encroachment continues unabated on many of the remaining glade parcels – large and small – that dot the south and southwest facing slopes in this area. It has been conservatively estimated that as much as 70% of the original high quality glades in Missouri are now covered in red-cedar. Many of these are privately held – their owners either do not recognize their ecological significance or are loathe to set fire to them. An example can be seen in the picture here – this small parcel is part of the Victoria Glades complex but lies on private land in red-cedar choked contrast to the Nature Conservancy parcel immediately to the south. Small numbers of herbaceous plants persist here, but without intervention by fire or chainsaw their numbers will continue to dwindle and the glade will die. Aside from the loss of these glades, the continuing reduction of glade habitat complicates management options for preserved glades as well. Many glade associated invertebrates are “fire-sensitive” – i.e., they overwinter in the duff and leaf litter above the soil and are thus vulnerable to spring or fall fires. While these fires are profoundly useful for invigorating the herbaceous flora, they can lead to local extirpation of fire-sensitive invertebrate species within the burn area. Recolonization normally occurs quickly from unburned glades in proximity to the burned areas but can be hampered if source habitat exists as small, highly-fragmented remnants separated by extensive tracts of hostile environment. Grazing also continues to threaten existing remnants in the Jefferson County complex. Grazing rates are higher now than ever before, with greater negative impact due to the use of fencing that prevents grazers from moving to “greener pastures”. Over-grazing eliminates native vegetation through constant depletion of nutrient reserves and disturbance of the delicate soil structure, leading to invasion and establishment of undesirable plant species. Eventually, the glade becomes unproductive for pasture and is abandoned – coupled with fire suppression this leads to rapid woody encroachment. It is truly depressing to drive through Jefferson County and recognize these cedar-choked glades for what they were, able to do nothing but watch in dismay as yet another aspect of Missouri’s natural heritage gradually disappears. The continued loss of these remnant glades makes careful use of fire management on Victoria and Valley View Glades all the more critical – ensuring that a patchwork of unburned, lightly burned, and more heavily burned areas exists at a given time will be critical for preventing invertebrate extirpations within these managed areas.

I close by sharing with you a few more of the many photographs I took during this visit – stiff tickseed (Coreopsis palmata), three-toed box turtle (Terrapene carolina triunguis), climbing milkweed (Matelea decipiens – see the excellent post about this plant on Ozark Highlands of Missouri), downy phlox (Phlox pilosa), green milkweed (Asclepias viridiflora), and a “deerly” departed native browser.

Saving Missouri’s tigers

For several years now, my friend and colleague, Chris Brown, and I have been studying the tiger beetles (family Cicindelidae) of Missouri in an attempt to characterize the faunal composition and in-state distributions of the included species. Our studies have relied on examination of specimens in museum collections along with several seasons of field work across the state. The data we’ve gathered so far have revealed a fauna that reflects the ecotonal position of Missouri, comprised of elements from the eastern deciduous forest, the southeastern mixed hardwood forests and pinelands, and the western grasslands. These beetles are most frequently associated with disturbed habitats containing sparse vegetation, such as sandbars and erosion cuts, but they also live in other habitats such as along muddy banks, on glades and in forest litter. Since European settlement of Missouri, drastic alterations have occurred in the abundance and distribution of these habitats across the state, and tiger beetle populations have been affected as a result. Dredging and straightening of natural water courses have impacted species that prefer the water’s edge, while fire suppression has impacted those that need dry, open habitats. Grazing has had a profound impact on species associated with sensitive, saline habitats. Conversely, some anthropogenic changes have benefited certain species – road, borrow sand pit and pond construction have increased habitat for species able to utilize such habitats. To date, our surveys have confirmed the presence of 23 species in Missouri (16% of the North American fauna). Some species are common and widespread, such as Cicindela sexguttata (six-spotted tiger beetle) (above, photo taken in a mature white oak forest in Warren Co.). Others have more specific habitat requirements, but their status within the state remains secure. A few are rare and highly localized, primarily representing species at or near the northern or western edge of their distributions that exist in the state as small, disjunct populations. Special conservation efforts may be warrented for these to ensure their continued survival within the state.

