In search of a lost natural community: the Ozark savanna edition

Calvin Maginel is the Ecological Resource Scientist at Shaw Nature Reserve in Gray Summit, Missouri.

Anyone hoping to join the articulate stream of Missouri articles about natural communities ought to lovingly reference Paul Nelson’s “The Terrestrial Natural Communities of Missouri” (2010). In that vein, we will start our journey with page 233, the Savanna.

Paul differentiates savannas largely by overstory, topography, and light level characteristics. Primarily, savannas are grasslands that happen to hold little pockets, family clusters, of trees, that mosey through the swaying grass like the slowest of turtles. The natural history of these clusters is as such: a mature parent hosts numerous offspring around her perimeter that shelter her from the repeated onslaughts of prairie fires, while she in turn nurtures offspring on the lee side which will eventually replace her. They are separate from woodlands in that savannas exhibit a tree canopy of less than 30%, while woodlands can range from 30% to 90% canopy. Paul further describes the ground flora layer of savannas as being highly indicative of a prairie, holding the majority of a site’s diversity, and being strongly adapted to frequent fire.

Of the six savanna communities Paul describes, as nostalgia blurs the typeset, two are considered S1 (critically imperiled) and four are SH, or state historic. A glass of cold water to the face: no known examples remain when something is classified as state historic. To put numbers on this, an estimated 6.5 million acres of savanna in Missouri are now represented by <1,000 recognized acres. Robin Wall Kimmerer aptly wrote: “If grief can be a doorway to love, then let us all weep for the world we are breaking apart so we can love it back to wholeness again.”

Stylized drawings of the prairie-forest continuum, borrowed from The Tallgrass Restoration Handbook by Packard and Mutel.

Recognizing a savanna

As nice as it is to reminisce about and romanticize processes long devastated by European colonizers, if there are (nearly) no savannas left, then why does it matter? Well, there still is hope! While Missouri has a fair percentage of public land (11.2%), most of which has received extensive visits by ecologists throughout the years, the other 88.8% of private lands in Missouri often harbor as-yet-undescribed natural communities that may classify as savanna. In an effort to heighten awareness of these potential gems in the fire-starved hills, I offer a photo tour of a private site in southwest Washington County, near the town of Courtois, that could be described as a savanna. A few points about this site: it is currently being managed for its ground flora character, with repeated fire and herbicide, specifically to the detriment of encroaching cedars and woody re-sprouts. For 25 years prior to the current ownership, it received two fires and periodic mowing to maintain its relatively shrub-free character. Prior to that, it is assumed that this was a hay meadow, cut annually for livestock that were grazed in the valley nearby but not itself grazed. There is a rusty but strong sickle-bar mower still parked in the grass that is set up for a mule to pull, with patent dates from the 1920s.

Since Paul begins with the overstory, so too will this tour. Anecdotal descriptions of certain areas in the Ozarks by foresters refer to “wolf trees”, trees with spreading branches that were removed from the woodlot since those individuals were considered to be exhausting resources around themselves, much as wolves were believed to be harmful predators that exhausted prey species. An example of this can be found in Photo 1, where a large white oak shows the breadth of branches characteristic of an “undesirable” wolf tree. As mentioned in the caption, the health of the lowest branches can tell something about a site’s history. Overgrazing by cattle or other domestic animals often defoliates these branches until the tree sheds them entirely, so an observation of a tree similar to this one might mean that this site was hayed but not grazed intensively.

Now that photos have been mentioned, we’ll begin the photo tour in earnest. All photos are from August 22nd, 2021, unless otherwise stated. To the right side of Photos 1, 2, and 3, you will notice a young shortleaf pine (Pinus echinata) with a wolfish future, and in Photos 2 and 3, there is a distinctive Eastern Red Cedar (Juniperus virginiana) that seems to have lost half its top. All other photos will contain at least a blurry version of those two distinctive trees, in an effort to maintain scale. Speaking of scale, the distance between the white oak wolf tree and the red cedar is a little over 250 feet (76 meters). Photos 2 and 3, of almost the same area at different phenologies, hold the first real hope of a savanna classification. The structure is distinctively grass- and forb-dominated. While clearly the floral display is greater during June, this is not unexpected in an intact prairie system where suitable micro-habitats are dominated by the best-adapted competitors for those micro-habitats. For example, the glade coneflower in Photo 3 is distributed between the foreground of the photo and the base of the pine tree, but seems to decrease in abundance towards the red cedar in the upper left of the photo. Presumably, soil or other characteristics make the former area highly suitable for glade coneflower, despite the fact that no bedrock or other glade indicators occur in those areas. That said, it stands to reason that glade coneflower, currently relatively restricted to glade communities, must have had a mechanism to lay claim to those communities. Possibly this species was historically as ubiquitous in Ozark savannas and prairies as it currently is in glades.

