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.

REFERENCES:

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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Friday flower – Krameria lanceolata

Photo details: Canon 100mm macro lens on Canon EOS 50D, ISO 100, 1/250 sec, f/9, MT-24EX flash 1/4 power w/ diffuser caps.

Photo details: Canon 100mm macro lens on Canon EOS 50D, ISO 100, 1/250 sec, f/9, MT-24EX flash 1/4 power w/ diffuser caps.

I encountered few insects this past June on the dry slopes of sand shinnery oak shrubland that just makes it into the northwestern corner of Oklahoma’s Four Canyon Preserve – insect population levels were still depressed from the wildfire that swept through the area in April of last year.  Plant life, however, was diverse and abundant, including this most unusual plant – Krameria lanceolata (many common names, including trailing krameria, trailing ratany [sometimes spelled “rhatany”], Texan ratany, prairie sandbur, sandspur, etc.).  A dicot in the monogeneric family Krameraceae, plants in this genus share several unusual traits, the most obvious being their distinctly orchid-like, zygomorphic flowers (i.e., capable of division into symmetrical halves by only one longitudinal plane passing through the axis).  The resemblance to orchids is strictly superficial – they are most closely related to plants in the family Zygophyllaceae.

Orchids, of course, are monocots with trimerous flowers that only appear to be five-petaled because of the three petal-like sepals and the third true petal being modified into a “lip” onto which pollinating bees land.  Krameria flowers also appear five-petaled with a lip, but in this case it is the five sepals that form the “petals,” while the five true petals are modified into a lip (three fused petals) and two lateral upright “flags” called elaiphores.  These eliaphores play a central role in Krameria‘s unusual pollination biology, whose flowers produce not nectar, but fatty oils as rewards for their visitors – female bees of the genus Centris (Anthophoridae) (Simpson and Neff 1977).  The bees collect the oils from the modified external surfaces of the eliaphores, pollinating the flower in the process, and mix the oils with pollen to feed their larvae.  Although the Krameria plants are wholly dependent upon Centris bees to effect their pollination, the relationship is not mutually exclusive – Centris bees utilize other oil-producing plants as well.

All species of Krameria examined to date are obligate semiparasites, forming haustoria on the roots of a broad range of host plants.  Of the 18 species currently known in the genus, five occur in the U.S., with K. lanceolata the most widespread (Kansas and Colorado south to Arizona, New Mexico, and Texas and east to Georgia and Florida) (Austin and Honeychurch 2004). It is distinguished from the other U.S. species by its herbaceous, prostrate form.

Update 8/10/09: Mike Arduser, my hymenopterist friend who visited Four Canyon Preserve with me, wrote the following in response to my query about collecting bees from these flowers:

Yes, collected several off Krameria at Four Canyons and at Packsaddle – all were the same species, and I’m trying to remember the name as I’m writing this (all notes and material are at home) –  it was Centris lanosa. They are best found by listening, as they have a distinctive buzz as they move from flower to flower at ground level (difficult to see there).

REFERENCES:

Austin, D. F. and P. N. Honychurch.  2004.  Florida ethnobotany. CRC Press, Boca Raton, Florida. 909 pp.

Simpson, B. B. and J. L. Neff. 1977. Krameria, free-fatty acids and oil-collecting bees. Nature 267: 150-151.

Copyright © Ted C. MacRae

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Friday flower

Photo details: Canon MP-E 65mm 1-5X macro lens on a Canon EOS 50D, ISO 100, 1/250 sec, f/16, MT-24EX flash 1/8 power through diffuser caps

Photo details: Canon MP-E 65mm 1-5X macro lens on a Canon EOS 50D, ISO 100, 1/250 sec, f/16, MT-24EX flash 1/8 power through diffuser caps

While photographing small Acmaeodera beetles on flowers of Tradescantia ohioensis at Packsaddle Wildlife Management Area, I thought I should take a photo of the flower itself.  Flowers of Tradescantia species, or spiderworts, are notable for their bright yellow anthers and filaments with numerous hairs. Each of the (usually) six stamens possesses around 70-100 hairs, which in turn are composed of a chain of about 20 large, single cells that are purple in color and contain a large, water-filled central vacuole. The cells can be seen easily with low magnification – click on the photo to see the larger version, with the individual cells that make up each hair clearly visible. I haven’t been able to ascertain the function of these hairs for the plant, but their usefulness in observing division in plant cells (the flowing cytoplasm and nucleus can be seen easily) and their sensitivity to radiation and chemical mutagens have been recognized for many years. The hairs turn pink when exposed to radiation, allowing them to be used as a sort of ‘natural’ Geiger counter.