One species of potential conservation concern is C. pruinina (loamy-ground tiger beetle), a grassland species normally found in Kansas, Oklahoma, and Texas (left, photo by Chris Brown). This beetle is sometimes treated as a synonym or subspecies of C. belfragei but was most recently considered a valid species by Spomer et al. (2008). When we began our studies, the species was known from Missouri only by a small series of specimens collected in one of the western counties and deposited in the collection of Ron Huber. An additional specimen in the Huber collection labeled “Columbia, Mo.” is considered to likely represent student mislabeling. Several attempts at relocating the western Missouri population were required before we found it, and intensive surveys to determine the extent of its distribution in Missouri were conducted during 2006 using a combination of pitfall traps and direct observation. Those surveys succeeded in detecting the beetle only along one 2.5-mile stretch of county road in Johnson Co. The beetle seems to be restricted to red clay embankments occurring in a small localized area of the county. This season we plan to refine our survey by focusing tightly on promising habitats in this area near the sites identified in 2006 to more precisely define the distributional limits of this population. Regardless of what we find, the flightless nature of the species, its highly localized Missouri occurrence, and the disjunct nature of such suggest that special conservation status is warranted for the species to ensure its continued presence in the state. Despite the relatively low numbers of individuals we have seen, the protected status of the land on which this beetle lives leaves us optimistic about its future.

A Missouri species about which we are far less optimistic is C. circumpicta johnsonii (saline springs tiger beetle) (right, photo by Chris Brown), long known from saline spring habitats in the central part of the state. The Missouri population is highly disjunct from the main population further west and exhibits a uniform blue-green coloration rather than the mix of blue, green, and red colors exhibited by the main population. These features suggest that separate subspecies status might be warranted for the Missouri population. Numerous historical collection records exist from a handful of sites in Howard and Cooper Counties; however, surveys conducted by us during recent years revealed that the populations had suffered severe declines. This appears to be largely due to cattle disturbance and vegetational encroachment of the sensitive saline spring habitats upon which the beetle depends, especially at sites located on private land. Ron Huber, in a letter to me listing the collecting records he had for this species, reported seeing “hundreds of the wary little buggers” at one locality on private land, but in our visit to the site not a single individual was seen around the spring – badly trampled and overgrown with the exotic pasture grass tall fescue (Festuca arundinacea). In all, we were successful in finding the beetle at only one of the historical localities and at one new site located nearby. While both of these sites are located on state protected land, we concluded that the longterm viability of the C. circumpicta johnsonii population in Missouri was in serious jeopardy. Based on our recommendation, the species was placed on the Missouri Species of Conservation Concern Checklist with a ranking of S1 (critically emperiled). While this affords the species legal protection under the Wildlife Code of Missouri, the benefit may be minimal since the Wildlife Code does not address the main threat to this beetle’s survival – habitat degradation. We have not surveyed for this beetle since but plan to make field observations this summer (no trapping!) to check on its status. I sincerely hope we will not have to hang our heads with the realization that we have succeeded in extirpating yet another beautiful and irreplaceable gem.

Last year we finally succeeded in locating C. cursitans (ant-like tiger beetle) in Missouri (left, photo by Chris Brown). Adults are flightless, and at less than 1 cm in length, are among the smallest of North American tiger beetles. The species has been recorded broadly but sporadically between the Appalachians and Great Plains – a distribution that is probably underestimated due to its small size and general resemblance to ants. A significant distributional gap exists between the eastern and Great Plains records, which Ron Huber believes may be indicative of two disjunct forms and potentially two species. Missouri falls within this gap, and although the species has not yet been formally recorded from the state, a single specimen collected in 1991 “nr. Portageville” is deposited in the Enns Entomology Museum, University of Missouri, Columbia. We had made several attempts over the past few years to locate this species by searching what we thought were promising habitats along the Mississippi River near Portageville, but the species was not located until last year, when I relayed this information to Portageville biologist and tiger beetle enthusiast Kent Fothergill. Kent not only located the beetle at the location I suggested, but quickly found another population on a nearby parcel of land managed by the Missouri Department of Conservation. Hurriedly, we visited the first site and observed a few additional individuals at a nearby location just to the south. The bottomland forest habitat within which all of these individuals were observed is fairly extensive along the Mississippi River in the southeastern lowlands of Missouri, but at this point we can only speculate whether C. cursitans occurs throughout this habitat. Other habitats have been reported for this species, including mesic and wet prairies and meadows (Brust et al. 2005). Such habitats are also found in the southeastern lowlands, and while C. cursitans has not been seen through cursory examinations in such areas, it is possible that the small size, cryptic habits, and narrow temporal occurrence of C. cursitans have allowed it to escape detection. Kent will be helping us this season with additional trapping and direct observation at several selected sites along the Mississippi and St. Francois Rivers to determine whether the beetle occurs more broadly in the southeastern lowlands and whether it utilizes these other habitats in addition to bottomland forests.