Photo 1. Forgive the valiantly bolting hickory grubs and mowed path, but this white oak (Quercus alba) exemplifies the spreading nature of a relatively open-grown specimen. Note how the lower branches actually touch the warm season grasses: despite 5 recent years of annual dormant season fire, they are not set back. In fact, one telltale of current or historical colonizer-style grazing is that these perpetually-stretching side limbs are defoliated until they succumb and die. Trees with this character can tell a lot about a site’s history.
Photo 2. With the same white oak as in Photo 1 to the left of the frame, this photo shows the vegetative structure of the site. You will notice a handful of woody re-sprout clumps, but this area is largely dominated by warm season grasses and prairie forbs.
Photo 3. June 14, 2016 is the date on this photo. Note the profusion of wild quinine (Parthenium integrifolium) and glade coneflower (Echinacea simulata), the latter of which is commonly identified by its yellow pollen. The more westerly species with white pollen, E. pallida, does not occur on this site.

In addition to the striking summer floral display in Photo 3, there are distinct waves of blooms throughout the season. Each species, present in profusion in its preferred micro-habitat and scattered elsewhere, blooms en masse and then fades into the background, letting another take the stage like a carefully-choreographed dance.

At this point, you may be noticing that the common names for many of the plants listed in the photo captions refer to a habitat (eg “glade” coneflower, “upland” white goldenrod, “prairie” coreopsis). This name-relation to a community can serve to help with identifying that community, but the overall assemblage of species tells a stronger story. When you consistently encounter species that occur within multiple habitats (Ozark woodlands, glades, and/or prairies), which is true for most of the species shown in these photos, it may be a telltale sign of the missing connection between all of those communities. Similar to the previously mentioned glade coneflower, both downy gentian and the upland white goldenrod are commonly found in glades and open woodlands. They tend to fall out in areas with >60% shade. Almost all of these species are considered highly conservative; species that we expect to maintain high fidelity to intact ecosystems. Missouri is one of the states that maintains a coefficient of conservatism list, with values ranging from 0 to 10, where 9-10s are virtually only found in the highest quality habitats. For example, the downy gentian, white upland goldenrod, savanna blazing star, and southern prairie aster are all c=9 species. Most of the grasses are 4 or 5, as well as the prairie dock, prairie blazing star, and Canada lousewort. When visiting a natural community, generally the more intact, remnant sites boast a bell curve of c-values, with the peak being a good diversity of c = 4-6 species. The distinctive composition at this site, with conservative prairie and glade species present (yet located deep in the Ozarks in an area not considered historic prairie), triggers the savanna vibe.

Photo 4. Savanna blazing star (Liatris scariosa var. nieuwlandii), wild quinine, big bluestem, and ashy sunflower. Savanna blazing star is currently listed as a species of conservation concern in Missouri.
Photo 5. Southern prairie aster (Eurybia hemispherica, old name Aster paludosus), forms a colony with leaves reminiscent of a graminoid until it blooms with striking purple discs.

