Revisiting the Swift Tiger Beetle – Part 3

…continued from Revisiting the Swift Tiger Beetle – Part 2.

The Oklahoma trip had been an unqualified success. Not only had I managed to find the rare Cylindera celeripes (swift tiger beetle) at Alabaster Caverns, I had also determined the population there was healthy and, in fact, occurred robustly across a large swath of red clay/gypsum hill habitat in the vicinity of the Cimarron River (Woodward and Major Counties) in northwestern Oklahoma. This is good news for the species, who some have regarded as a potential candidate for federal listing on the endangered species list. There is no doubt that the species has suffered greatly in many parts of its range during the past century – most likely due to loss of habitat; however, the presence of a strong population in Oklahoma gives reason for optimism about its long-term prospects. It would have been even better had I found the species at the two Nature Conservancy preserves (Four Canyon and Tallgrass Prairie) that I had targeted, and the reasons for its apparent absence at those two sites despite an abundance of apparently suitable habitat remain a mystery to me (although I have my suspicions). Nevertheless, I returned to St. Louis happy, with new localities in my database and live individuals in containers of native soil for another attempt at rearing.

Our work with this species was not done, however. While C. celeripes has never been recorded in Missouri, my colleague Chris Brown and I have long suspected that it might occur here – most likely, we felt, in extreme northwestern Missouri where the Loess Hills landform reaches its southern terminus. We had looked for it in this area a few times before on the few remaining dry, hilltop prairie relicts that are so common further north in Iowa, and we had also looked for it in the larger tallgrass prairie remnants of west-central Missouri. None of these searches were successful, and with each unsuccessful effort it seemed less and less likely that the species actually occurred within the state – especially considering the declines that the species has experienced throughout its range. However, when we managed to find a small, newly discovered population of the species last summer in the Loess Hills of southwestern Iowa, just 60 miles north of Missouri (see The Hunt for Cicindela celeripes), we decided that one more thorough effort to locate the species in Missouri was in order.

Star School Hill Prairie Natural Area (north tract), Atchison Co., Missouri.

Star School Hill Prairie Natural Area (north tract), Atchison Co., Missouri.

Our plan was straightforward – we would travel to northwestern Missouri each weekend beginning in late June and search the most promising hilltop prairie relicts that still remain in Missouri. There aren’t many of these, so I contacted Tom Nagel of the Missouri Department of Conservation – who probably knows more about Missouri’s hilltop prairie relicts than anyone else – for assistance in identifying these parcels. Tom graciously sent me descriptions and aerial photographs of the highest quality relicts still remaining in Missouri. None of these are large (12 contiguous acres or less), and all have been impacted to some degree by woody encroachment and are in various stages of restoration. We had already searched one of these tracts (Star School Hill Prairie) a few times, but two others were new to us. So, on a Friday evening before the first of three planned weekends for our study (and only two weeks after returning from Oklahoma), Chris and I made the long drive across Missouri and north along the Missouri River and began our search the next morning.

Fieldmate Chris Brown surveys loess hilltop prairie habitat at Star School Hill Prairie Natural Area, Atchison Co., Missouri

Fieldmate Chris Brown surveys loess hilltop prairie habitat at Star School Hill Prairie Natural Area (south tract), Atchison Co., Missouri

Our first stop was High Creek Hill Prairie in Brickyard Hill Conservation Area (Atchison Co.). We had been to Brickyard Hill a few times but had not previously found this particular hilltop prairie. We found the tract, a long, narrow series of ridge tops and southwest-facing slopes, thanks to Tom Nagel’s map and began searching with all the enthusiasm and optimism that accompanies any new search. Our optimism waned with each hilltop ridge that we traversed not seeing the beetle, until we reached the easternmost ridge amidst a jumble of eastern red-cedar cadavers that halted any further progress or promise. As we stood atop that last hill, we debated our next move. Chris had noted apparently good habitat on the lower slopes below us, while I had spotted another very small hilltop tract across a wooded ravine and disjunct from the main prairie. We decided these areas should be explored before moving on to the next site, but as we searched those lower slopes our optimism continued to wane. The habitat was perfect based on what we had seen in Iowa last year and what I had seen in Oklahoma earlier in the month – small clay exposures amongst clumps of undisturbed little bluestem and grama, but still no beetles. Chris, refusing to accept defeat, continued to search the slope, while I worked my way over to the smaller hilltop tract I had seen from above. After crossing through the wooded ravine, I found an old 2-track running along the base of the tract and began walking along it. The small slope above the 2-track was littered with large cadavers of the invasive eastern red-cedar (Juniperus virginiana), apparently left in place after chainsawing to provide fuel for a planned, future burn. As I walked, a white-tailed deer bolted from a nearby cadaver, giving me a bit of a start, and I veered towards the cadaver to have a look at where it had bedded down. By this time I almost wasn’t even really thinking about C. celeripes anymore – we had been there for about an hour and a half and searched the most promising habitats without success – the small tract where I was now working was almost a last gasp before moving on. As I approached the deer’s bedding site, a “flash” in the thick vegetation caught my eye, and I knew instantly what it was. Immediately I dropped to my knees and tried to “trap” the evasive little beetle (I’ve found that forming a “trap” between the crotches of my hands and gradually closing my hands together forces the beetle to run up and over one of my hands, at which time I can try to pin it down with my other). The beetle behaved exactly as expected, running over my left hand – but I missed it. I trapped it again, and once again it ran over my hand too fast to pin down. I tried to follow it as it zigzagged erratically through the thick vegetation, but in the blink of an eye it was gone. I spent the next several minutes frantically pulling apart the vegetation in a 2-foot radius around the spot in what I knew was a vain attempt to relocate the beetle before ultimately accepting that I had missed it. No matter – I had seen it and had absolutely no doubt about what it was – C. celeripes does indeed occur in Missouri! Wow – big news! I knew if I had seen one, I had a good chance of seeing another, so I began searching the area again – now with much more deliberation. I walked back and forth along the old 2-track, up and down the cadaver-littered slope, and back to the original spot several times. As time passed, a gnawing fear began to grow inside me that this new state record might lack a voucher. Suddenly, very near the original spot, I saw another. This time I pounced with authority and made no mistakes, and after securing the live beetle in a vial I gloated and congratulated myself unabashedly inside while bursting to give the news to Chris. I searched the slope some more, but I couldn’t take it anymore – I had to tell someone. I pulled out my cell phone and began texting a message to my daughter Mollie (who really doesn’t care about beetles but loves to receive text messages). As I was texting, Chris appeared on the lower slope, obviously noting that my net had been left on the ground purposely to mark a spot. As I finished texting I told Chris to come here, I wanted to show him something, and then non-chalantly handed him the vial. I would give anything to have a video of the look on Chris’ face as it changed from quizzical dumbfoundedness to shocked elation. Chris, too, had reached a low point in his optimism after thoroughly searching the previous slope without success, but now we were both as giddy as school boys – our long efforts had finally paid off with a new state record for one of North America’s rarest tiger beetles (the way we were acting, you’d have thought we’d just discovered plutonium!). We searched the slope for another half hour or so, with Chris seeing one more individual very close to where I had seen the first one. Whether it was the same or a different individual is unknown, so we decided that we had seen at least two individuals at this site. The discovery of C. celeripes here caused us to once again search the lower slope that Chris had previously searched so thoroughly, but again the beetle was not seen. Our giddiness was beginning to give way to concern over the few individuals we had seen and how localized they seemed to be. We had been at the site now for about three hours, and I was famished. I hiked back to the truck, noting some habitat at the far western end of the main prairie where we had begun our search that looked like it deserved another search. As I ate, Chris worked his way over to that spot, and after a period of time I heard him yell down to me and give me the “thumbs up.” I hurriedly finished eating and worked my way up to where he stood, and together we located two more individuals – taking one as a voucher for the site and ganging up on the other to keep it pinned into an open area where each of us could take field photographs before we finally let it “escape.” Seeing the species on the larger parcel had relieved our concern a little bit, and we felt a little less worried about its status here now.

