Botanizing at Prairie Fork Conservation Area

Restored claypan prairie around Crow Pond.

Today’s outing for the Webster Groves Nature Study Society (WGNSS) Botany Group was Prairie Fork Conservation Area, a newish Missouri Department of Conservation (MDC) property that is open to the public only by appointment. The 911-acre property lies within the historic 9-mile long prairie of Callaway County, Missouri—namesake for Nine Mile Prairie Township. Prior to MDC ownership, the property was owned by Ted Jones, son of the founder of Edward Jones Financial Company, and his wife Pat, who began dedicated conservation practices beginning in the 1950s. Obviously, conservation in those days—with its reliance on plantings of many now-invasive exotics such as Lespedeza sericea (sericea lespedeza), Lonicera mackii (bush honeysuckle), and Elaeagnus angustifolia (Russion olive), meant something very different than it does today. Nevertheless, the foresight and generosity of Ted and Pat Jones ultimately led to the creation a conservation area where more than half of its area has been or is being restored to claypan prairie resembling as much as possible its presettlement character.

Crow Pond.

The group engaged in two short hikes—the first through several garden plantings and then at the interface between tallgrass prairie and upland forest surrounding the perimeter of Crow Pond. Almost immediately we were treated to the sight of Gentiana andrewsii (closed gentian) in bloom. To the uninitiated, the mature flowers appear to be still-unopened buds. In fact, the corolla in this species remains closed, and the flowers are pollinated by bumblebees, which must force their way through the closed corolla. We would see numerous plants as we traveled around the pond, with most being difficult to access and photograph due to the thick, surrounding vegetation.

Gentiana andrewsii (closed gentian) mature inflorescences.
Gentiana andrewsii (closed gentian) mature inflorescences.

At pond’s edge, we found several examples of Symphyotrichum laterifolium (calico aster), one of the few white fall asters that can be identified fairly easily in the field due to its profusion of smaller-than-average-sized flowers along lateral branches with relatively few rays (9–16) and disc florets turning from pale yellow to purple, lance-elliptic leaves with a “Mohawk” (i.e., hairs only along the midvein) on the underside, and stems covered in soft, white hairs.

Symphyotrichum laterifolium (calico aster).

At the far end of the pond, the group noticed Taxodium distichum (bald cypress) trees (native to southeastern Missouri but not this far north) with strange-looking galls on the twigs. There was some discussion about whether they were caused by an insect or a fungus—their rusty-brown color and numerous spikes brought to mind the galls on Juniperus virginiana (eastern red-cedar) caused by the fungus Gymnosporangium juniperi-virginianae (cedar-apple rust). The fact that both Taxodium and Juniperus are in the family Cupressaceae made the idea of a rust fungus being the culprit seem even more likely. Nevertheless, a little online sleuthing (thank goodness for smart phones!) revealed that the actual culprit was, indeed, and insect—specifically Taxodiomyia cupressiananassa (cypress twig gall midge), a tiny fly. The spongy galls are snow-white at first and then turn brown with age, eventually dropping from the tree as leaves are shed. Maggots pupate inside the galls and adults may emerge from galls that are still on the tree later in the season. Maggots usually overwinter inside fallen galls to pupate and emerge as adults the following spring.

Galls caused by Taxodiomyia cupressiananassa (cypress twig gall midge) on Taxodium distichum (bald cypress).
Galls start out snow-white but turn brown with age.

As we walked, some of the more adventurous among us partook in freshly fallen fruits of Diospyros virginiana (persimmon). Those that were plucked from trees just as they were ready to drop were found to be the tastiest and least stringent, and sucking on the five or so seeds per fruit stretched out the flavor as long as possible. As we enjoyed the impromptu snack, a few still-flowering plants of Bidens aristosa (bearded beggarsticks or tickseed sunflower) were seen near the pond margin.

Bidens aristosa (bearded beggarsticks or tickseed sunflower).
Bidens aristosa (bearded beggarsticks or tickseed sunflower).

Eventually, we spotted from afar a large patch of bright, yellow-flowered composites in the distance. Their brown-purple discs brought the genus Rudbeckia immediately to mind, and our initial thought was R. subtomentosa (sweet coneflower) after seeing some of the large leaves with lobes. Still, the time of bloom seemed very late, and the plants were not as tall nor the leaves as “gray” as would be expected for that species. In reality, the majority of leaves were not lobed, and we decided they instead represented a form of R. fulgida (orange coneflower)—a highly variable and taxonomically difficult complex of populations.

Rudbeckia fulgida (orange coneflower).
Rudbeckia fulgida (orange coneflower).
Rudbeckia fulgida (orange coneflower) involucre.
Rudbeckia fulgida (orange coneflower) leaf.
Rudbeckia fulgida (orange coneflower) stem.