I would be most interested in any additional reports of these rare tiger beetles in Missouri (contact me at the email address shown in the left sidebar copyright statement). Remember, C. circumpicta johnsonii is critically emperiled in Missouri – please do not collect it.

Saving turtles, one by one…

“You should be able to push him from behind,” said the other motorist, who had also seen the young snapping turtle sitting in the middle of the exressway right after I did and pulled up as I was taking some “pre-rescue” pictures. “Nyeah, I think I’ll go ahead and get something to push him with anyway,” I said, hoping that the tone of my voice did not betray my true thoughts, “What, are you crazy? I’m not putting my bare fingers or sandal-clad feet anywhere near that thing!” I’ve rescued plenty of snappers over the years, and I know first hand just how surprisingly quick they can be. Truth be told, snapping turtles can be safely moved by hand – apparently they cannot reach the back of or underneath their shell. However, it takes considerably more temerity than I possess to actually try this. I grabbed a bicycle pump from the back of the truck and hooked the base of it under its shell. Immediately, the young turtle snapped at the pump – startling the man as well as two other cyclists who had stopped to watch the goings on. I admit to feeling more than a little vindicated as they all stepped back a few steps. I was hoping the turtle would maintain his grip on the pump so I could just carry him over to the roadbank, but every time I tried to lift he let go. So I had to just keep hooking the pump under his shell and pulling him towards the side of the road – the turtle fought every bit of the way, hissing and snapping and clawing against the road. At last he was in the grass – it was then an easy matter to roll him over a few times down the bank and safely (for now) away from the road.

Road mortality is suspected to have contributed to widespread population declines in turtles across the United States. This seems especially true for freshwater aquatic species, which often make land migrations for breeding. Vehicles often do not stop for turtles in the road, and I have seen some (usually a pickup truck with very large tires) swerve deliberately in an attempt to hit them (or even more sadistically, “shoot” them across the roadway). Conincident with these declines has been a demographic change towards male-biased populations in many freshwater species. Adult female freshwater turtles make nesting migrations that males do not and are often attracted to road shoulders and embankments as nesting habitat, making them disproportionately more vulnerable to road mortality. The resulting male-biasing surely represents an additional risk factor to their populations, especially in areas where high traffic occurs in proximity to wetlands. In such places, mitigation measures such as barriers and wildlife underpasses are clearly warranted (Steen et al. 2006).

I’ve always been a little awed by snappers – so grizzled and ancient, almost dinosaurian, and while I doubt that my sporadic rescues have near as much impact as barriers or underpasses, I do know that they cannot possibly hurt. As for this turtle, whether it continued on its way or turned around and crawled back onto the road (due to my unwittingly placing it on the side from which it just came) will remain unknown. I was heartened to see that I wasn’t the only person who stopped, intent on saving this grotesquely beautiful creature. But as I scanned this miles-long stretch of very recently constructed roadway, which now enables St. Louis countians to rapidly zip along next to newly created wetlands in the Missouri River bottoms while avoiding the stop-and-go on I-270, I couldn’t help but wonder why barriers and underpasses, seemingly simple protective measures, weren’t also included on the final blueprints of the roadway before they were sent to the printers. If such had been done, then I would not have had this encounter. But I could’ve lived with that.