Cylindera celeripes - High Creek Hill Prairie, Brickyard Hill Conservation Area, Atchison Co., Missouri (new state record)

Cylindera celeripes - High Creek Hill Prairie, Brickyard Hill Conservation Area, Atchison Co., Missouri (new state record)

Later in the day we would see the species again at Star School Hill Prairie Natural Area , the northernmost substantial loess hilltop prairie within Missouri, and one that we had searched at least twice previously for the species. Again, we saw only two individuals in almost three hours of searching, confirming the impression first gained at Brickyard Hill that the species is not present in very high densities. Like Brickyard Hill, the beetles at this site were found in areas of undisturbed hilltop prairie with moderately thick shortgrass vegetation and were seen only when they IMG_0789_1200x800ran from one grass clump to another after being disturbed by our approach. We also looked for it at a smaller disjunct parcel just to the north, but the lateness of the hour limited the time we had to explore this site. Star School Hill Prairie is some 6 miles north of Brickyard Hill, thus, finding C. celeripes at two sites not in close proximity increased our optimism that the species might actually occur in many of the loess hilltop prairie remnants still remaining in northwestern Missouri. This optimism was further increased the next day when we saw two more individuals at one of Missouri’s southernmost hilltop prairie relicts at McCormack Loess Mounds Natural Area in Holt Co. However, our optimism is tempered by the fact that, again, we saw only two individuals, both of which were seen in a small, unburned spur extending northward off the main prairie, while none were seen in the much larger main parcel that appeared to have been recently burned in its entirety.

Cylindera celeripes macrohabitat at Star School Hill Prairie.  Beetles were seen along the narrow trail in the foreground and on the mild upper slopes (below bur oak in upper left).

Cylindera celeripes macrohabitat at Star School Hill Prairie. Beetles were seen along the narrow trail (foreground) and on the mild upper slopes (below bur oak, upper left).

The presence of this rare Great Plains species in Missouri’s critically imperiled hilltop prairies is cause for both excitment and concern. Cylindera celeripes represents a unique and charismatic addition to the state’s rich natural heritage. However, like soapweed yucca (Yucca glauca var. glauca), skeletonweed (Lygodesmia juncea), and the dozen or so other plant and animal species of conservation concern found within the hilltop prairies of IMG_0774_1200x800Missouri’s Loess Hills, C. celeripes appears to be entirely dependent upon these habitats for its survival within the state. Ensuring its continued survival will require careful reconsideration of the management approaches used for these rapidly shrinking natural communities. Prescribed burning has been and will continue to be an important tool in restoring our hilltop prairies; however, nonjudicious use of fire could lead to local extirpaton of C. celeripes within these habitats. Should that occur, recolonization from nearby parcels is unlikely due to the small, highly disjunct, and upland character of Missouri’s hilltop prairie remnants and the flightless nature of C. celeripes. As a result, rotational cool-season burns should be utilized as much as possible to avoid localized extirpations, especially on smaller parcels (Panzer 2002).

Hilltop prairie at McCormack Loess Mounds Natural Area, Holt Co., Missouri.  The main tract (pictured) was recently burned - beetles were found in a small unburned spur (off left center).

Hilltop prairie at McCormack Loess Mounds Natural Area, Holt Co., Missouri. The main tract (pictured) was recently burned - beetles were found in a small unburned spur (off left center).

Photo details:
Beetles: Canon 100mm macro lens w/ 68mm extension on Canon EOS 50D, ISO 100, 1/250 sec, f/18, MT-24EX flash 1/2 power through diffuser caps.
Landscapes: Same except Canon 17-85mm zoom lens (17mm at Star School, 20 mm at McCormack), 1/60 sec, f/8-9 (Star School) or f/13 (McCormack), natural light.


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

Copyright © Ted C. MacRae 2009

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Four Canyon Preserve, Oklahoma


Looking SE into lower Horse Canyon towards Canadian River

On my recent week-long collecting trip, the first three days were spent at Four Canyon Preserve in far northwestern Oklahoma.  This nearly 4,000-acre preserve features a stunning landscape of rugged, wooded canyons dissecting ridges of mixed-grass prairie which provide critical habitat for several rare plants and animals.  Despite years of overgrazing, fire suppression, and invasion by exotic plants, The Nature Conservancy (TNC) recognized the restoration potential of this landscape and began management practices to restore its ecological function and integrity after acquiring it in 2004.  The land was rested until April 2008, at which time a wildfire swept through the area and burned approximately 90% of the property.  This event was actually welcomed by TNC, who was already in the process of initiating a prescribed burn – they simply pulled back and let it rip!  The burn, combined with mechanical removal of eastern redcedar (Juniperus virginiana) that had invaded many areas of the preserve, did much to confine woody growth to the canyons proper, and good rains during the past two springs following that burn have resulted in a lush, green, diverse landscape brimming with prairie wildflowers.  The vivid contrast between the green vegetation and the red clay canyons with their white gypsum exposures has created spectacular vistas of a rugged landscape.  This year, cattle have been reintroduced at low levels to simulate the irregular, patchy disturbance experienced in pre-settlement times when native grazers (bison and elk) dotted the landscape.

The flora (Hoagland and Buthod 2007) and avifauna (Patten et al. 2006) of the preserve are well characterized, but (as nearly always seems the case) arthropod and other micro faunas need much additional study.  My hymenopterist colleagues and I were welcomed enthusiastically by TNC staff, who are anxious to incorporate the results of our insect surveys into an overall fauna.  Apoid hymenopterans appear to have benefited greatly from the recent rejuvenation of the preserve’s floral character.  Results for the beetle populations that I encountered, however, were more mixed. Certain groups, such as scarabaeine dung beetles, were quite abundant and diverse (due to the reintroduction of cattle), but others, including the tiger beetles, jewel beetles, and longhorned beetles that I was most interested in finding, existed at rather low and not very diverse levels.  I had hoped to find the very rare Cylindera celeripes (swift tiger beetle) running amongst the clumps of vegetation on the preserve’s red clay exposures but instead saw only the ubiquitous Cicindela punctulata (punctured tiger beetle), and the few jewel beetles that I managed to beat off the lower branches of hackberry (Celtis laevigata) and soapberry (Sapindus saponaria var. drummondii) trees were found only in the small parts of the preserve that escaped last year’s burn.  This seems fairly typical – I generally don’t find many insects in these groups whenever I survey areas that have experienced a significant amount of recent burning.  Some ecologists might take exception to this statement, and they would have little difficulty citing studies that show rapid recolonization of prairies by a majority of prairie insect specialists within two years after a prescribed burn.   Nevertheless, the impact of prescribed burning on invertebrate populations and its potential for causing local extirpations has become a contentious issue among ecologists and entomologists in recent years.  While my experience hardly passes for rigorous investigation, I am becoming increasingly convinced that a certain amount of caution is warranted when designing burn management plans for prairie relicts.

I’ll discuss more about the beetles and other insects (and even some vertebrates) that I saw during my three-day visit to Four Canyon Preserve in future posts.  In the meantime, I share with you some of my photos of this spectacularly beautiful landscape (note the abundance of woody cadavers from last year’s burn in some of the photos).


Looking S into upper reaches of Mulberry Canyon


Looking S into upper reaches of Mulberry Canyon


Looking E across upper Harsha Canyon


Looking SE into Harsha Canyon towards Canadian River


Looking E across lower Harsha Canyon

View of Mulberry Canyon bluffs from Canadian River valley

Looking NE towards Mulberry Canyon bluffs from Canadian River valley


Hoagland, B. W., and A. K. Buthod.  2007.  Vascular flora of the Four Canyons Preserve, Ellis County, Oklahoma.  Journal of the Botanical Research Institute of Texas 1(1):655–664.

Patten, M. A., D. L. Reinking, and D. H. Wolfe.  2006.  Avifauna of the Four Canyon Preserve, Ellis County, Oklahoma.  Publications of the Oklahoma Biological Survey (2nd Series) 7:11-20.

Copyright © Ted C. MacRae 2009

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Saving endangered species with herbicides

ResearchBlogging.orgThe latest issue of Cicindela (a quarterly journal devoted to tiger beetles), which arrived in my mailbox last week, features an article coauthored by my good friends Kent Fothergill and Kelly Tindall of Portageville, Missouri, along with lead author Stephen Bouffard of Boise, Idaho (Bouffard et al. 2009).  The article reports the results of a vegetative management pilot test for using herbicides to restore habitat for the critically imperiled St. Anthony dune tiger beetle, Cicindela arenicola.  This species is endemic to Idaho, primarily the St. Anthony Dunes area in the southwestern part of the state (Pearson et al. 2006), and like the Coral Pink Sand Dunes tiger beetle (Cicindela albissima, recently covered in this post) it is restricted to sand dune habitats that are threatened by a variety of land-use practices, including motorized vehicle use, livestock trampling, intentional stabilization of dunes by grass seeding, conversion of dune habitats to agriculture, and disposal of public lands by transfer to private ownership (Idaho State Conservation Effort 1996).