Update: While writing this post, I sent an email to George Yatskeivych, botanist at the Missouri Botanical Garden and author of Steyermark’s Flora of Missouri (1999, 2006), asking if he knew the function of the filamental hairs.  After reading his response (below), I don’t feel quite so bad for not being able to determine the answer myself:

I don’t know that I have ever heard anyone express a particular use for the hairs on the filaments of Tradescantia species.  Sometimes, hairy filaments help to trap pollen from visiting insects in proximity to the stigma or act as nectar guides, but I do not think that anyone has determined such “uses” in Tradescantia.  There may not be a selective advantage to hairy filaments in the genus.

If any botanist happens to read this post and has some insight about this, a comment would be most appreciated.

REFERENCES:

Yatskievych, G. 1999. Steyermark’s Flora of Missouri, Volume 1. Missouri Department of Conservation, Jefferson City, 991 pp.

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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Answers to “Winter botany quiz #2”

Finally, I present to you the answers to “Winter botany quiz #2 “. The delay in providing these answers was two-fold. Firstly, I knew this would be a hard test, so I wanted to give people plenty of time to figure out the answers. Secondly, the answers were delayed an extra day due because of some debate that arose among the experts I consulted about #3 – more on that below. I thank all those who participated, and while there was no clear-cut “winner”, several honorable mentions are deserved:

  • Doug Taron, who was the first to properly deduce the South African nature of these plants.
  • James C. Trager, a myrmecologist (yet still my friend!) who correctly identified the genus of #1.
  • Everyone, for guessing that #2 was “an orchid” – although Tom @ Ohio Nature was the only one to use the formal scientific name for the family, and Doug Taron was the only one to attempt a generic identification (and came close – Oncidium and Ansellia are both assigned to the tribe Cymbidieae in the subfamily Epidendroideae).

#1.  Ornithogalum seineri (family Hyacinthaceae)
Ornithogalum is a large genus occurring mostly in the drier habitats of southern Africa and around the Mediterranean.  The genus and its relatives were formerly included in the Liliaceae (as many of the participants guessed), but the group is now given familial status as the Hyacinthaceae.  This genus contains numerous species of horticultural note.  One is (as James noted) O. umbellatum, or  “star of Bethlehem”, which in North America has escaped cultivation as a garden ornamental and gained status as an invasive weed.  Another is O. longibracteatum (syn. caudatum), a popular houseplant with the common name “pregnant onion”.  This species, native to the Cape and Natal Provinces of South Africa, is easily recognized by its bulb that “gives birth” to tiny replicas of itself just beneath a thin, transparent ‘onion’ skin (as shown in the photo at right from Trans-Pacific Nursery).  At flowering, a long spike grows from the center of the green strap leaves, eventually giving rise to a spearhead of tiny white flowers situated at the end.

While I couldn’t find much information about O. seineri, I did find this spectacular photo of numerous blooming plants in bushveld habitat amongst grazing zebra (photo by ingrid1968 in this post at SANParks.org Forum).  My view of this species was not quite so spectacular, as I saw only the lone plant in the photographs posted earlier.

#2.  Ansellia africana (family Orchidaceae)
Ansellia is an African genus of orchid commonly called Leopard Orchid or African Ansellia.  There is some degree of morphological, geographical and ecological variation in Ansellia populations, with the result that several species, subspecies and varieties have been described.  Flower color varies from pure yellow to variably splotched with brown to almost completely black with finely indicated yellow divisions.  Recent taxonomic work has concluded that there are no discontinuities within the spectra of variations exhibited and the populations are thus attributable to the single, polytopic species, A. africana (Khayota 1999).

