habitat management

The hunt for Cicindela celeripes

/ Ted C. MacRae / 12 Comments

One of the more enigmatic tiger beetle species in North America is Cicindela celeripes LeConte (swift tiger beetle). This small (6-8 mm), flightless species has been recorded from a restricted area of the eastern and southern Great Plains – from eastern Nebraska and westernmost Iowa south through Kansas to western Oklahoma and the Texas panhandle (Hoback and Riggins 2001, Pearson et al. 2006). Unfortunately, populations of this species appear to have suffered severe declines. It apparently is holding strong in the Flint Hills region of Kansas, but many of the records from outside of that area date back more than a century. Reportedly once common on the bluff prairies along the Missouri River, it has not been seen in Nebraska since 1915 and may have been extirpated from that state (Brust et al. 2005). The reasons for this decline undoubtedly involve loss of preferred habitat – upland prairies and grasslands with clay or loess soils and sparse or patchy vegetation. Areas supporting these native habitats have been drastically reduced since European settlement of the region, and suppression of fire – so vital to prairie ecosystems – has led to extensive woody encroachment on the few prairie relicts that do remain. Unlike many other tiger beetle species that have been able to adapt to these anthropogenic changes, this species apparently cannot survive in such altered habitats.

Chris Brown and I have been interested in this species ever since we began surveying the tiger beetles of Missouri. It has not yet been recorded from the state, but we have long suspected that it might occur in extreme northwest Missouri. It is here where the Loess Hill prairies along the Missouri River reach their southern terminus. (Incidentally, the Loess Hills are themselves a globally significant geological landform, possessing natural features rarely found elsewhere on earth. They will be the subject of a future post). We have searched several of what we consider to be the most promising potential sites for this species in Missouri, though without success. Nevertheless, we remain optimistic that the species might eventually be found in Missouri and has simply been overlooked due to the limited temporal occurrence, small size, rapid running capabilities, and tendency of adults to dart rapidly to the bases of grass clumps where they hide (Pearson et al. 2006). Furthermore, even though the species has not been seen recently in adjacent areas of Nebraska where it has been recorded in the past, it has been seen recently in a few Loess Hill prairie remnants just to the north in Iowa.

A few weeks ago, I was fortunate to receive specific locality information for one of the recently located Iowa populations. Armed with site descriptions, Google maps, photographs, and whatever book learnin’ I had gained about this species, my colleague and I made the long drive to southwestern Iowa in hopes of locating the population for ourselves, seeing adults in their native habitat, and using the learnings we would gain about their habitat preferences and field behavior to augment our efforts to eventually locate the species in northwestern Missouri. At mid-July, we were nearing the end of the adult activity period, but adults had been observed at the site the weekend prior, so we felt reasonably confident that adults might still be found. Additionally, fresh off of our recent success at locating the related Cicindela cursitans in Missouri (another small, flightless, fast-running species), we were hopeful that we now possessed the proper “search image” to recognize C. celeripes in the field should we have the good fortune to encounter it.

Walking into the area, I was impressed at the extensiveness of the prairie habitat – much larger than any of Missouri’s Loess Hill prairies. The presence of large, charred red-cedar cadavers on the lower slopes revealed active management for prairie restoration. We later learned from the area manager that the restoration area had been acquired from a neighboring landowner who had used the land for grazing and sold it when it became unproductive. I can only imagine the second thoughts that landowner must have had when subsequent burn regimes and woody growth removal prompted a return to the beautifully lush sea of prairie vegetation that now covered the hills. As we approached the area where we decided the beetles must have been seen, we started searching slowly and deliberately – looking carefully for any movement between the clumps of grass. It didn’t appear to be prime habitat for C. celeripes – the vegetation was just so thick, with only small openings among the plants. We continued to scour the area closely but saw nothing, and my optimism began to wane. Wrong spot? – I don’t think so. Bad search image? – hard to imagine, considering its similarity to C. cursitans. Too late? – could be.