Cicindela arenicola, copyright © Kent Fothergill 2008

Bouffard et al. conducted their study at Minidoka National Wildlife Refuge in southern Idaho. Cicindela arenicola was recorded on small remnant sand dunes within the refuge during the mid-1990’s but had not been detected in more recent opportunistic searches. The authors noted that the sand dunes appeared to have become overgrown with the invasive annual grass, downy brome (Bromus tectorum). Their study comprised three elements: 1) herbicide treatment on dune habitats to reduce downy brome density; 2) surveys of treated versus untreated plots during the following season to assess the efficacy of the herbicide in reducing downy brome density and any effect it might have on native vegetation, as well as the presence of C. arenicola; and 3) laboratory bioassays to evaluate the acute toxicity of herbicides on a surrogate tiger beetle species, Cicindela repanda (common shore tiger beetle). The laboratory bioassays were necessary, because toxic effects by a herbicide against tiger beetles would negate its potential usefulness for habitat improvement. For the herbicide treatment plots, Imazapic (trade name Plateau®) was selected because of its effectiveness against downy brome, minimal effects on native vegetation, and low toxicity to animals, including insects. Imazapic is labeled for control of downy brome and for use on rangeland. A nonselective herbicide, glyphosate (trade name Touchdown®) was also evaluated in the laboratory bioassay, even though it was not used in the field test, because glyphosate-based herbicides also have low animal toxicity and have been shown to be effective in assisting the establishment of native plant species in prairie restorations.

The authors were successful in observing live adult C. arenicola in both of the test plots where adults of this species were last seen in the mid-1990s. Moreover, larval burrows – putatively representing this species – were also noted in the plots. No adults or larvae were seen in a third plot; however, no previous records of the species exist in the area where that plot was located. They noted the presence of residual downy brome stems from the previous season’s treatment in the sprayed plots but no new growth, while the untreated controls exhibited extensive new downy brome growth. More importantly, no negative impacts on native vegetation – principally rabbitbrush (Chrysothamnus sp.) and Indian ricegrass (Achnatherum hymenoides) – were noted in the sprayed plots. The third plot had only a light downy brome invasion prior to treatment, and no apparent negative effects were observed on the native bunchgrasses, rabbitbrush, and sagebrush (Artemisia tridentata) in this plot after treatment. In the laboratory, neither imazapic nor glyphosate showed evidence of acute toxicity against the surrogate tiger beetle, C. repanda.

While the use of herbicides for conserving endangered species may seem counterintuitive, this study demonstrates a potential use for herbicides in restoring and improving sand dune habitat for a critically imperiled species of tiger beetle. Herbicides that are effective in reducing invasive annual grasses with minimal effects on both native vegetation and tiger beetles could greatly facilitate habitat management for a number of critically imperiled western U.S. sand dune tiger beetles besides C. arenicola, including C. albissima in southwestern Utah, C. waynei (Bruneau tiger beetle) in western Idaho, and C. theatina (Colorado Great Sand Dunes tiger beetle) in southern Colorado. Where vegetational encroachment presents a threat to critical sand dune habitat, broad spectrum or grass selective herbicides may offer an effective and convenient alternative to habitat restoration. Additional research will be needed to determine if repeat applications of herbicides will be necessary to prevent reinvasion, and if so with what frequency, as well as the chronic or behavioral effects of herbicides on both larval and adult forms of the insects targeted for conservation.

I thank Kent Fothergill for allowing me to use his beautiful field photograph of C. arenicola, which also graces the cover of the current issue of Cicindela.


Bouffard, S. H., K. V. Tindall and K. Fothergill. 2009. Herbicide treatment to restore St. Anthony tiger beetle habitat: a pilot study. Cicindela 41(1):13-24.

Idaho State Conservation Effort.  1996.  Habitat conservation assessment and conservation strategy for the Idaho Dunes Tiger Beetle.  Report No. 7, Boise, ID.

Pearson, D. L., C. B. Knisley and C. J. Kazilek. 2006. A Field Guide to the Tiger Beetles of the United States and Canada. Oxford University Press, New York, 227 pp.

Copyright © Ted C. MacRae 2009

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Blackjack oak “flower”


This blackjack oak (Quercus marilandica) was found on one of southeastern Missouri’s finest sand prairie relicts a couple of weeks ago on my ‘Annual Birthday Season Opener Bug Collecting Trip.’ Growing near the edge of the prairie at the transition to dry sand forest (Nelson 1985), the arrays of soft, red, newly-expanding leaves at each branch tip had a distinctly floral quality to them. Of course, as with all oaks, the actual flowers of blackjack oak are much less conspicuous, with the staminate (male) flowers borne on drooping catkins, the pistillate (female) flowers on separate spikes on the branch, and pollination accomplished by wind.

Missouri is oak country – nearly a quarter of North America’s 90 oak species (Nixon 2009) occur naturally within the state. This high diversity is explained partly by Missouri’s ecotonal continental position – straddling the east-west transition from the great eastern deciduous forest to the western grasslands. The boundary between these two great biomes is a dynamic, ever-changing interdigitation of woodland, savanna, and prairie that ebbs and flows with the prevailing climatic conditions. Unlike the more mesic forests further east, these dry woodland habitats were often subjected to fire during presettlement times – to which oaks in general (and blackjack oak in particular) are supremely adapted with their thick bark and ability to resprout repeatedly after being burned or grazed back. Sadly, the suppression of these fires post-settlement has caused many of these unique, fire-mediated natural communities to shrink drastically amidst a choking growth of junipers (“cedars” ’round these parts), maples, and other fire-intolerant species. Only on publicly owned preserves and a few private parcels under progressive ownership (such as the sand prairie relict where this photograph was taken) is fire once again shaping the landscape.

Oaks are among my favorite trees, and among the oaks I have several favorites. White oak (Quercus alba) – tolerating many forest types but forming nearly pure stands in high-quality, mesic sites, its tall symmetrical crown, pale bark, and brilliant fall colors are unparalleled among Missouri’s other oaks. Post oak (Q. stellata) as well – lacking the elegance of white oak but achieving its greatest character in fire-adapted savannas and open woodlands as squat, gnarled, massively-trunked trees with broad, spreading crowns¹. Blackjack oak has none of these qualities, yet somehow, it is still one of my favorite Missouri oaks. Stunted and gnarled (‘scrub oak’ to some), it occurs mostly in sandstone and limestone glades, savannas and woodlands on dry, nutrient-poor soils that support few other tree species. The dark green of its tough, waxy (to limit the loss of water), pear-shaped leaves contrasts beautifully with its rough, blocky, almost black bark. Blackjack oak has virtually no timber value, although it is sometimes used for charcoal and firewood. Nevertheless, for me, it is almost an icon for the unique natural communities in Missouri in which it occurs – communities that face ever-increasing pressure from human and forest encroachment.

¹ Please refer to this lovely essay about post oaks in Missouri, by the talented Allison Vaughn.


Nelson, P. W. 1985. The Terrestrial Natural Communities of Missouri. Missouri Natural Areas Committee, Jefferson City, 197 pp.

Nixon, J. C.  2009. Quercus in Flora of North America, Vol. 3.

Copyright © Ted C. MacRae 2009

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Trees of Lake Tahoe – The “Other” Conifers

The inviting openness of the Sierra woods is one of their most distinguishing characteristics. The trees of all the species stand more or less apart in groves, or in small, irregular groups, enabling one to find a way nearly everywhere, along sunny colonnades and through openings that have a smooth, parklike surface.–John Muir, The Mountains of California (1894)


In a previous post (Trees of Lake Tahoe – The Pines), I discussed the six species of pine that can be found within the Tahoe Basin. These include Jeffrey pine (Pinus jeffreyi) – dominant around the lake at lower elevations, lodgepole pine (P. contorta ssp. murrayana) – common in meadows at lower elevations and replacing Jeffrey pine at higher elevations, ponderosa pine (P. ponderosa) – uncommon in the basin due to its preference for lower elevations, sugar pine (P. lambertiana) – the magnificent giant of high quality mesic sites along the western shore, western white pine (P. monticola) – co-occurring with lodgepole pine at higher elevations, and whitebark pine (P. albicaulis) – covering the highest peaks with its gnarled and twisted form.  In this post, I will cover the five “other” coniferous trees that can be found growing in the Tahoe Basin.  These other conifers belong to several different genera in two gymnospermous families – the Pinaceae and the Cuppressaceae.  Together with the pines, these trees comprise what John Muir described as one of the most diverse and appealing coniferous forests in the world. I am most inclined to agree with him.  The diversity of conifers found in the Tahoe Basin is reflective of the wide range of conditions occurring there as a result of differences in elevation (from 6,200 ft to more than 10,000 ft), exposure, and moisture.