An additional, striking character of this site is the height of the vegetation (Photos 6 and 7). In particular, Photo 6 includes a species called ashy sunflower (Helianthus mollis). Various botanists and restorationists have used disparaging terms for this species, even the socially problematic term “thuggish,” since this species tends to form thick 2-4 foot tall monocultures to the detriment of other species. Surprisingly, the ashy sunflower at this site is a whopping 0.5 – 1 foot high and comfortably interwoven with other species. The matrix grasses, consisting of mostly of big bluestem (Andropogon gerardii), little bluestem (Schizachyrium scoparium), prairie dropseed (Sporobolus heterolepis), and Indian grass (Sorghastrum nutans) are consistently knee-high or shorter, barring their flowering stems of around 5 feet. In many prairie reconstructions, the big bluestem and Indian grass commonly attain heights of more than 9 feet and encountering each clump of bunchgrass is like climbing up a small mima mound. Here, the grass ramets have presumably reached old age and no longer exhibit the mounding character. Many ecologists attribute the presence of hemi-parasitic species like Canada lousewort (Pedicularis canadensis), scarlet paintbrush (Castilleja coccinea), or blue hearts (c=10!, Buchnera americana) to decreased robustness of warm season grasses. All three of these hemiparasitic species are present at this site, yet the truth is that the science of ecology is still learning about what actually makes remnant sites look consistently different than reconstructed sites. Is it nutrient limitation, due to all niches being occupied in remnants? Maybe it’s mycorrhizal associations determining community composition and structure, since Arbuscular Mycorrhizal Fungi have been shown to strongly affect plant communities. What about beneficial or pathogenic bacteria, or soil structure, maybe parent material, or surely it’s the site’s aspect and moisture profiles? The obvious answer is that it’s a combination, and that we have much to learn about our natural communities. The quote by J. K. Rowling, “Understanding is the first step to acceptance, and only with acceptance can there be recovery,” might as easily have been about natural communities as it was directed at Harry Potter’s life.

Photo 6. Upland white goldenrod/prairie goldenrod (Oligoneuron album) blooms amongst two Silphium species, prairie coreopsis (Coreopsis palmata), and well-mannered ashy sunflower (Helianthus mollis) stems. Rarely is the term well-mannered used in conjunction with ashy sunflower.
Photo 7. Downy gentian (Gentiana puberulenta) looking disheveled prior to its glorious frost-triggered blooms, amidst prairie dock, prairie blazing star, and a grass/sedge matrix. A dominant sedge species here is few-flowered nut rush (Scleria pauciflora).

The last point regarding vegetative species groups are those considered woodland species. Just like in prairies and glades, there are a handful of woodland indicator species that assist with identification of the natural community we call a woodland in Missouri. As a reminder, woodlands have a canopy cover of >30%, all the way up to 90% cover, yet have an open mid-story maintained most commonly with frequent fire. Some characteristic species present at this site that are considered common woodland indicators include deerberry (Vaccinium stamineum), Samson’s snakeroot (Orbexilum pedunculatum), and stiff aster (Ionactis lineariifolia). The last species is especially striking, as botanists and plant geeks commonly observe it in acidic, poor-nutrient woodlands or power line rights-of-way. Yet keep in mind that glade coneflower, a known calciphile, is hanging out with the stiff aster. Whatever processes are allowing this site to host such a mish-mash of Ozark woodland, glade, and prairie flora, it seems to support the understudied idea that there really was a thriving prairie-forest ecotone amongst these aged hills.

Photo 8. Prairie willow in the foreground left (Salix humilis), vying for growing space with prairie dock (Silphium terebinthinaceum), tall tickseed (Coreopsis tripteris), little-leaf tick trefoil (Desmodium ciliare), and others.

Wrapping Up

As outlined above, there are few to no known savannas left in Missouri. While many agencies are trying valiantly to re-create open or closed woodlands, the sawdust of Missouri’s logging culture weighs heavily on our boots and generally there are fewer restoration practitioners aiming for savannas and their lack of timber products. The Nature Conservancy comes to mind, but the majority of their sites classify as true prairie, except maybe Bennett Spring Savanna. That site, like Ha Ha Tonka State Park, tends to maintain characteristics more similar to open woodland, but has lovely intact ground flora with a solid assemblage of prairie species. The critical missing piece is that for most natural community restorations, we have a goal in mind, dictated and informed by multiple examples of that community. With savannas and the lack of high-quality examples, we are left with a great deal more speculation. The hope mentioned in the beginning comes into play with each of you. There is a plethora of private lands that are largely inaccessible to state and federal biologists. If you get a chance to visit a friend’s farm, do so with a thought to some of the characteristics described above. Citizen science really does work, and maybe the next branch of citizen science is natural community identification! As Rachel Carson said, “The more clearly we can focus our attention on the wonders and realities of the universe about us, the less taste we shall have for destruction.”

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