Ansellia africana is a large, perennial, epiphytic species that usually grows attached to the branches of tall trees but is sometimes found growing on rocks.  This genus is immediately recognizable by its large, cane-like pseudobulbs that arise from a basal rhizome and is notable for the white, needle-like, upward pointing aerial roots that form a sort of “trash basket” around the clump.  The term is surprisingly appropriate, since the root basket seems to function in catching dropping leaves, flowers and detritus which provide nutrients for the plant as they decay.  This species can grow to enormous size and often forms spectacular clumps, some of which have an estimated weight of more than one ton.

Of the three plants featured in the quiz, this was the one I expected someone would guess, since the species is popularly cultivated by orchid enthusiasts.  Unfortunately, the pressures of wild collection for commercial purposes has caused declines in its population.  The problem is exacerbated by the unsustainable methods use to harvest, transport, and cultivate wild-born plants.  Host trees are usually cut down and sections with the orchid removed, resulting in wholesale destruction of both orchids and hosts. After harvesting, plants are cut up and transported slowly in open handcarts, to be sold along roadsides where they may sit exposed to full sun for days or weeks.  Cutting the clumps damages the roots, and exposure results in dessication, making it difficult for harvested plants to recover once in cultivation.  Plants that do survive harvest and transplant suffer high mortality rates in cultivation due to improper attention to light and moisture regimes.

#3. Adenia sp., poss. glauca (family Passifloraceae)
To be completely honest, not only did I not expect anyone to guess this one, I didn’t think I was even going to be able to provide an answer. I sent the photos to my friend and colleague, George Yatskievych, director of the Flora of Missouri Project (and author of the recently published Steyermark’s Flora of Missouri, 1999 and 2006), who forwarded the photographs to several more colleagues, and at the same time I posted the photos on SANParks.org Forum (a fantastic resource, which I just recently discovered myself, for those interested in South Africa National Parks and their natural history). It took some time for these sources to weigh in with their opinion, which in the end were in agreement that it represented a species of African passion flower in the genus Adenia of the family Passifloraceae (not to be confused with Adenium, a genus of flowering plants in the family Apocynaceae – also occurring in Africa). As for which species, the choices had been narrowed down to either A. glauca or A. fruticosa. According to Imberbe, a photo of the leaves would have been diagnostic, and the flowers are also different (A. glauca has yellow flowers while those of A. fruticosa are green). Fred Dortort, in an article on the University of California at Berkely Botanical Garden website titled, “Passion and Poison“, notes that A. fruticosa has a tall, spindle-shaped caudex topped with a few thin, sparsely-leafed, arching branches, while in A. glauca the caudex is roughly globose and can become quite large. This description seems to favor A. glauca, which Imberbe also noted was known to occur in the area where I took the photographs.

Species identification aside, the genus Adenia is notable for its bizarre adaptations for water storage. Most of the 100 or so species in this Afrotropical and Indomalaysian genus have underground tubers. Those of species adapted to drier environments have grown proportionately larger, with some turning into above ground caudices that can take several different forms and that, in some species, may reach up to eight feet in diameter and height. Even more notable than these succulent adaptations are the poisonous properties that many plants in the genus possess. Not all species have been analyzed (and I found little or conflicting information about A. glauca and A. fruticosa), but one species in the genus – A. digitata – has gained notoriety as perhaps the most poisonous plant in the world. Two different toxins are found within its tuber, one a cyanogenic glycoside, the other a particularly potent toxin called modeccin. The latter is a 57kD protein that resembles ricin and acts a powerful inhibitor of protein synthesis by binding to ribosomes (Gasperi-Campani et al. 1978). Imberbe, in her comments about the photos I posted on SANParks.org Forum, noted the following about plants in this group:

…take heed of the Afrikaans name “Bobbejaangif” (Baboon poison)… It has been used as a fish poison, as well as in suicide and murder. It causes nausea, fits and liver and kidney damage.

REFERENCES:

Gasperi-Campani, A., L. Barbieri, E. Lorenzoni, L. Montanaro, S. Sperti, E. Bonetti, & F. Stirpe. 1978. Modeccin, the toxin of Adenia digitata. Biochemistry Journal 174:491-496.