After it became obvious we were searching the same gaps in the vegetation repeatedly, I started walking towards a small cut further down the hillside that I had noticed earlier (just visible in the previous photo). I had thought, “That’s tiger beetle land down there!” My optimism increased when I reached the cut, seeing the remains of an old, overgrown 2-track leading through the cut and on down the hillside. Vegetation was much sparser within and below the cut – it looked perfect. Chris had become distracted taking photographs of something, so I began searching. I’d been in the cut a few minutes when I thought I saw something flash across a bare patch out of the corner of my eye – was that it? It had to be. I carefully inspected around the base of every clump of vegetation at my feet but found nothing. It must have been wishful thinking – just another spider. I continued on down the cut, and within a few more minutes I saw the flash again – this time there was no doubt as to what it was, and I had a lock on it. I started slapping the ground frantically as the little guy darted erratically under, around, and over my hands. In the few seconds while this was happening, I was simultaneously exuberant at having succeeded in finding it, utterly astounded by its speed and evasiveness, and desperately afraid that it was getting away – swift tiger beetle, indeed! Persistence paid off, however, and eventually I had it firmly in my grasp.

We would see a total of seven individuals that day. Most of them were within or immediately below the cut, while another individual was seen much further down the 2-track. Mindful of the population declines this species has experienced, we decided to capture just three individuals (even though by this point in the season mating and oviposition would have been largely complete) in hopes that at least one would survive the trip back to the lab for photographs. Our primary goal – to see the species in its native habitat – had been accomplished. We now turned our attention to attempting in situ field photographs. This would prove to be too difficult a task – each beetle we located immediately ran for cover, and flushing it out only caused it to dart to another clump of vegetation. This scenario repeated with each beetle until eventually it simply vanished. We would have to settle for photographs of our captured specimens in a confined arena – a few of which are shown here. The beetles were photographed on a chunk of native loess taken from the site, and no chilling or other “calming” techniques were used. Spomer et al. (2008, Field Guide to the Tiger Beetles of Nebraska and South Dakota) state that C. celeripes is a delicate species that does not do well in captivity. It has never been reared, and the larva is unknown. Nevertheless, I placed the chunk of native loess in a plastic tupperware container and transplanted into one corner a small clump of bluegrass from my yard. The soil around the grass clump is kept moist, and every few days I have placed various small insects in the container. Of the three individuals that we brought back, two died within two days. The third individual (these photographs), however, has now survived for four weeks! Moreover, it is a female, and during the past two weeks six larval burrows have appeared in the soil (and another egg was seen on the soil surface just yesterday). Indeed, an egg can be seen in the upper right of the first photo. It remains to be seen whether I will be successful at rearing them to adulthood; however, I’m hopeful this can be accomplished using methods described for C. cursitans (Brust et al. 2005).

Do I still think C. celeripes occurs in Missouri? I don’t know – on one hand, the mixed grass Loess Hill prairie habitats in which the beetle lives in Iowa do extend south into Missouri, and the beetle could be inhabiting them but be easily overlooked for the reasons I’ve already mentioned. However, Missouri’s Loess Hill prairie relicts are small, both in number and in size, and highly disjunct. Such features increase the likelihood of localized extinctions and hamper recolonization through dispersal, especially in flightless species that must traverse unsuitable habitat. With its adult activity period winding down, renewed efforts to locate this species in Missouri will have to wait until next season. Hopefully, the knowledge we gained this season will help this become a reality. For now, the hunt continues…

(closing photo by C. Brown)

Sand Prairie Conservation Area

/ Ted C. MacRae / 11 Comments

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.

Glades of Jefferson County

/ Ted C. MacRae / 13 Comments

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.

Pipestone National Monument

/ Ted C. MacRae / 5 Comments

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.