Family-level identification of Tahoe Basin conifers is relatively straightforward – those with needle-shaped leaves belong to the Pinaceae (the pine family), while those with scale-like leaves belong to the Cuppressaceae (the cypress family).  There are other characters that distinguish members of these two families, but leaf shape is the most useful for purposes of field identification.  Nine of the eleven species of conifers found in the Tahoe Basin belong to the Pinaceae, while only two are members of the Cuppressaceae.  Within the families, the genera can be distinguished most readily by the following characters:


  • Pines (Pinus) – needles linear, arranged in bundles or clusters of up to 5 needles held together at the base by sheath of papery bark (discussed in Trees of Lake Tahoe – The Pines).
  • Firs (Abies) – needles more or less flattened, growing directly and singly from the branch and with a plump base that leaves a round depression on the branch.  Cones upright, on upper branches.
  • Hemlocks (Tsuga) – needles more or less flattened and growing directly and singly from the branch like firs, but narrowly stalk-like at the base where they are joined to tiny wooden pegs.  Cones pendant, on outer branches.


  • Incense-cedars (Calocedrus) – scale-like leaves 4-ranked, twigs branching in one plane to form flat sprays, cones > ½” in length, consisting of two large scales separated from a closed center.
  • Junipers (Juniperus) – scale-like leaves arranged in circles of 3, twigs not forming flat sprays, cones < ½” in length, berrylike.

There are three additional coniferous genera in the Sierra Nevada – each represented by a single species and found along the western slope – that do not occur in the Tahoe Basin.  These include: Douglas-fir (Pseudotsuga menziesii) – widespread at elevations from 2,500 ft to 6,000 ft (higher at the southern end of its range); giant sequoia (Sequoiadendron giganteum) – primarily in Giant Sequoia National Monument, and California nutmeg (Torreya californica) – of scattered occurrence.

White fir (Abies concolor)

As old age creeps on, the bark becomes rougher and grayer, the branches lose their exact regularity, many are snow-bent or broken off,…but throughout all the vicissitudes of its life on the mountains, come what may, the noble grandeur of the species is patent to every eye.


White fir is second only to Jeffrey pine as the dominant conifer at the lower elevations within the Tahoe Basin¹.  It is immediately recognizable as the only non-pine member of the Pinaceae to occur at these elevations – red fir and mountain hemlock are found only at higher elevations in the basin.  Young trees have a nearly perfect pyramidal shape, with silvery gray bark that is thin, smooth, and p1020588_2covered with resin-filled blisters that can be “popped” to shoot out the resin. Older trees develop a more cylindrical and slightly irregular crown, and the bark becomes thick and roughly furrowed, changing to a dark gray or brown color. The foliage has a gray frosted appearance from below, and crushing the needles releases a delightful citrus smell that I found myself partaking in repeatedly. In the narrow elevational zone where white fir and red fir co-exist, white fir may be recognized by its more flattened needles (cannot be “rolled” in the fingers) which are distinctly twisted near the base, causing them to appear 2-ranked. White fir was seen throughout the Tahoe Basin at elevations below around 7,500 ft, and especially along the western shore and southern shores where the greater moisture and protection of north and east facing slopes are to this species liking.

¹ This post by Watching The World Wake Up provides an excellent introduction to the characteristics and distribution of white fir and its relatives. It also contains what must be the best tangent to ever appear in a botanical blog – the connection made between white fir and the alluring Salma Hayek (annoyingly mispelled “Selma” Hayek), softly singing Siente Mi Amor, is pure brilliance!

Despite its “noble grandeur,” white fir may be regarded as somewhat of a pest species. The suppression of fires that have been the hallmark of 20th century forest management have encouraged the replacement of pines throughout the Sierra Nevada by this species. White fir does not tolerate fire as well as the pines with which it occurs, but unlike those species it does well in shadier conditions. The suppression of fires has resulted in dense stands of white firs growing up in the spaces between the pines. Since it tends to retain its lower branches as it grows, when fires do occur the white firs can act as “fire ladders” that allow the fires to reach the upper canopies of the pines. Pines are not as shade tolerant as firs and are thus unlikely to become established beneath the dense canopy of firs. The result of these fire suppression policies are mixed-conifer forests that are denser and contain a much higher proportion of white fir than in the past, making the forests more vulnerable to stand-replacing fires as well as stress-induced insect and disease outbreaks. These counterproductive management policies are beginning to change – and I saw two controlled burns taking place during the week while I was in Lake Tahoe – but there is still much progress yet to be done if we are to once again see large expanses of the “inviting openness” that so captivated John Muir.

Red fir (Abies magnifica)

This is the most charmingly symmetrical of all the giants of the Sierra woods, far surpassing its companion species [white fir] in this respect… Happy the man with the freedom and the love to climb one of these superb trees in full flower and fruit.

p1020785_2I suspected I had seen this magnificent relative of the white fir in the higher elevations at Heavenly Ski Resort on my first trip back to the area last year, but lacking any real knowledge or field guides at the time it remained only a suspicion. When I returned to Heavenly this year, I was ready for it, and I recognized it instantly when I reached elevations around 8,000 ft. The massive trees with deeply reddish bark were unmistakable, and my only disappointment in seeing this species was that I was unable to approach them closely enough to allow a more thorough examination of their needles and bark. Like the white firs I saw at lower elevations, these massive trees had developed a bit of irregularity in their long, cylindrical crowns.

Younger trees can appear more similar to white fir because of their thin, smooth gray bark with elliptical resin blisters. However, in trees both young and old, the foliage is a more boldly colored blue-green than the paler foliage of white fir. p1020784_2Both species develop thick, deeply furrowed bark as they age, but the bark of red fir is distinctly reddish-brown or reddish purple, compared to the dark gray or brown bark of white fir – almost ashen in appearance. In the hand, the needles are not so flattened as white fir – almost quadrangular in cross-section and able to be rolled in the fingers – nor are they distinctly twisted near the base. The photo at right shows a stately red fir on the left next to a Jeffrey pine on the right at Lakeview Lodge on the California side of Heavenly (elevation 8,250 ft – the highest at which I saw the latter species).  I found this species growing in the company of western white pine (Pinus monticola), lodgepole pine (P. contorta ssp. murrayana), and mountain hemlock (Tsuga mertensiana), as well as in groves of its own kind (unfortunately, seen only from my perch upon a ski lift).

Mountain hemlock (Tsuga mertensiana)

The Hemlock Spruce is the most singularly beautiful of all the California coniferæ. So slender is its axis at the top, that it bends over and droops like the stalk of a nodding lily. The branches droop also, and divide into innumerable slender, waving sprays, which are arranged in a varied, eloquent harmony that is wholly indescribable.

p1020804_2I hadn’t a clue whether I would succeed in finding mountain hemlock – I knew it was a denizon of the snowy high mountains, though less common than some of the other high country conifers, and I didn’t recall noticing anything that might be this species during last year’s visit to the slopes of Heavenly. Of course, being a long-time resident of the Midwest I have little experience with hemlocks in general – eastern hemlock (T. canadensis) is on occasion planted in urban landscapes here, but mountain hemlock is markedly different from that species, as well as its Pacific counterpart western hemlock (T. heterophylla), due to its needles growing out of the twigs in all directions rather than in two flat planar sprays. Additionally, the needles are square in cross-section like spruce (Picea), a genus that does not now occur in the Sierra Nevada. These features caused 19th century botanists to suspect that mountain hemlock might have originated from an intergeneric hybridization event, as evidence by John Muir’s reference to it as “Hemlock Spruce.” However, no crosses between genera in the Pinaceae have ever been substantiated, and no compelling evidence of the presumed crossing events proposed for mountain hemlock has been brought forth (Lanner 1999).

p1020803_2Perhaps being primed by the readings I had done beforehand, I knew instantly I had found this species while riding a ski lift and seeing what looked at first like small junipers, but with a Tolkienesque appearance due to the gracefully nodding leader and drooping branch tips.  My hurried attempts to snap photographs of the trees from the moving ski lift produced nothing but skewed views marred by lift cables and passing cars, but once at the summit I was able to ski down to a little grove next to the ski run for closer inspection.  I immediately noticed the many cones clustered at the branch tips and was struck by their pine cone-like appearance. They were quite large – nearly 2” long (massive by hemlock standards).  Sadly, the only examples I would see of this species would be these small trees that only hinted at the charms of the massive specimens with trunks up to six feet in diameter that so enamoured John Muir.  Like the rare Washoe pine (Pinus washoensis) that occurs just outside Tahoe Basin on the eastern slopes of Mt. Rose, attempts to find some of these graceful 100-footers will have to await my next year’s visit.