Khayota, B. N. 1999. Notes on systematics, ecology and conservation of Ansellia (Orchidaceae), pp. 423-425. In: J. Timberlake & S. Kativu (eds.), African Plants: Biodiversity, Taxonomy and Uses, Royal Botanic Gardens, Kew.

Copyright © Ted C. MacRae 2009

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Winter botany quiz #2

In the first winter botany quiz, I learned that I have some rather astute botanists amongst my readership. They were not only able to quickly identify to species every plant I had pictured but also identify their commonality, sometimes from quite afar. As a result, this one is harder.  I use the term “winter botany quiz” in the broadest possible sense – just because it’s winter here doesn’t mean it’s cold everywhere! All of the photos were taken in the same general (for now unspecified) locality during late November and early December (this paragraph simply reeks of clues!).

To give everyone a fair chance, I’ve turned on comment moderation so people can submit their answers without seeing what has already been submitted.  I’ll remove moderation after a couple days or so.  First one with all the right answers wins the admiration and jealousy of their peers!

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#1A

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#1B - closeup of flowers in #1A

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#2A

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#2B - closeup of flowers of #2A

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#3A - the vine, not the trees

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#3B - closeup of vine base

Copyright © Ted C. MacRae 2009

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Top Ten of 2008

For the first post of 2009, I begin with a look back at some of my favorite photos from 2008 (idea stolen from Alex Wild and others).  I initially hesitated to do a “best photos” post since I’m not really a photographer – just an entomologist with a camera.  Nevertheless, and with that caveat in mind, I offer ten photos that represent some of my favorites from this past year. To force some diversity in my picks, I’ve created “winning” categories (otherwise you might just see ten tiger beetles!). Click on the photos to see larger versions, and feel free to vote for your favorite. If so, what did you like about it? Was there a photo I didn’t pick that you liked better?  Enjoy!

Best tiger beetle

Cicindela formosa generosa

From “All the better to see you with, my dear!” (September 2008).  Picking a top tiger beetle photo was tough with so many to choose from.  Ultimately, I decided I really like these face-on shots, and of the several I’ve posted this one of Cicindela formosa generosa has the overall best composition, balance and symmetry.  I considered this one of Cicindela formosa formosa – with its half-cocked jaws, it probably has better personality.  However, the one above got the final nod because it is a true field shot of an unconfined, unmanipulated individual.

Best jewel beetle

Aegelia petelii

From Buppies in the bush(veld) (December 2008).  Although taken back in 1999, I just recently scanned and posted this photo of Agelia petelii from South Africa.  I like the bold, contrasting colors of the beetle combined with the soft colors of the host foliage.  Runners up included these photos of Evides pubiventris with its sumptuous iridescent green blending beautifully with the green background (but suffering slightly from shallow depth of field) and Chrysobothris femorata with its intricate surface sculpturing.

Best longhorned beetle

Tetraopes femoratus

From Rattled in the Black Hills (September 2008).  This was an easy choice – none of the other longhorned beetle photos that I posted during 2008 matched this photo of Tetraopes femoratus for clarity, composition, and the striking contrast between the red color of the beetle and the green color of the host plant.  I especially like the detailing of the body pubescence.

Best non-beetle insect

Proctacanthus milbertii

From Magnificently Monstrous Muscomorphs (November 2008).  I do like other insect besides beetles, and robber flies are hard to beat for their charisma.  This photo of Proctacanthus milbertii (which, as Chris Taylor pointed out, literally translates to “Milbert’s spiny butt”), has great composition and nice, complimentary colors.  I like contrast between the fine detail of the fly and the soft background.

Best non-insect arthropod

Argiope aurantia

From Happy Halloween! (October 2008). I didn’t have many non-insect arthropod photos to choose from, but this photo of a female Argiope aurantia (yellow garden spider) would be deserving of recognition no matter how many I had to choose from. I like the bold, contrasting colors and symmetry of the spider in front of the dappled background of this photo.

Best non-arthropod animal

Prairie rattlesnake (Crotolus viridis)

Another one from Rattled in the Black Hills (September 2008).  This is admittedly not the best photo from a purely technical perspective – it’s a little out of focus, and the color is a bit off.  However, no photo could better convey the moment – confronted with a live, angry prairie rattlesnake (Crotalus viridis) (among the more aggressive species in the genus).  The forked tongue and rattle – blurred in motion – were icing on the cake.