After completing the circuit around Crow Pond, the group caravanned to the northernmost and most recently renovated quarter on the property. Most of the plants established in this large tract of now-tallgrass prairie were well on their way to fall seed (and, in fact, had been combine-harvested for such), but mowing along the gravel access road allowed some still-green regrow the—the most dramatic being Helianthus maximiliani (Maximilian sunflower). This uncommon sunflower is most similar to H. grosseserratus (sawtooth sunflower) due to its large attractive flowers and narrow, curved leaves. However, leaves of the former are grayish due to the presence of many tiny hairs covering the surface of the leaf (in the latter leaves are glabrous), and the stem is rough—also due to the presence of hairs (in the latter the stem is glabrous and has a glaucus coating that can be rubbed off).

Helianthus maximiliani (Maximilian sunflower).
Helianthus maximiliani (Maximilian sunflower) involucre.
Helianthus maximiliani (Maximilian sunflower) stem/leaves.
Helianthus maximiliani (Maximilian sunflower) leaf.

The group agreed that three weeks earlier would have been a good time to see the area, and next spring or summer would be even better!

©️ Ted C. MacRae 2021

Promiscuous Plants

Naturalists have long been aware of the greater tendency for plants than for animals to create viable interspecies hybrids. This is attributable not only (as some might expect) to a higher likelihood of passive plants whose mating is mediated by pollen-hungry insects, or the wind, to hybridize more often, but rather to a greater ability of plants, with the simpler design of their anatomies, successfully to build a functioning organism with a Gemisch of genes from parents of different species. Such hybrids occur naturally, and are often reported in regional floras. Further, the advent of modern techniques for characterizing DNA has revealed that hybridizations of yore have given rise to numerous species, and higher lineages, in plants, in fungi, and to a lesser extent in animals.

My recent wanderings in quest of fall flora photos at Shaw Nature Reserve really brought this phenomenon of admixture of species to mind as I was examining populations of the three Gentiana species that live at the reserve. All three are fairly recent introductions at SNR, added to the flora in several locations in our prairie and wetland habitat reconstruction program. Hybridization among these gentian populations was first brought home in my observation over the last three years of increasing numbers of purplish and bluish and outright blue individuals in a population that was originally pure white gentian – Gentiana alba. This population was sowed in the mid-1990s as part of a mesic prairie reconstruction in the watershed of our wetland complex.

Gentiana alba, G. andrewsii and their lavender tinted hybrid growing side by side at Shaw Nature Reserve.

Pale or white bottle gentian, in "pure" form.

A few years later, 50 or so meters distant, separated by a dense row of trees and shrubs, and in a much wetter habitat in which water pools after every rain and seeps subsurficially much of the year, blue bottle gentian – G. andrewsii — was sowed into a wet prairie / sedge meadow reconstruction.

The rich blue flowers of the blue bottle gentian, Gentiana andrewsii

At first the two populations grew independently and remained separate, but what I surmise was a combination of water borne seed transport (along the shore of a pond whose edge both populations are near), and bumblebee borne pollen transport, conspired to bring gametes of the two species together, creating what population geneticists call a hybrid swarm, and what taxonomists call a — well, I can’t write it in polite company such as my readers.

Observe in the sequence of images above how a bumblebee gyne (a potential queen of one of next year’s annual bumblebee colonies) pries open a bottle gentian flower and dives in for a long drink of nectar at the base of the large vessel. Apparently the nectar is copious, because bumblebees may remain in a single gentian flower for up to a minute.

The result of pollen transport among pale and blue bottle gentians, a hybrid of intermediate characteristics.

While there are other populations of both species on the reserve (one hopes, out of bumblebee range from each other) that may retain their genetic integrity, the rampantness of the admixture at this site does give me pause.

And it gets worse! — On drier ground up the slope, among a dense planting dominated by prairie dropseed and little bluestem grasses,  a third gentian known as downy or prairie gentian – Gentiana puberulenta – was established from a seed mix sowed 10 years ago to convert the watershed of the reserve’s wetlands to prairie vegetation.

Unlike the two previously mentioned species and their hybrids, the downy gentian's petals open wide at anthesis, admitting entry to small bees and even to spindly-legged potential pollinators such as syrphid flies.

And now those perverse bumblebees have gone and defied the laws of speciesness and created what appear to be hybrids of this third gentian species with the other two. Honestly, I don’t know whether to feel that I have done some sort of wrong by creating the situation that allowed this to happen … or simply to be intrigued by this unforeseen outcome of my work, and to wonder what will come of it after I’m gone?

The gentian in the upper photo appears to be the offspring of a cross between white and downy gentian parents, while the one in the lower photo appears to be the result of a cross between blue bottle and downy gentian.