Incense-cedar (Calocedrus decurrens)

Casting your eye over the general forest from some ridge-top, the color alone of its spiry summits is sufficient to identify it in any company.


The incense-cedar is my favorite of all the Tahoe Basin conifers. The bright, cinnamon-red bark of mature trees, deeply-furrowed, fibrous and peeling, is reminiscent of California’s two most iconic conifers – redwood (Sequoia sempervirens) and giant sequoia (Sequoiadendron giganteum), respectively the world’s tallest and most massive trees. Incense-cedar is neither as tall as redwood nor as massive as giant sequoia – p1020590_2indeed, it is not even very closely related (redwood and giant sequoia belong to yet another coniferous family, the Taxodiaceae, containing also the graceful but much smaller resident of southeastern U.S. swamps, baldcypress – Taxodium distichum). Nevertheless, old trees – veterans of centuries of fires and storm damage – are stunning specimens to behold, their massive, buttressed trunks often draped in yellow-green mosses and bearing deep basal fire scars, their spired crowns often broken and forked.  Their flattened sprays of foliage give the tree a delicate, lacy appearance in beautiful contrast to its grizzled, gnarled bark. Indeed, even in death these trees stand out for their stark beauty.

Incense-cedar is common at lower elevations in the Tahoe Basin, especially down close to the lake and in the communities ringing the shore. It rarely forms “stands” like white fir and the pines, p1020670_2but rather most often occurs singly – as if to emphasize their distinctiveness. I found it most common along the western shore, where it grows scattered amongst white fir and Jeffrey, sugar, and ponderosa pines. Some of the most massive incense-cedars I have ever seen were found down near the lakeshore along the Rubicon Trail in Emerald Bay State Park. Common on these trees were what I take to be incense-cedar mistletoe (Phoradendron libocedri) (family Santalaceae), which is apparently rare in the Tahoe Basin but known to occur in the mesic forests of the west shore.

Incense-cedar is another of the so-called “wrongly named” conifers – it is not a true cedar (thus, the hyphen in the name), a group of conifers belonging to the genus Cedrus in the family Pinaceae that is found across Eurasia². While somewhat resembling the true cedars, incense-cedar’s closest relatives are restricted to China and Taiwan. p1020640_2Early botanist-explorers, when they first encountered this tree, named it for what it most resembled to them – the old world cedars. This distinctiveness makes older trees the easiest Tahoe Basin conifer to identify. Even it’s cones that litter the ground under mature trees are unique – slender, spindle-shaped, and about an inch long, with the two longest scales bending back at maturity in a manner resembling a wide-open duck’s bill with the tongue sticking out. Young trees resemble Sierra juniper by their scale-like leaves and peeling bark, but the flattened, yellow-green sprays of incense-cedar and shiny reddish coloration of the bark of twigs and younger branches are immediately distinctive.

² There are actually numerous examples of such wrongly named conifers – Douglas-fir (Pseudotsuga menziesii) is not a true fir; eastern redcedar (Juniperus virginiana), western redcedar (Thuja plicata) and Alaska-cedar (Chamaecyparis nootkatensis) are not true cedars; and baldcypress (Taxodium distichum) is not a true cypress. Long live scientific names!

Like white fir, the Sierra Nevada has seen a bit of a population explosion of incense-cedar due to the fire-suppressive forest management practices of the past century. Despite the thick, fire-resistant bark of older trees, the thin-barked seedlings and saplings are intolerant of fire and grow more slowly than the fire-adapted pines. As a result, the frequent low-intensity fires of the past kept seedling establishment to a minimum, resulting in spot occurrences of mature, fire-resistant specimens. The suppression of these fires, combined with the ability of incense-cedar to germinate in shade and thick layers of duff, have allowed this species to increase in incidence throughout the Sierra Nevada. Along with white fir, it is gradually replacing the pines. This may seem like a good thing from the perspective of foresters and loggers, who value the wood of incense-cedar for its use in making pencils and cedar chests, but from an ecological perspective this has the same negative consequences discussed above for white fir.

Sierra juniper (Juniperus occidentalis ssp. australis)

Its fine color and odd picturesqueness always catch an artist’s eye, but to me the Juniper seems a singularly dull and taciturn tree, never speaking to one’s heart.


This was another conifer that I didn’t recall seeing on my previous visits, but from what I had read I really hoped I did. Gnarly and burly, mature specimens have a weather-beaten, picturesque quality that is unmatched by any other Tahoe Basin conifer save whitebark pine (P. albicaulis). While I did not find this tree to be common in the Tahoe Basin, I did find it in the most surprising of places – Emerald Bay overlook, where I had gazed in admiration at Lake Tahoe on so many previous occassions. This enduring dweller of exposed granite crags grows where no other trees can, anchored to crevices with only the tracest amounts of soil, p1020613_2seemingly thriving on nothing more than rock, snow, and sunshine. Old trees, with their massively short trunks supporting wind-pruned crowns, cannot be mistaken for any other Tahoe Basin conifer. The wood, it seems, is almost as hard as the granite upon which the trees grow, accounting for John Muir’s impression of this tree as without expression – not even the strongest Sierra winds evoke the slightest of shudders or the quietest of whispers in its unyielding bows.

I did not find this species commonly in the areas of the Tahoe Basin that I visited (which were mostly lower elevation sites below 7,000 ft). In addition to the specimens seen at Emerald Bay State Park, I also found this species near Upper Truckee River before the climb to Echo Summit, and I found a number of fine mature specimens outside of the basin proper at Pyramid Creek Geological Area.  Where I did find it, Jeffrey pine was the most common associate, but in most cases the trees stood alone in their own starkness.  Among the Tahoe Basin conifers, the small scale-like leaves are recognizable to almost any easterner as those of juniper, immediately placing it in the family Cuppressaceae alongside incense-cedar.  Even the young trees can be distinguished from that species by their non-glossy foliage borne on twigs that radiate out from the branches in all directions.  p1020602_21The bark of young trees is shreddy and peeling like that of incense-cedar, but it is dull brown to reddish-brown rather than the shiny purple-red color of incense-cedars.

Sierra Nevada populations of Juniperus occidentalis are considered a separate subspecies due to differences in reproduction and elevational preference.  Trees in nominotypical populations, found in northeastern California and up through eastern Oregon and Washington, are found at somewhat lower elevations (4,000 ft to 5,000 ft) and have cones of both sexes on the same tree; while those of subspecies australis, limited to higher elevations (usually from 6,500 ft to over 10,000 ft) in the Sierra Nevada, have either all male cones or all female cones.


Arno, S. F. 1973. Discovering Sierra Trees. Yosemite Association, Yosemite National Park, California, 89 pp.

Graf, M. 1999. Plants of the Tahoe Basin. Flowering Plants, Trees, and Ferns. A Photographic Guide. California Native Plant Society Press, Berkeley, 308 pp.

Muir, J. 1894. The Mountains of California. The Century Co., New York, xiii+381 pp.

Lanner, R. M. 1999. Conifers of California. Cachuma Press, Los Olivos, California, 274 pp.

Peterson, P. V., and P. V. Peterson, Jr. 1975. Native Trees of the Sierra Nevada. University of California Press, Berkeley, 147 pp.

Copyright © Ted C. MacRae 2009

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Illinois Butterfly Monitoring Network

For those of you who plan to be in the Chicago area on Saturday, March 7th, perhaps you’ll be interested in attending the 2009 Illinois Butterfly Monitoring Network Annual Indoor Workshop. I’ll be giving a talk entitled, “From Hilltops to Swamps: Insects in Missouri’s Rarest Prairies,” in which I’ll focus on the natural history and some associated insects in two of Missouri’s most critically imperiled natural communities – the loess hilltop prairies in the northwestern corner of the state, and the sand prairies of the southeastern lowlands. How a beetle guy ended up being invited to talk to a butterfly group is still a little confusing to me, but apparently IBMN Director, Doug Taron (author of Gossamer Tapestry) put in a good word for me.

