Friday Flower – Dwarf Spiderwort

Living at the foothills of the Ozark Highlands, I sometimes forget how unique the biota of this ancient landscape truly is. More than 200 species of plants and animals are largely restricted to the region, with around 160 of these being true Ozark endemics found nowhere else on earth. The biodiversity of the region stems from the landform’s unusual geology, topography and hydrology, it’s ectotonal position within the North American continent, and its distinction as the only significantly elevated landform between the Appalachian and Rocky Mountains. Many Ozark endemics are found in the region’s abundant caves and sinkholes, formed by underground dissolution of its massive limestone/dolomite bedrocks. Others represent isolated populations of more typically northern plants and animals that found refuge here during the Pleistocene glacial advances. Still others evolved during periods of isolation when vast inland seas covered much of the continent’s interior.

Tradescantia longipes, known locally as dwarf spiderwort or wild crocus, is a particularly exquisite Ozark endemic found scattered in dry igneous woodlands of the Missouri’s St. Francois Mountains and Arkansas’ Ouachita Mountains. I first saw this species two years ago in May at Crane Lake in the heart of the St. Francois Mountains, and the plants shown here were seen this past April in the igneous woodlands of Sam Baker State Park at the southernmost extent of the St. Francois Mountains’ igneous exposures. The genus to which this plant belongs contains some much more widely distributed (though no less striking) members (e.g. T. ohioensis, which I featured in my first “Friday Flower” post). Tradescantia longipes flowers are similar to those of T. ohioensis, but the plant differs from most others in the genus by its short, squat habit of growth and strictly basal leaves.

One feature shared by T. longipes with all other members of the genus is the dense fringe of hairs arising from the stamen filaments.  I discussed these in my first Friday Flower post, noting that each of the 70-100 hairs per filament is composed of a chain of about 20 large, single cells – easily seen with low magnification. While their sensitivity to radiation and chemical mutagens has been recognized for many years (the hairs turn pink when exposed to radiation), less seems to be known about their natural function for the plant.  It is interesting to note, however, that the flowers of Tradescantia and related genera rely heavily on insects for pollination (primarily bees and bee flies), yet they do not produce nectar.  Faden (1992) has speculated that the stamen hairs might combine with floral scents and the nearly pollenless anthers to deceptively attract insects, provide footholds, retain pollen fall, and influence the pollen-collecting behavior of the insects.

Photo Details: Canon 50D (ISO 100, 1/250 sec, f/14-18), Canon 100mm macro lens, Canon MT-24EX flash (1/4 ratio) w/ Sto-Fen diffusers. Post-processing: levels, unsharp mask, minimal cropping.

REFERENCE:

Faden, R. B.  1992. Floral attraction and floral hairs in the Commelinaceae.  Annals of the Missouri Botanical Garden 79(1):46–52.

Copyright © Ted C. MacRae

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Friday Flower – Pawpaw

As my friend Rich and I stood in the verdent understory admiring the spectacular panicles of red buckeye punctuating the green lushness, a small brown flower on a leafless branch above me caught my eye.  “Pawpaw!” I exclaimed, perhaps partly in amazement that it took us awhile to notice the trees that were, in fact, all around us.  Pawpaw (Asimina triloba) is a member of the only temperate genus in the otherwise exclusively tropical and subtropical family Annonaceae (Custard Apple Family).  Although not nearly as restricted in occurrence in Missouri as the red buckeyes with which they were growing, they are nevertheless always a treat to see. Scattered throughout the state, they can be found growing in bottomland forests, ravines in mesic upland forests, along woodland streams, and at bases of bluffs (Yatskievych 2006).

Pawpaws are, of course, famous for their large edible fruits, sometimes called Indian bananas, Missouri bananas, Michigan bananas, [insert eastern state here] bananas, etc.  Technically, however, the pawpaw fruit is a berry, since it is derived from a single pistil and has multiple seeds embedded within the pulpy matrix.  I’ve not tried the fruit myself, not for lack of desire but rather an inability to find them when they ripen in fall before the birds and mammals get to them.  Some effort has been made to cultivate the plant for fruit production, but low fruit set seems to be a persistent problem due to reproductive self-incompatibility.