The joys of ecological restoration

Indian paintbrush and lousewort now dominate patches of SNR

I moved to Missouri in the summer of 1988, having experienced 8 years of generous support of my family’s livelihood by my research on the infamous imported fire ants of the US Southeast, and their relatives in South America. When I arrived in the Midwest, I  hoped to land a job as an insect taxonomist in a university or museum, a goal of mine since before entering college. But this dream was one that even before moving to Missouri was dimming, and then receded ever further from the realm of possibility for me (and for traditionally trained taxonomists, generally), once here. So, I began to re-think what I might do with my work life. It would be something, I hoped, that would make some use of all the course work (mostly in entomology and botany) and research (on ant systematics) I had done during my 24 years (!) of getting educated and four additional years as a post-doc. As or more important, whatever job I ended up in would somehow have to allow me to share my life-long love of nature with others.

A museum drawer of ant specimens mounted for taxonomic study, the ants no doubt frustrated by the years of inattention they have received as I have tended to the duties of my day job.

Early in my residence in eastern Missouri, I made the acquaintance of the naturalist at a 2500-acre (1000-hectare) nature reserve outside of St. Louis. Shaw Arboretum, as it was then known, is country cousin to the world-renowned Missouri Botanical Garden, and is named after the Garden’s founder Henry Shaw. Long story short, in the summer of 1990 the naturalist mentioned to me that he would soon retire, the position would become available, and that I ought to apply. So I applied, and was hired as the arboretum’s naturalist in January 1991.

A dolomite glade plant endemic to a few counties in eastern Missouri, this leatherflower was established at SNR in the 1930s, but expanded exponentially after prescribed fire was introduced in the 1990s. Here, an ant characteristic of glades and dry prairies forages on the flower.

When I came on board, the “Arboretum” had mostly ceased to be an arboretum (a formal collection of trees for display, breeding and research), and most folks seemed unable to either pronounce or define the word. Indeed we learned, through a public survey, that the strange name and the stone wall in the front actually dissuaded people unfamiliar with it from entering! Yes, there were a few patches of exotic trees scattered around the property, especially in the conifer collection near the front entrance know as the “Pinetum”, but ever since the Garden had decided around 1930 that it would not, afterall move all of its horticultural operations to this then very rural site (the original intent of its purchase), formal arboretum and botanical garden type activities had been few and far between, and the site began gradually reverting from abandoned farmland to a wilder sort of place, as well as a haven for native biota. Thus, on its 75th anniversary in the year 2000, Shaw Arboretum was renamed Shaw Nature Reserve.

Colony-founding queen bumblebees are the primary actors in loosening pollen with ultrasound from shootingstar anthers, and distributing it about the plant population.

Around that time, my title changed too, to “Restoration Biologist”. The job is multifaceted; presenting public programs and classes on various aspects of the site’s natural history, writing and reviewing articles, acting as liaison to the vigorous regional group of academic ecologists who use the site for research and teaching, a very intermittent personal research program on ants resulting in sporadic publications, and last but certainly not least, ecological restoration.

Ecological restoration, in the broad sense, consists of  two primary practices:

  • Restoration of a natural community to structure and species composition presumed characteristic of an  ;;earlier condition (however arbitrary or ill-defined).
  • Reconstruction of regional, native-like habitats, de novo, using locally acquired native plant propagules in the appropriate settings of soil, hydrology,  slope aspect and climate.

Both  require essentially perpetual, follow-up maintenance, including invasive species control, mowing, haying, grazing, selective timber removal, species richness enhancements, and prescribed burning. All of these have many variations and nuances in application, and there can be impassioned arguments about their implementation in the literature, at conferences, and in forums and blogs on ecological restoration, native plants, butterflies, beetles, etc..

An ecologically conservative lily ally of undisturbed moist soil habitats now thrives in prairie plantings at the Reserve.

Attitudes about ecological restoration vary, among practitioners, among sociologists and philosophers, and in the general public. One broad attitudinal schism lies along the lines of  whether ecological restoration activities are some sort of primitivist, grand-scale gardening, or do they represent ecologically valid landscape conservation? Another question some pose is to what extent we should interfere with “natural successsion”? Be this as it may be, most people with functioning sensory perception agree the results can be very beautiful. The loveliness of the mosaic of colors in the herb layer of a spring woodland is inarguable, especially so after it has had its woody stem density reduced, and had the leaf litter burned off, to allow more light, rain and seeds to the soil surface — even where there is genuine concern about damage to invertebrate assemblages residing in forest duff. A waving meadow of grasses and flowers in a tallgrass prairie planting, intended to replace just a few of the tens of millions of acres of this ecosystem that have succumbed to the plow, has its own grand beauty, though its per-square-meter species density of plant species remains less than half that of a native prairie remnant and it is dominated mainly by habitat-generalist insect species rather than prairie specialists, even after 30 or more years.

A self-introduced grassland ant forages among a thriving, human-introduced population of this wet prairie gentian.