Best wildflower

Victoria Glades

From Glades of Jefferson County (July 2008).  I had several wildflower closeups to choose from, but I kept coming back to this field shot of pale purple coneflower (Echincea simulata) and Missouri evening primrose (Oenethera macrocarpa).  The eastern redcedars (Juniperus virginiana) in the background are at once indicative of their preferred habitat (limestone/dolomite glades) and also testament to their threatening encroachment.

Best tree

Calocedrus decurrens

From the very simply and aptly named Lake Tahoe, California (March 2008).  Incense cedar (Calocedrus decurrens), with its reddish, deeply furrowed bark and great height, is one of the most majestic of western conifers.  I was captivated by this tree – beautiful even in death and contrasting nicely with the surrounding green foliage.

Best rockscape

Pipestone National Monument, Old Stone Face

From Pipestone National Monument (April 2008).  “Old Stone Face” is one of Pipestone’s most recognizable geologic features, and the short angle of the sun on this early spring day provided nice detail to the cracks and fissures of the rock – almost appropriately adding a weathered “age” to this old man.

Best landscape

Emerald Isle, Lake Tahoe

Another one from Lake Tahoe, California (March 2008).  Few places on earth are more photogenic than Lake Tahoe, and this perspective overlooking Emerald Bay is among the finest views I’ve seen.  Brilliant blue skies and majestic snow covered mountains reflected perfectly from the still surface, with Fannette Island providing a perfect focal point for the photo.

Best miscellaneous

Water drops, Ozark Trail, Trace Creek SectionFrom Ozark Trail, lower Trace Creek Section (December 2007).  While technically not a 2008 photo, it’s close enough.  This was one of the first macro photographs I took with my camera, and it remains one of my favorites.  A chance occurence of an unlikely subject, created by cold temperatures and heavy moisture-laden air. I like the contrast between the water drops – sharp, round, and clear – with the vertical shapes of the leaf petioles and background trees.  Viewing the image full-sized reveals the reflection of the photographer in the leftmost water drop.

Subsequent edit: Okay, so after I put this post together, I realized I actually featured eleven photos – too much difficulty choosing, I guess. Let’s call it a baker’s ten.

A sand prairie autumn

Splitbeard bluestem seed headsAsk any astronomer when autumn begins, and they will likely tell you it begins at the autumnal equinox – when shortening days and lengthening nights become equal as the sun crosses over the celestial equator. According to them, fall began this year on September 22 – at 11:44:18 A.M. EDT, to be precise. I agree that autumn begins at a precise moment, but it is not at the equinox. Rather, it is that unpredictable moment when a sudden crispness in the air is felt, when the sky somehow seems bluer and shadows seem sharper, and hints of yellow – ever so subtle – start to appear in the landscape. Butterfly pea blossomIn Missouri, with its middle latitudes, this usually happens a few weeks before the equinox, as August is waning into September. It is a moment that goes unnoticed by many, especially those whose lives and livelihoods have lost all connection with the natural world. To plants and animals, however, it is a clear signal – a signal to begin making preparations for the long cold months of winter that lie ahead. Plants that have not yet flowered begin to do so in earnest, while those that have shift energy reserves into developing seeds. Animals take advantage of their final opportunities to feed before enduring the scarcities of winter, digging in to sleep through them, or abandoning altogether and migrating to warmer climes. Insects begin hastily provisioning nests for their broods or laying eggs – tiny capsules of life that survive the harsh winter before hatching in spring and beginning the cycle anew.