The Illinois Butterfly Monitoring Network (IBMN) is a citizen scientist program monitoring the health of butterfly populations throughout northeastern and central Illinois.

The IBMN was initiated in 1987 by The Nature Conservancy to explore the effects of habitat management on invertebrates. From 7 sites in the Chicagoland area in its first year, the program has expanded greatly and is now monitoring more than 100 sites throughout Illinois.  Butterflies are ideal “indicator organisms” with which to monitor the effects of prescribed burning and other management techniques, since many species are restricted to intact prairie and savanna remnants by narrow habitat requirements.  The fact that they are relatively easy to identify allows them to be monitored in a cost effective manner with the help of dedicated amateurs.  Much the same can be said for tiger beetles (which will – surprise! – be featured prominently my talk).

The workshop will be held Saturday, March 7, 2009, 9:30 AM until 3:00 PM at the Gail Borden Public Library, 270 North Grove Avenue, Elgin (directions).  Registration is required, contact Mel Manner at (847) 464-4426 or by email.

Copyright © Ted C. MacRae 2009

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Sanctuary for the Betulaceae

Nestled on the eastern side of the St. Francois Mountains, where the craggy exposures of the Ozarks most ancient rocks begin to subside underneath the Cambrian sandstones laid down over them, lies Hawn State Park – considered by many to be the loveliest of Missouri’s state parks. I have written previously about Hawn – in fact, it was the subject of my very first post on this blog. I have long treasured Hawn for its excellent insect collecting, diversity of plants and habitats, and unbridled beauty. I have hiked the incomparable Pickle Creek and Whispering Pine Trails many times – far more than any other trail in the state, and each time I fall more deeply in love with what, to me, represents the essence of the Missouri Ozarks in their most pristine state.

Lamotte sandstone cutThe charm of Hawn results from a unique combination of geological features. The Lamotte sandstone outcrops that dominate Hawn’s landscape are the oldest sedimentary rocks in the state, formed from coarse sand deposits that were laid down over the Precambrian rhyolites and granites that form the core of the St. Francois Mountains. These sand deposits were themselves buried under limestone and dolomite layers formed at the bottom of vast seas that later covered much of the interior of the continent. Subsequent periods of uplift and erosion once again exposed these sandstones, whose unique ability to hold groundwater has resulted in the formation of spring-fed streams that have cut deep into their soft layers to create canyon-rimmed valleys with tall vertical cliffs. rhyolite shut-ins One of these streams is Pickle Creek, which is fed throughout the year by Pickle Spring and has in some places cut all the way down to the underlying igneous rock to form “shut-ins.” In contrast to the slow, sandy bottomed stretches where Pickle Creek is still cutting through sandstones, the water in these igneous shut-ins rushes through narrow openings in the highly resistant rock. The igneous and sandstone exposures found in Hawn are spectacularly beautiful and support a unique flora due to the acid soils they produce. One group of plants that have taken sanctuary in these moist, acid soils is the Betulaceae, or birch family. Missouri is home to five native species of Betulaceae¹, and while none of them are extraordinarily uncommon they are limited in their occurrence to natural communities with sufficient moisture and exhibit a clear preference for acidic soils. This confluence of conditions occurs perfectly along Pickle Creek, allowing all five native species to grow here side-by-side – a betulaceous “hot spot” that represents not only the full diversity of the family in Missouri, but also the total generic diversity of the family in North America. In fact, only one other genus (Ostryopsis, shrubs related to Corylus and restricted to China) is assigned to the family on a global basis (Furlow 2004).

¹ Dr. George Yatskievych, in his recently published Steyermark’s Flora of Missouri (2006), regarded the presence of Corylus cornuta in Missouri as unlikely despite earlier reports of such. Dr. Yatskievych also recorded a single escape of the European species Alnus glutinosa from Springfield, Missouri.

The Betulaceae are deciduous trees and shrubs that occur primarily in the boreal and cool temperate zones of the Northern Hemisphere, although outposts are also known from high elevations in the Neotropics and, as mentioned above, China. Fossils of this ancient lineage of flowering plants are traceable to the late Mesozoic (upper Cretaceous), and the family appears to form a clade with hamamelidaceous plants. As would be expected from a group with boreal affinities, most species exhibit adaptations for survival in cold climates, such as small stature, shrubby growth habits, and small leaves. Several of Missouri’s species have performed well and gained acceptance as ornamental trees and shrubs, while others are important as sources of hazelnuts (genus Corylus) or ecologically for their ability to fix nitrogen (genus Alnus). My interest in these plants has nothing to do with their economic importance, but rather in their role as host plants for several rarely encountered species of woodboring beetles. Often, insects in this group may be collected on foliage of their hosts during the summer, making host identification fairly easy due to the presence of leaves. This is not always possible, however, due to limited periods of adult activity or low population densities. Rearing these insects from their hosts provides additional opportunity to document their occurrence, and winter is often the best time to collect the dead branches in which they breed, since by that time they have nearly completed their development and will be ready to emerge as soon as temperatures rise during spring. Identifying woody plants without foliage can be a challenge, but the ability to distinguish host plants by non-foliage characters such as bark, growth habit, bud shape, etc. greatly facilitates studies of wood boring beetles through rearing. In the past I have relied heavily on Cliburn and Klomps’ (1980), A Key to Missouri Trees in Winter, which utilizes mostly details of the twigs and buds to discriminate among Missouri’s 160+ species of trees. However, after a certain level of familiarity is gained, one eventually learns to recognize winter trees and even downed logs or fallen branches simply by their “look”.

Betula nigra - habit

Betula nigra - habit

Betula nigra - old bark

Betula nigra - old bark

Betula nigra - sapling

Betula nigra - sapling

Betula nigra (river birch) is the only member of this largely boreal genus found in the middle and southern latitudes of the U.S. and, thus, cannot be confused with any of Missouri’s other betulaceous species². It is the largest of the five and, along with the following species, is the most demanding in terms of keeping its “feet” wet. Trees are usually encountered right at the water’s edge, with tall, slender, often twisted or leaning trunks. Young trees and large branches on older trees exhibit gorgeous reddish brown bark peeling in thin, papery sheets, becoming thick and scaly on the main trunks of older trees. Small branches are dark, purplish brown in color with smooth bark and distinctly horizontal lenticels.  I have reared a small jewel beetle from fallen, dead branches of this tree collected at several locations in Missouri – this beetle turned out to be new to science, which I described and named Agrilus betulanigrae in reference to its (then) only known host (MacRae 2003).  I have also reared tremendous series of another jewel beetle, Anthaxia cyanella, which at the time was not known to utilize this host and was considered uncommon.  As it turns out, Betula nigra is its preferred host, and the rearing of large series from many locations resulted in improved knowledge about color forms and variability in this species (MacRae & Nelson 2003).

² The widely planted but dreadfully non-adapted Betula pendula (European white birch) and B. papyrifera (paper birch) can be recognized by their distinctly white bark. These species are limited to urban landscapes where they rarely achieve significant stature before declining and eventually succumbing to insect pests such as Agrilus anxius (bronze birch borer). River birch provides an equally attractive and much more durable choice!

Alnus serrulata - habit

Alnus serrulata - habit

Alnus serrulata - sapling

Alnus serrulata - sapling

Alnus serrulata - old cones

Alnus serrulata - old cones

Alnus serrulata (common alder, hazel alder, smooth alder, tag alder…) also demands to be next to (or even in) the water.  Unlike B. nigra, however, this species rarely reaches true tree status, instead usually forming shrubby thickets along the water’s edge.  Saplings can resemble those of B. nigra due to their smooth brownish bark, but the latter is usually more purplish, and the lenticels of A. serrulata are not distinctly horizontal as in B. nigra. The large purple-red buds also differ from the small brown buds of B. nigra, and during winter A. serrulata is adorned with numerous staminate catkins.  The persistent woody cones also cannot be mistaken for those of any other species of Betulaceae in Missouri. Associated with this plant is the longhorned beetle, Saperda obliqua, which reaches its southwesternmost distributional limit in Missouri on the basis of a single specimen collected some 25 years ago right here along Pickle Creek and given to me by lepidopterist George Balogh. Numerous attempts to find this species here since then have not (yet!) been successful.