Pipestone National Monument

Grandson, do not expect to accomplish much in this lifetime, for no one shares your vision… – The Oracle


Wednesday was my birthday, and it has been my custom for many years now to take the day off and go hiking/bugging somewhere. Coming as it does in early spring, it is usually the first real bug collecting trip of the year. This year, however, I was roped into a short business trip to visit a USDA lab in Brookings, South Dakota, so tradition would have to take a back seat. My visit at the lab ended early, though, and my flight back home from Sioux Falls didn’t leave until that evening, so I studied the map to look for any possible nearby points of interest in this landscape that has, for the most part, been unforgivingly converted to fields of corn, soybean, and wheat. I quickly noted a place called Pipestone National Monument just over the border into Minnesota. I love stopping at national monuments while traveling – they usually have some significant historical or geological interest, and their typically (though not always) small size means one can fully explore the area in a relatively short time. I did not know or had never heard of this place, but what I found was a charming little jewel tucked within a remnant of tallgrass prairie. At this far northern latitude, spring is still in its earliest of states. Few insects would be seen, but nevertheless I felt thankful for the chance to spend time outdoors and in a place of beauty where I could reflect on the years gone by and those (hopefully) still to come.

The area is named for a thin layer of catlinite – pipestone – exposed in this small area that has been quarried for centuries by Native Americans for carving into pipes (both war and peace). Quarrying within the monument continues to this day, with permission to do so reserved by law only to registered Native Americans. The area is identified as a sacred site associated with Native American spiritual beliefs and is preserved as a significant cultural and ethnographic landscape. Of particular interest to me was the site’s distinct hydrologic/geologic landscape and the native tallgrass prairie associated with it. A short ¾-mile trail loops through the area, providing a diverse glimpse of the area’s unique features. Pipestone may have provided the area’s namesake, but a narrow exposure of Sioux quartzite is the area’s most prominent geologic feature. Sioux quartzite is derived from billion and a half year-old layers of sand/silt sediments deposited thickly on the floors of ancient, Precambrian seas and compressed over the vastness of time into a hard, reddish metamorphic rock. Normally covered in this area by glacial till, the layers at this site are tilted upward 5–10 degrees towards the west and break through the surface to form a jagged, mile-long west-facing escarpment 23-30 feet high. Underneath the quartzite is the pipestone, a thin layer of metamorphosed shale. This fine-grained rock is derived from clay deposits, thus it is much softer and redder than the harder-than-steel quartzite. Pipestone Creek bisects the escarpment, giving rise to the lovely Winnewissa Falls, flowing over the escarpment and running down to a small, natural empoundment (Hiawatha Lake) before continuing its journey back into the glacial till and tallgrass prairie (for anybody surprised that there should be “falls” in this part of the country, it is interesting to note that nearby Sioux Falls is named after a grander example of of such flowing over quartzite exposures in its downtown).

Precious little remains of the expansive tallgrass prairie that once extended from horizon to horizon in this area. A few small parcels managed to escape the plow, but even in those tiny remnants dramatic alterations in plant communities have occurred due to fire suppression and the introduction of more than 70 non-native plant species. Prescribed burning programs are now being used at the Monument to restore the prairie’s native plant composition and appearance. Looking out over the tallgrass prairie remnants above and below the quartzite escarpment, it I was tempted to visualize circles of teepees on the higher ground away from the quarries (all Native American tribes worshipped this site and would never camp directly within it), with herds of American bison dotting the landscape in the distance. Contrasting with the openness of the prairie, the escarpment itself is densely studded with trees – American elm (Ulmus americana) along the top edge, and bur oak (Quercus macrocarpa) in the escarpment itself. Unlike the large, sometimes towering examples of their kind found further to the east, the trees here are dwarfed and spreading, almost gnarled. Below the escarpment, woodland quickly gives way to pure stands of smooth sumac (Rhus glabra) and choke cherry (Prunus sp.), which just as quickly yield to the surrounding sea of prairie. Along Pipestone Creek below the escarpment, lower layers of exposed quartzite provide nooks and crannys where enough moisture collects to support the growth of green ash (Fraxinus pennsylvanica) trees, until glacial till once again covers the quartzite, and riparian woodland yields to grasses and forbes. It’s not hard to imagine why this became a special place to the Native Americans, even before they discovered the pipestone that was to become so important to their culture.