Sand prairie in early September.Sand prairie in early October.  Note abundance of splitbeard bluestem seed heads.Across much of Missouri, in the Ozark Highlands and in riparian ribbons dissecting the northern Plains, autumn brings an increasingly intense display of reds, purples, oranges, and yellows, as the leaves of deciduous hardwoods begin breaking down their chlorophyll to unmask underlying anthocyanins and other pigments. Small southern jointweedIn Missouri’s remnant prairies, seas of verdant green morph to muted shades of amber, tawny, and beige. This subtle transformation is even more spectacular in the critically imperiled sand prairies of the Southeast Lowlands, where stands of splitbeard bluestem (Andropogon ternaries – above) turn a rich russet color while fluffy, white seed heads (1st paragraph, 1st photo) appear along the length of each stem, evoking images of shooting fireworks. Small southern jointweed (Polygonella americana – right) finds a home at the northern extent of its distribution in these prairie remnants and in similar habitats in nearby Crowley’s Ridge, blooming in profusion once the cooler nights arrive. Butterfly pea (Clitoria mariana – 1st paragraph, 2nd photo) blooms add a gorgeous splash of soft purple in contrast to the muted colors of the plants around them.

Kent Fothergill, Ted MacRae, and Rich ThomaAfter first becoming acquainted with Missouri’s sand prairies this past summer, I knew a fall trip (or two) would be in order. The extensive deep, dry sand barrens were ideal habitat for sand-loving insects, including certain spring/fall species of tiger beetles that would not be active during the summer months. The cooler nights and crisp air of early fall make insect collecting extraordinarily pleasurable, so it took little effort to convince friends and colleagues Kent and Rich to join me on another excursion to these extraordinary remnant habitats, along with my (then 8 yr-old) daughter Madison (who would likely characterize this as “tallgrass” prairie). Madison MacRae, age 9 (almost)I was, as ever, on the lookout for tiger beetles; however, temperatures were cool, skies were overcast, and the fall season was just beginning, greatly limiting tiger beetle activity during this first fall visit. We did see one Cicindela formosa (big sand tiger beetle), which cooperated fully for a nice series of photographs. We also found single specimens of the annoyingly ubiquitous C. punctulata (punctured tiger beetle) and a curiously out-of-place C. duodecimguttata (12-spotted tiger beetle), which must have flown some distance from the nearest dark, muddy streambank that it surely prefers. Of greatest interest, we found two specimens of C. scutellaris (festive tiger beetle), which in this part of Missouri is represented by a population presenting a curious mix of influences from two different subspecies (more on this in a later post…). Despite the scarcity of tiger beetles, other insects were present in great diversity, some of which I share with you here.

Ululodes macleayanusThis bizarre creature, sitting on the stem of plains snakecotton (Froelichia floridana), is actually a neuropteran insect called an owlfly (family Ascalaphidae). Looking like a cross between a dragonfly and a butterfly due to its overly large eyes and many-veined wings but with long, clubbed antennae, this individual is demonstrating the cryptic resting posture they often assume with the abdomen projecting from the perch and resembling a twig. The divided eyes identify this individual as belonging to the genus Ululodes, and Dr. John D. Oswald (Texas A&M University) has kindly identified the species as U. macleayanus. As is true of many groups of insects, their taxonomy is far from completely understood. Larvae of these basal holometabolans are predaceous, lying on the ground with their large trap-jaws held wide open and often camouflaging themselves with sand and debris while waiting for prey. The slightest contact with the jaws springs them shut, and within a few minutes the prey is paralyzed and can be sucked dry at the larva’s leisure.

Ant lion, possibly in the genus Myrmeleon.Another family of neuropteran insects closely related to owlflies are antlions (family Myrmeleontidae, sometimes misspelled “Myrmeleonidae”). This individual (resting lower down on the very same F. floridana stem) may be in the genus Myrmeleon, but my wanting expertise doesn’t allow a more conclusive identification [edit 4/12/09 – John D. Oswald has identified the species as Myrmeleon immaculatus]. Strictly speaking, the term “antlion” applies to the larval form of the members of this family, all of whom create pits in sandy soils to trap ants and other small insects, thus, it’s occurrence in the sand prairie is not surprising. Larvae lie in wait beneath the sand at the bottom of the pit, flipping sand on the hapless prey to prevent it from escaping until they can impale it with their large, sickle-shaped jaws, inject digestive enzymes that ‘pre-digest’ the prey’s tissues, and suck out the liquifying contents. Finding larvae is not easy – even when pits are located and dug up, the larvae lie motionless and are often covered with a layer of sand that makes them almost impossible to detect. I’ve tried digging up pits several times and have failed as yet to find one. Larvae are also sometimes referred to as “doodlebugs” in reference to the winding, spiralling trails that the larvae leave in the sand while searching for a good trap location – these trails look like someone has doodled in the sand.