Carpinus caroliniana - habit

Carpinus caroliniana - habit

Carpinus caroliniana (blue beech, hornbeam, musclewood) is one of my favorite betulaceous species. The beautifully fluted trunks and smooth, light gray bark are remniscent of the limbs of a sinewy, muscular person – every time I see this tree I cannot resist the temptation to grab and stroke the hard limbs (should I be admitting this?). This character begins to show even in very young trees, making its identification during winter quite easy. These trees also like to be near water, but they are not so demanding to be right at the water’s edge as are the previous two species. They usually form small trees, often in clumps with multiple trunks.  There are some notable insect associations that I’ve found with this plant.  One is a small jewel beetle, Agrilus ohioensis, which I reared from dead branches of this plant collected along Pickle Creek (Nelson & MacRae 1990), and which after more than 20 years still remain the only known Missouri specimens of this species.  Another is the longhorned beetle, Trachysida mutabilis, a single adult of which I reared from a dead (almost rotting) branch of this plant collected not too far from Pickle Creek in Iron Co.  This beetle also is the only representative of its species known from Missouri (MacRae & Rice 2007).

Ostrya virginiana - habit

Ostrya virginiana - habit

Ostrya virginiana - trunk

Ostrya virginiana - trunk

Ostrya virginiana (hop hornbean, American hornbeam) has a form and growth habit very similar to C. caroliniana, but its leaves that persist through the winter make it instantly recognizable from afar.  In Missouri, this habit is most often seen with the oaks (Quercus spp.).  This species can be found even further away from the water than the previous species, and its small stature combines with the orangish, persistent leaves to form a distinctive understory layer during winter.  Also, in contrast to the smooth gray bark of Carpinus, this species exhibits scaly, light reddish brown to brownish gray bark.  I have succeeded in rearing one of the two known Missouri specimens of another jewel beetle, Agrilus champlaini, from O. virginiana collected along Pickle Creek (the other specimen was reared from wood collected at Graham Cave State Park, another site where sandstone bedrocks favor an O. virginiana understory).  Unlike most other jewel beetles, A. champlaini forms galls in small living branches of its host.  I have collected the distinctive swellings during winter on many occasions but managed to rear only these two individuals (plus one ichneumonid parasitoid).  I have also noted similar swellings on Carpinus but have not yet managed to definitely associated them with this beetle.

Corylus americana (hazelnut, American hazelnut) is the smallest of Missouri’s five betulaceous species, always forming shrubs, sometimes in thickets, and never assuming the form of a tree. Its staminate catkins present during winter immediately identify plants of this species as Betulaceae, but the small, globe-shaped buds are unlike the more pointed buds of Ostrya and the elongated, reddish buds of Alnus. This species is the least demanding in terms of being near water and can be found even in upland prairies and glades. I haven’t yet associated any woodboring beetles with this plant in Missouri, but there are several jewel beetles known from the eastern U.S. that utilize Corylus (Agrilus corylicola, A. fulgens, and A. pseudocoryli) and could occur in Missouri.

pine savanna - fire managementThe upland habitats at Hawn are of interest as well. Lamotte sandstones are the dominant bedrock, creating acid soils that support a canopy dominated by Missouri’s only native species of pine, Pinus echinata (shortleaf pine), several species of oak, and a diversity of acid-loving shrubs primarily in the family Ericaceae (including the stunningly beautiful Rhododendron prinophyllum, or wild azalea). Historically, so-called “pine savanna” was prevalent in this area, a natural community in which periodic fires maintained an open structure amongst the fire-adapted pines and allowed a diverse herbaceous layer beneath the open canopy. Much of Hawn has closed up after decades of fire suppression; trail through pine savannahowever, the Department of Natural Resources has implemented a rotational burn management regime to recreate pine savanna habitat within Hawn’s Whispering Pines Wild Area. Evidence of what appeared to be very recent burns could be seen at several places as I hiked along the Whispering Pines Trail, and while many visitors might have been alarmed at the apparent “damage” they were observing, my heart sang with the prospect of seeing mature pine savanna communities taking hold throughout my beloved Hawn. As I stood atop this ridge and looked back down from where I had come, I could almost see Henry Schoolcraft and Levi Pettibone in the distance on horseback, perhaps pausing to gaze at an elk.


Cliburn, J. and G. Klomps. 1980. A Key to Missouri Trees in Winter, 2nd edition. Missouri Department of Conservation, Jefferson City, 43 pp. (subsequently revised)

Furlow, J. J.  2004. Betulaceae in Flora of North America @

MacRae, T. C. 2003. Agrilus (s. str.) betulanigrae MacRae (Coleoptera: Buprestidae: Agrilini), a new species from North America, with comments on subgeneric placement and a key to the otiosus species-group in North America. Zootaxa 380:1–9.

MacRae, T. C., and G. H. Nelson. 2003. Distributional and biological notes on Buprestidae (Coleoptera) in North and Central America and the West Indies, with validation of one species. The Coleopterists Bulletin 57(1):57–70.

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

Nelson, G. H. and T. C. MacRae. 1990. Additional notes on the biology and distribution of Buprestidae (Coleoptera) in North America, III. The Coleopterists Bulletin 44(3):349–354.

Yatskievych, G. 2006. Steyermark’s Flora of Missouri, Volume 2. The Missouri Botanical Garden Press, St. Louis, 1181 pp.

Copyright © Ted C. MacRae 2009

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The Loess Hills in Missouri

The term Mountains in Miniature is the most expressive one to describe these bluffs. They have all the irregularity in shape, and in valleys that mountains have, they have no rocks and rarely timber. – Thaddeus Culbertson, missionary, 1852

One of the things I enjoy most about the natural history of Missouri is its diversity. Lying in the middle of the North American continent, it is here where the eastern deciduous forest yields to the western grasslands. Coinciding with this transition between two great biomes is a complex intersection of landforms – the northern plains, recently scoured by glaciers; the southeastern lowlands, where the great Mississippi River embayment reaches its northern extent; the Ozark Highlands, whose craggy old rocks comprise the only major landform elevation between the Appalachian and Rocky Mountains; and the eastern realm of the vast Great Plains. This nexus of east and west, of north and south, of lowlands and highlands, has given rise to a rich diversity of natural communities – 85 in all according to Paul Nelson (2005, Terrestrial Natural Communities of Missouri). Despite the overwhelming changes wrought upon Missouri’s landscape during the past 200 years, passable examples of most of these communities still exist in many parts of the state and provide a glimpse of Missouri’s rich natural heritage.

Last month I talked about the critically imperiled sand prairie community in extreme southeast Missouri. This month, we travel 500 miles to the distant northwestern corner of the state to visit another critically imperiled community – the dry loess prairie. These communities are confined to thin slivers of bluff top along the Missouri River in Atchison and Holt Counties. The bluffs on which they lie are themselves part of a unique landform called the Loess Hills. Like the sand prairies of the southeastern lowlands, this angular landscape owes its birth to the glacial advances of the Pleistocene epoch (2.5 million to 10,000 years ago), when streams of meltwater – swollen and heavily laden with finely ground sediments (i.e., glacial “flour”) – filled river valleys throughout the Midwest during Pleistocene summers. Brutal cold during winter reduced these flows to a trickle, allowing the prevailing westerly winds to pick up the sediments, left high and dry, and drop them on leeward upland surfaces across Iowa and northern Missouri. The thickest deposits occurred along the abrupt eastern border of the Missouri River valley – at least 60 feet deep, and in places up to 200 feet. Loess (pronounced “luss”) is a homogeneous, fine-grained, quartz silt – undisturbed it is highly cohesive and able to stand in near vertical bluffs. It is also extremely prone to erosion, and as a result for 10,000 years now the forces of water have reshaped the Loess Hills into the landform we see today. Loess itself is not rare – thick deposits can be found in many parts of the world and over thousands of square miles across the Midwest. It is here, however, along the western edge of Iowa and northern Missouri – and nowhere else in North America – where loess deposits are deep enough and extensive enough to obliterate any influence by the underlying bedrock and dictate the form of the landscape.