The pipestone quarries are located a short distance to the west of the escarpment – where the hard quartzite layer is thin enough to break through – and, thus, have had little impact on altering the physical appearance of the escarpment itself. Winnewissa Falls (meaning “Jealous Maiden” in the Dakota language), lies at the center of the escarpment, providing a stunning centerpiece. Despite its beauty, it is but a shadow of what it was before early settlers in the area blasted away the top 18 feet of the ledge to create a reservoir for drinking water. A century of weathering and recolonization by lichens and mosses have softened the scars on the rocks, leaving little to indicate that such a dramatic alteration took place. However, standing in front of the falls, finding that “zone” where the temperature suddenly drops and cool wet mist blows on the face, and thinking about the significance of this place to the Native American tribes who held it so sacred, I was left feeling bewildered at how such drastic measures could have been contemplated for so beautiful a place.

In addition to the falls, nature has created some striking sculptures in the rock. “Old Stone Face” can hardly be mistaken for anything else – despite its human likeness, it was created entirely by natural forces. “The Oracle” is another naturally-formed human likeness found (though not as easily as Old Stone Face) in the outlines of the rocks. Tribal Shamans (Medicine Men) believed it served as a guardian of the valley and that voices issued from it’s cold stone lips. I stared for awhile and strained to listen, trying to imagine what words it might have spoken. At first, it seemed as if all was silent. Then I noticed the sound of the wind rolling over the prairie and twirling through the gnarled oaks. I heard the falls in the distance. I heard birds in the midst of frantic early-spring songs. I thought perhaps these might be the voices that guided the Shamans – spoken so loudly, yet so easily unheard.

The first U.S. government expedition to the quarry occurred in 1838 with Joseph Nicollet, a French scientist who was sent to map the upper Mississippi country. He and the members of his expedition carved their names in the rocks atop the escarpment, as did many of the early pioneers that first settled in the area. In studying the surface of these rocks, I couldn’t help but notice the incredible diversity of lichens to be found. Around 75 species are known from the area, and as shown in the photos I share below they come in a fantastic array of forms and colors. Lichens are primary colonizers of rock surfaces, able to do so as a result of their nutritional autonomy. Lichens are merely fungi that have evolved a specialised mode of nutrition: symbiosis with photosynthetic microalgae or cyanobacteria. Often, the algal component is capable of fixing nitrogen from the atmosphere, while the fungal organism attacks the rock with organic acids to release minerals. This is the basis of soil formation. Over time, enough soil accumulates in small depressions to allow mosses to colonize the rock surface. As successive generations of moss grow and die, more and more organic material accumulates on the rock surface, eventually supporting the growth of vascular plants (which extract nitrogen from the soil, rather than from atmospheric sources). These cycles of growth and death act in concert with the forces of erosion to ultimately convert barren rock to tallgrass prairie, hardwood forest, or other climax habitat. Mind you, this is an extraordinarily slow process – it can take a full century for a lichen to grow one inch! As I looked at the abundance and diversity of lichens on the rock surfaces, I tried to visualize the breadth of time encompassed by what was before me and quickly became lost in eternity.

While the trail that loops through the area is less than a mile in length, it took me an hour and a half to complete it. What started out as a few hours to kill ended as a hurried rush through the museum and interpretive center, trying to cram a few last morsels of knowledge into my head in those final moments before I would have to submit to the drive back to Sioux Falls. As I left the area, I noticed these oddly out-of-place boulders known as “The Three Maidens.”
Native Americans believe that these boulders shelter the spirits of maidens who demand offerings before permitting them to quarry the pipestone. Science tells us that the boulders are composed of granite and were likely carried here by glaciers during the past 1 million to 10,000 years ago. Originally a single boulder some 50 feet in diameter, repeated freezing and thawing over the millenia since it was dropped here have split the boulder into the several pieces seen here. Perhaps only The Oracle knows which is true.

For a more detailed, yet highly readable account of the geology of this area, please consult Minnesota Geology, Field Trip, Summer 2000 and Other MN DNR Workshops, by Arlyn DeBruyckere.