Lake Tahoe, California

/ Ted C. MacRae / Leave a comment

…at last the Lake burst upon us — a noble sheet of blue water lifted six thousand three hundred feet above the level of the sea, and walled in by a rim of snow-clad mountain peaks that towered aloft full three thousand feet higher still! It was a vast oval, and one would have to use up eighty or a hundred good miles in traveling around it. As it lay there with the shadows of the mountains brilliantly photographed upon its still surface I thought it must surely be the fairest picture the whole earth affords. – Mark Twain, Roughing It (1872)


Mark Twain may not have liked the name “Lake Tahoe” – preferring its then-official, patronimic designation as “Lake Bigler.” However, he was clearly overwhelmed by its beauty, and surely no person who has ever seen this place can find fault with the words he so eloquently penned almost a century and a half ago. The view above of Emerald Bay, on the south side of the lake, may not be where Twain first viewed Lake Tahoe, but for me it is the most iconic place from which to view it. I first fell in love with Lake Tahoe almost 18 years ago, when my then fiancée and I first moved to Sacramento. We married up there, and for the 5 years we lived in California we spent many a weekend enjoying Tahoe’s 4-season charm. It has been 12 years since we moved back to St. Louis, and I hadn’t been back — until this past weekend. The reasons for the delay are many, but returning to this place reminded me why I consider it the most beautiful place in the world. I shall not let so long a time pass before my next visit.

Lake Tahoe is a relatively young lake, forming within the last several million years (in contrast, the block of granite that was to become the Sierra Nevada mountains – and in which Lake Tahoe lies – began forming during the Paleozoic Era and was then exposed by erosion beginning about 130 million years ago). The basin in which the lake lies was formed by fault-induced block slippage between two uplifted blocks, with the lake itself forming after magma upwellings dammed the northern part of the basin. Glacial action in more recent years (2 million to 20,000 years ago) caused additional damming, causing drastic fluctuations in the lake level — maximum levels reached nearly 800 feet higher than present. The most recent glaciations (~10,000 years ago) carved out Donner Lake (just east of Lake Tahoe), Emerald Bay (above), and nearby Fallen Leaf Lake (below — the frozen lake surface can just be seen above the trees in the foreground).


Emerald Bay is actually part of a glacial “staircase” featuring intermittent flat stretches containing lakes and meadows before ultimately ending at Emerald Bay. Eagle Lake lies immediately above Emerald Bay on one of these “steps,” and the 1-mile trail to it is one of the most popular hikes in the area — below is a view towards Eagle Lake from Emerald Bay:


On the day we arrived (Sat 3/15), a late winter storm was dumping new snow on the surrounding mountains, as seen in this view across the south end of the lake towards the city of South Lake Tahoe. Heavenly Ski Resort was shrouded from view on this day, but the fresh powder being dumped there would provide for some delightful spring skiing over the next few days.


In the meantime, there would be plenty of activities to keep ourselves occupied. With the amount of snow on the ground, one might think there would be little opportunity for botanizing. However, I favor the woody flora, and I was excited about the chance to begin reacquainting myself with some of the western conifers for a change. Of these, one of my favorites is incense-cedar (Calocedrus decurrens) — mature trees develop thick, deeply furrowed, brick red bark that stands out in beautiful contrast from the other trees. Even dead trees maintain a rustic and majestic beauty, and this large dead snag is as stately as any I’ve seen:


On Monday we rented snowshoes and hiked the cross-country ski trails at Camp Richardson. None of us had ever snowshoed before, but the girls quickly got the hang of it (note the live incense-cedar in the background):


We encountered a few cross-country skiers during our hike, but for the most part we spent the day in solitude. Shortly after beginning our hike, however, we came upon this impression in the snow. At first we thought someone had attempted to make a “snow angel,” but after studying it more carefully we realized it was made by a cross-country skier who had fallen and then struggled to get back up:


At this altitude, conifers dominate the flora. I was a little rusty on my knowledge of western U.S. plants, but I think I have things figured out (please let me know if you see any needed corrections to my identifications). The aforementioned incense-cedar was a conspicuous component of this lake-level forest, and its foliage – arranged in flattened, elongated, rumpled sprays – makes this tree easily identifiable amongst the other coniferous genera with which it grows:


Huge pine trees also dominanted the forest in this area. At first I thought they were ponderosa pines (Pinus ponderosa) due to their large size, irregular crown, and large plate-like patterns on the trunk caused by deep cross-checked fissuring of the bark. Eventually, however, I decided they must instead be Jeffrey pine (Pinus jeffreyi), a closely related species (that was once considered a variety of ponderosa pine), since the bark was more orange than yellow.