It is this form that makes the Loess Hills so unique. The depth of the soil, its cohesiveness, its natural tendency to slump on steep slopes and sheer in vertical planes, and the action of water over the past several millenia have created a landscape of narrow undulating ridges flanked by steep slopes and numerous side spurs, intricate drainages with sharply cut gullies, and long, narrow terraces called “catsteps” cutting across the steep upper hillsides. It’s a sharp, angular, corrugated landscape, stretching 200 miles north and south in a narrow band of varying width from north of Souix City, Iowa, to its southern terminus in northwestern Missouri. Its western boundary is sharply delimited by the Missouri River valley, where lateral erosion (now halted by channelization of the river) and vertical sheering have created precipitous bluff faces. The eastern boundary is harder to delimit and is dependent upon the thickness of the loess. Deposits that fall below 60 feet in depth are unable to mask and reshape the rolling terrain of the eroded glacial till lying beneath. In general, this happens at distances of only 3 to 10 miles from the western edge of the landform.

Its southern terminus in Missouri, however, is the most arbitrary boundary. Discontinuous patches of deep loess terrain do occur as far south as Kansas City, but the dry hilltop prairies, common in the north, are gradually replaced by woodland in the south and disappear completely just north of St. Joseph. It is this interdigitation of two great biomes – the great deciduous forest to the east, and the expansive grasslands stretching far to the west – that give the Loess Hills such a fascinating natural history. This is due as much to the physical character of the Loess Hills themselves as to their ecotonal position at the center of the continent. Rapid drainage of rainwater off the steep slopes combines with direct sun and prevailing southwesterly summer winds to create very dry conditions on hilltops and south and west facing slopes, especially on the steeper slopes along the landform’s western edge. Such xeric conditions favor the growth of more drought-tolerant species derived from the western grasslands. North and east facing slopes and valley floors, protected from direct sun and drying winds, are able to retain more moisture, favoring the growth of woody plant species more common in the eastern forests. Seasonal moisture also shows a north-south gradient, with southern latitudes receiving higher annual rainfall totals that also favors the growth of woody plants, while the lower rainfall totals further north result in larger, more expansive grassland habitats. The steep slopes and rapid drainage create much more xeric conditions than those found further south in the flat to rolling terrain of the unglaciated Osage Plain, resulting in a more drought-tolerant mixed-grass prairie rather than the tallgrass prairie of western and southwestern Missouri. The distribution patterns of prairie versus woodland are dynamic and ever-changing, influenced by both natural and anthropogenic processes. Climatic conditions over much of the Loess Hills are capable of supporting either community type, both of which repeatedly expand and shrink as the balance tips in favor of one versus the other. In the past, the major influence was shifting periods of greater or lesser rainfall. During drier periods, grasslands expanded and woodlands shrank, finding refuge in only the moistest streamside habitats. Wetter periods allowed woody plants to migrate out of the valleys and up the slopes, especially those facing north and east. One particular very dry “hypsithermal” began about 9,000 years ago and lasted for several thousand years. Tallgrass prairies expanded as far east as present day Ohio, and todays tallgrass praires in the eastern Great Plains were invaded by even more drought-tolerant species from the shortgrass prairies further west. Eventually the hypsithermal abated, moisture levels increased, and the grasslands retreated in the face of the advanding forest. Not all of the drought-tolerant species were driven back, however, and scattered populations of these “hypsithermal relicts” still remain on locally dry sites far to the east of their normal range of distribution. Conspicuous examples of such in Missouri’s Loess Hills are soapweed yucca (Yucca glauca var. glauca) and the leafless-appearing skeletonweed (Lygodesmia juncea) (plant above, flower right). Both of these plants are normally found further west in the mixed grass prairies of the western Great Plains but are considered endangered in Missouri due to the great rarity of the dry loess prairies on which their survival depends. (Incidentally, note the crab spider legs extending from behind the petals of the skeletonweed flower). In total, more than a dozen plant species occurring in Missouri’s dry loess prairies are listed as species of conservation concern, along with one reptile (Great Plains skink) and one mammal (Plains pocket mouse).

As is typical, the insect fauna of the Loess Hills has been far less studied than its plants, but many of the species that have been documented in its prairies also show affinity to the Great Plains fauna. Both soapweed and skeletonweed have insect associates that rely exclusively on these hosts for reproduction, and as a result they are also highly restricted in Missouri. Evidence of one of these – a tiny cynipid wasp (Anistrophus pisum) that forms small spherical galls on the stems of skeletonweed – can be seen in the photo above. However, my purpose for visiting the Loess Hills this summer was to look for the rare and possibly endangered tiger beetle, Cicindela celeripes (see this post). Cicindela celeripes has not yet been recorded from Missouri but is known to occur in the Loess Hills of southwestern Iowa, and while I have not succeeded in finding it (yet!) I did observe several adults of this unusual May beetle species, Phyllophaga lanceolata. This May beetle occurs throughout the Great Plains in shortgrass prairie communities. Larvae feed in the soil on roots of grasses and other plants, while adults feed above ground on flowers and foliage. The heavy-bodied adults are unusual in the genus due to their conspicuous covering of scales (most species of Phyllophaga are glabrous or with sparsely scattered and indistinct setae) and by being active during the day. They are also relatively poorer fliers and are thus usually observed moving about on foot – as seen with this individual who was found on bare soil below a vertical cut. This snakeweed grasshopper (Hesperotettix viridis, ID by Eric R. Eaton) is another species more typically seen in the western United States, although populations have been found from across the continent. Preferred host plants include a variety of asteraceous shrubs, but as suggested by the common name snakeweeds (Xanthocephalum spp.) are highly preferred and account for its greater abundance in the west. Populations in northern and eastern portions of its range, which would include northern Missouri, are considered subspecies pratensis, while the more southern and western populations are considered the nominotypical subspecies. Interestingly (and unlike many grasshoppers), this species is considered beneficial by ranchers, since the plants on which it prefers to feed are either poisonous to livestock or offer little nutritional value while competing with more desirable forage plants for soil moisture. While exploring the upper slopes, I encountered sporadic plants of two of Missouri’s more interesting species of milkweed – whorled milkweed (Asclepias verticillata) and green milkweed (Asclepias viridiflora), raising my hopes that I might encounter one of the many Great Plains species of milkweed beetles (genus Tetraopes). However, the only species I observed was the common milkweed beetle, Tetraopes tetrophthalmus, which occurs broadly across eastern North America on the equally broadly distributed common milkweed (Asclepias syriaca).

It is a familiar refrain, but Missouri’s dry loess hill prairie communities are critically endangered. Historically, these communities were probably never as well developed as those further north, and only a few small remnants remain today due to significant woody encroachment following decades of fire suppression. Much of this encroachment has occurred in the past 50 years – Heinman (Woody Plant Invasion of the Loess Hill Bluff Prairies. M. A. Thesis, University of Nebraska at Omaha, 1982) used aerial photographs to show a 66 percent encroachment of shrubs and trees into the loess hill mixed-grass prairies between 1940 and 1981. Additional threats include overgrazing, erosion, invasion by exotic plant species and homesite development. Fewer than 50 acres of native dry loess hill prairie remain in Missouri – only half of which are now in conservation ownership. The majority of these can be found at Star School Hill Praire and Brickyard Hill Conservation Areas in Atchison County and at McCormack Conservation Area just to the south in Holt County. Controlled burning and selective cutting are being used at these sites to control woody plant invasions, but even these management techniques present challenges. Spring burns have been shown to promote the growth of big bluestem (Andropogon gerardii), which could allow it to encroach drier areas where mid-grasses such as little bluestem (Schizachyrium scoparium) and sideoats grama (Bouteloua curtipendula) typically dominate (Rushin 2005). Increases in tall grasses could shade out and eliminate some of the rarer low-growing forbs such as downy painted cup (Castilleja sessiliflora), locoweed (Oxytropis lambertii) and low milkvetch (Astragalus lotiflorus). Fall or winter burns may be more beneficial to forbs because the plants are allowed to complete flowering and seed set, but the steep slopes on which these communities occur make erosion a potential concern. Clearly, all factors must be considered when designing management plans for this rare and significant slice of Missouri’s natural heritage.

In addition to the links and references provided above, I highly recommend Fragile Giants: A Natural History of the Loess Hills, by Cornelia F. Mutel (1989). All of the above photographs were taken at Star School Hill Prairie Conservation Area on July 12, 2008. Additional photographs of Loess Hill habitats in extreme southwestern Iowa appeared in my earlier post, The hunt for Cicindela celeripes. The plants shown in photographs 5-7 are purple praire clover (Dalea purpurea), white prairie clover (D. candida), and lead plant (Amorpha canescens), respectively. Lastly, I would like to apologize for the length of this post – a consequence of my inability to temper my utter fascination with the natural world and desire to understand the depths its connectedness.