Bembix americanaThis digger wasp, Bembix americana (ID confirmed by Matthias Buck), was common on the barren sand exposures, where they dig burrows into the loose sand. Formerly included in the family Sphecidae (containing the better-known “cicada killer”), members of this group are now placed in their own family (Crabronidae). Adult females provision their nest with flies, which they catch and sting to paralyze before dragging it down into the burrow. As is common with the social hymenoptera such as bees and paper wasps, these solitary wasps engage in active parental care by providing greater number of prey as the larva grows. As many as twenty flies might be needed for a single larva. I found the burrows of these wasps at first difficult to distinguish from those created by adults of the tiger beetles I so desired, but eventually learned to distinguish them by their rounder shape and coarser, “pile” rather than “fanned” diggings (see this post for more on this subject).

Stichopogon trifasciatusRobber flies (family Asilidae) are a favorite group of mine (or, at least, as favorite as a non-coleopteran group can be). This small species, Stichopogon trifasciatus (ID confirmed by Herschel Raney), was also common on the barren sandy surface. The specific epithet refers to the three bands of alternating light and dark bands on the abdomen. Many species in this family are broadly distributed but have fairly restrictive ecological requirements, resulting in rather localized occurrences within their distribution. Stichopogon trifasciatus occurs throughout North America and south into the Neotropics wherever barren, sandy or gravely areas near water can be found. Adults are deadly predators, swooping down on spiders, flies and other small insects and “stabbing” them with their stout beak.

Chelinidea vittigerPrickly pear cactus (Opuntia humifusa) grows abundantly in the sandy soil amongst the clumps of bluestem, and on the pads were these nymphs of Chelinidea vittiger (cactus bug, family Coreidae). This wide-ranging species occurs across the U.S. and southward to northern Mexico wherever prickly pear hosts can be found. This species can either be considered a beneficial or a pest, depending upon perspective. On the one hand, it serves as a minor component in a pest complex that prevents prickly pear from aggressively overtaking rangelands in North America; however, prickly pear is used by ranchers as emergency forage, and fruits and spineless pads are also sometimes harvested for produce. In Missouri, O. humifusa is a non-aggressive component of glades, prairies, and sand and gravel washes, making C. vittiger an interesting member of the states natural diversity.

Ammophila sp., possibly A. proceraThis wasp in the genus Ammophila (perhaps A. procera as suggested by Herschel Raney) was found clinging by its jaws to a bluestem stem in the cool morning, where it presumably spent the night. One of the true sphecid (or “thread-waist”) wasps, A. procera is a widespread and common species in eastern North America. One of the largest members of the genus, its distinctive, bold silver dashes on the thorax distinguish it from most other sympatric congeners. Similar to the habits of most other aculeate wasp groups, this species captures and paralyzes sawfly or lepidopteran caterpillars to serve as food for its developing brood. Females dig burrows and lay eggs on the paralyzed hosts with which the nests have been provisioned. Adults are also found commonly on flowers, presumably to feed on nectar and/or pollen.

Dusty hog-nosed snakeRich is a bit of herpatologist, so when he brought this hog-nosed snake to our attention we all had a good time pestering it to try to get it to turn upside down and play dead. I had never seen a hog-nosed snake before but knew of its habit of rolling over and opening its mouth with its tongue hanging out when disturbed, even flopping right back over when turned rightside up or staying limp when picked up. We succeeded in getting it to emit its foul musky smell, but much to our disappointment it never did play dead, instead using its shovel-shaped snout to dig into the sand. Dusty hog-nosed snake - head closeupWe had assumed this was the common and widespread eastern hog-nosed snake (Heterodon platirhinos); however, in our attempts to turn it over I noticed its black and orange checker patterned belly. I later learned this to be characteristic of the dusky hog-nosed snake (H. nasicus gloydi), only recently discovered in the sand prairies of southeast Missouri and regarded as critically imperiled in the state due to the near complete destruction of such habitats. Disjunct from the main population further west, its continued survival in Missouri depends upon the survival of these small sand prairie remnants in the Southeast Lowlands.

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.