A closeup of the needles, which are in bundles of three and measure around 6-8 inches in length:


Another dominant coniferous component of this forest, also reaching massive size, was white fir (Abies concolor). The first photo below shows a large, mature tree in the distance, while the second shows a closeup of the foliage. At first I thought this might be Douglas-fir (Pseudotsuga menziesii), as the needles appeared to be irregularly 2-ranked; however, I asked Prof. Ronald Lanner to take a look, and he confirmed it is white fir. He said Douglas-fir needles are shorter, thinner, darker green, and have a skinny stalk, while fir needles have a fat round base and are wider and flatter. The latter also have a citrusy smell when crushed, which he describes as one of the best smells in the woods! Too bad I did not try it.



This decaying stump also represents white fir based on the scaly gray bark. I suspect the outer layers of the lower portion of the trunk (core still standing) were ripped off over time by animals looking for grubs and insects as decay progressed, eventually weakening it to the point that the upper portion (laying on the ground) finally broke off and fell:


As we hiked, I realized what an important part fire plays in the ecology of these forests. During the drive up from Sacramento, we passed several areas along Hwy 50 that had suffered severe damage due to the wildfires that swept through Lake Tahoe recently. One such area was even seen in the far eastern slopes of Heavenly Ski Resort itself. The forests around Camp Richardson had largely escaped these fires, and I wondered if fire management had contributed to this. Along the trail, evidence of fire was common on the trunks of trees, but few trees – even small ones – had been killed. I presumed the charring was evidence of fires that had been intentionally set and managed by the Forest Service with the objective of preventing fuel accumulation that could lead to the larger conflagrations that caused so much damage in other parts of the basin. These small incense-cedars trunks show obvious fire charring but otherwise looked healthy:


In a few areas it appears even these “cool” fires burned a little hot, killing some of the smaller trees but still avoiding the “torched-earth” damage seen in areas affected by uncontrolled burns:


I’m not much of a birder, but I do love woodpeckers. I got a glimpse of one during our hike, but I didn’t see it well enough to identify it. We did find this woodpecker hole in the trunk of a large, dead Jeffrey pine — a feather can even be seen clinging to the upper rim of the hole. The Lake Tahoe basin is home to several species of woodpeckers — whether this hole belongs to the black-backed woodpecker (Picoides arcticus), white-headed woodpecker (P. albolarvatus), or (more likely) hairy woodpecker (P. villosus) I can’t say for sure:


At the beginning of our hike, signs warning of bears and pleading not to feed them caught the girls attention. I told them it was winter and that they would be hibernating, but I wondered if at this late stage they might actually be starting to become active. It wasn’t long before we encountered these unmistakably bear tracks, made fresh in the new-fallen snow, and the more we looked the more abundant the tracks were to be found. I secretly (and the girls outwardly!) hoped we would see a live bear, but I don’t think the girls would have handled such an encounter very calmly:


I had intended to photograph some of the conifers seen at higher elevations while skiing at Heavenly Ski Resort, but I decided not to bring my camera. Pity, as I not only saw nearly pure stands of what I presume to be red fir (Abies magnifica), but also beautifully twisted and wind-gnarled pines at the highest elevations (+10,000 ft) that probably represent whitebark pine (Pinus albicaulis), judging by their highly forked trunks and upswept limbs. These magically grotesque trees were made even more beautiful by the previous day’s storms, which had deposited thick cakes of ice on their windward sides.

We coudn’t leave Lake Tahoe without one final visit to Emerald Bay. Below is a close up photograph of Fannette Island, the only island to be found in all of Lake Tahoe, and its famed “Tea House”:


We concluded our visit to Lake Tahoe by driving up Hwy 89 to Tahoe City for dinner at the Bridgetender Cafe before heading back to Sacramento. Next up — Muir Woods!