The smaller, daily rewards of restoration, to the practicing ecological restorationist and to those who visit his or her work, are many. Over 20 years, in the opened-up woods, restored glades and prairie and wetland plantings at SNR, I repeatedly have enjoyed the “sudden appearance” and increase in populations of ant species (of course) that I never observed during my early years of working at SNR (then scouring it for purposes of preparing an annotated ant list). The feeling I get upon discovering that a grouping of shooting star, royal catchfly, bunch flower or bottle gentian plants, are in bloom at a site where I spread their seeds five, seven, or even ten years earlier is a bit like that one feels when a child is born. The spontaneous colonization of SNR grassland plantings by prairie ragged orchid never fails to amaze me. Bird, or frog, or katydid and cricket songs in a former crop field or pasture, as the “restored” vegetation fills in and matures, is as pleasing to my ear as it is to my soul.

A few days ago (in early July), the director of the Reserve came to my office asking if I had noticed a purply pink, “possibly orchid” flower growing on a section of a berm (planted with native vegetation) in our 32-acre wetland complex. I had not been in the area recently, but headed right out to see what it was. Joyously, and not a little surprised, I learned that seeds of the purple fringeless orchid, sowed at a location nearby 17 years previously, had washed to this site, taken root, and as terrestrial orchids are wont to do, flowered after so many years!

The black-legged greater meadow katydid thrives in low areas and near bodies of water in SNR

The prairie ragged orchid began to appear in old fields and prairie plantings where prescribed burning occurs at SNR. It has not been seen in fields maintained exclusively by mowing or haying.

The purple fringeless orchid surprised the restorationist and St. Louis area botanists by flowering in the SNR wetland area 17 years after the original sowing.

Copyright © James Trager 2010

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Friday flower – Sabatia angularis

Photo details: Canon 100mm macro lens on Canon EOS 50D, ISO 100, 1/60 sec, f/22, MT-24EX flash 1/4 power w/ diffuser caps.

Photo details: Canon 100mm macro lens on Canon EOS 50D, ISO 100, 1/60 sec, f/22, MT-24EX flash 1/4 power w/ diffuser caps.

During my explorations of the glades in the White River Hills in southwestern Missouri this past July, I noticed large populations of a flower that I couldn’t recall having ever seen before.  Vivid, striking pink petals with contrasting yellow anthers and a curiously recurved style, it seemed difficult to believe that I had simply overlooked it during my many previous visits to the area over the past 25 years.  Perhaps it was the time of year – I’ve generally avoided these glades during the month of July – normally hot, dry, and baked to a crisp.  This year and the last, however, have been different, with timely rains resulting in unusually lush July vegetation.  I also had no clue as to the identity of the plant – the square stems and opposite branching suggested a mint of some kind, but the flowers were definitely not “minty.”  I would have to simply take photographs and hope that I captured enough key characters to allow its identification once I returned home.

As it turns out, I was able to easily identify the plant as Sabatia angularis¹ (rose pink, rose gentian) using the late Dan Tenaglia’s excellent Missouri Plants website, and I wasn’t the only person to notice an apparent population explosion of this beautiful species across the Missouri Ozarks (see Justin Thomas’ excellent essay, A Sabatia Induced Rant).  As suggested by the common name, this species is in the family Gentianaceae, but it doesn’t resemble other gentians in general appearance, especially the iconic Gentianopsis crinita (greater fringed gentian) and, closer to home, Gentiana puberulenta (downy gentian), that usually come to mind upon mention of this plant family.

¹ Sabatia, for Liberato Sabbati, an 18th Century Italian botanist; angularis, Latin for angular, referring to the angled stem.

This plant occurs in the eastern U.S. west to Wisconsin in the north and Texas in the south.  Denison (1978) and Kurz (1999) both mention a preference by this species for acid soils, usually in rocky open woods, glades, old fields, and upland ridges – habitats which occur primarily across southern Missouri.  The opposite pattern of branching distinguishes this species from the alternately branched, somewhat smaller, and much less commonly encountered S. campestris (prairie rose gentian), which is most commonly encountered in the unglaciated plains of west-central Missouri.

These plants were common throughout the many glades that I visited during my two trips to the White River Hills in July, adding a vibrant splash of color to the glades after most of the other flowering plants found in these habitats have long flowered out and contrasting beautifully against the green background of uncommonly lush July grasses.


Denison, E.  1978.  Missouri Wildflowers.  A Field Guide to Wildflowers of Missouri and Adjacent Areas, 3rd revised edition.  Missouri Department of Conservation, Jefferson City, 286 pp.

Kurz, D.  1999.  Ozark Wildflowers.  A Field Guide.  Globe Pequot Press, Guilford, Connecticutt, 262 pp.

Copyright © Ted C. MacRae 2009

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