Pawpaw also famously serves as the larval food plant for the zebra swallowtail butterfly, Eurytides marcellus (family Papilionidae).  Beyond this, however, there seem to be not many insects associated with the plant.  I have collected dead wood of pawpaw in an effort to determine the species of wood-boring beetle species that are associated with it.  The only species I’ve reared is the longhorned beetle, Elaphidion mucronatum (whose common name “spined oak borer” belies the fact that it is one of the most polyphagous of all North American species), and two other longhorned beetles, Eupogonius pauper and Urgleptes querci (also highly polyphagous), have also been reported being reared from dead wood of this plant.  I have not associated any jewel beetles with pawpaw, nor have any such associations been reported in the literature.  It would appear that woodboring beetles are not fond of the soft, weak wood of pawpaw, perhaps due to the plant’s annonaceous acetogenins with known pesticidal qualities (Ratnayake et al. 1993) (acetogenins are also under investigation as anti-cancer drugs).  Other poisonous compounds, chiefly alkaloids, are found in various parts of the plant, especially the seeds and bark, and likely play a role in herbivore defense. Insect pollinators also seem to be infrequent, as I have not noted any insects on its flowers. Most members of the family are pollinated by beetles (Yatskievych 2006), but the meat-colored, downward-facing, not-so-sweet-smelling flowers of pawpaw suggest pollination by flies, perhaps those attracted to carrion.

REFERENCE:

Ratnayake, S., J.K. Rupprecht, W.M. Potter, and J.L. McLaughlin. 1993. Evaluation of the pawpaw tree, Asimina triloba (Annonaceae), as a commercial source of the pesticidal annonaceous acetogenins. p. 644-648. In: J. Janick and J.E. Simon (eds.), New Crops. Wiley, New York.

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

Copyright © Ted C. MacRae 2010

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Friday Flower – Ozark Witch Hazel

Spring is beginning its “march” across the nation, and in typical fashion the month started out with the promise of pleasant weather but is throwing a few tantrums before giving way to April. For most folks in the lower Midwest, spring began a week or so ago when daffodils began popping up from nowhere and dotting the suburban and semirural landscapes with their yellow smiles. Forsythia are also set to burst forth, their appearance temporarily put on hold by this latest cold/wet snap, but when they do most people here will be satisfied that spring has finally come. For me, spring comes much earlier, and it’s not planted ornamentals that mark its beginning, but native trees.  Silver maples (Acer saccharinum) and American elms (Ulmus americana) are first, bursting open in the very first warm days of early March.  These are followed by the sugar maples (A. saccharum) and red maples (A. rubrum) that are in full bloom now, which will themselves give way to the redbuds (Cercis canadensis) and serviceberrys (Amelanchier arborea) that will close out the month before flowering dogwood (Cornus florida) dominates the area’s understories in April.

There is one tree in this part of the country, however, that shows its amazing blooms in January and February while winter’s grip is still strong.  Ozark witch hazel (Hamamelis vernalis) is restricted to the Ozark Highlands of Missouri and Arkansas, where it grows along the rocky creeks and streams that dissect this ancient landscape.  I have long wanted to see its striking blooms, but despite my many wintertime hikes throughout the Ozarks, I have never found myself in the right place at the right time – until a few weeks ago when I hiked the Mina Sauk Trail at Taum Sauk Mountain State Park.  I found these plants growing below Mina Sauk Falls and along Taum Sauk Creek below, and even though it was the first weekend of March (and the very first warm day of the season), many of the plants had already passed their peak bloom.  Fortunately, I was able to find these several plants with flowers still in good shape.

There is only one other species in the genus – eastern witch hazel (Hamamelis virginiana).  Although distributed widely across eastern North America, it is restricted in Missouri to these same St. Francois Mountains where I saw H. vernalis.  The two species are very similar by the characteristics of their foliage but can be easily distinguished by floral characters.  Hamamelis virginiana blooms in fall rather than winter, and its flowers, while nearly twice the size, rarely show the amount of red on the inner calyx that is seen in this species.  Hamamelis vernalis flowers are also quite fragrant, having what has been described as a “vanilla” scent.  The photographs here show the rather unusual color range of the flowers of this species, which can vary from orange to deep red to deep yellow.  I suspect that flower color also changes with age, in that petals are initially deep red and later fade to yellow, as in the photo below.  It’s difficult to explain why H . vernalis is restricted to the Ozark Highlands while H. virginiana occurs so broadly, but the Ozarks are a well-known refugium for a number of other plants and animals, especially Ice Age relicts.

Sitting on a rhyolite ledge overlooking Taum Sauk Creek as I ate lunch, I wondered about the pollination biology of a plant that flowers during winter.  It was a warm day – certainly an unusual occurrence during the period in which this plant flowers – and even still it was too early in the season for a lot of insect activity.  I watched one of the nearby plants as I ate to see what insects came to the flowers, and for a time all I saw were a couple of European honey bees.  Clearly, the plant did not evolve in association with this now ubiquitous insect.  I continued watching, and at last I saw a native insect visiting the flowers – a large species of hover fly (family Syrphidae), perhaps something in the genus Helophilus.  After taking a few more photographs (unfortunately, none of the fly), another of the same species visited the plant.  Flies in general are famous for appearing during warm days in winter, and I wonder if the unusually extended bloom period of this species is intended to take advantage of those few, unpredictable days during winter when temperatures are sufficient for flies to become active.

Photo Details: Canon 100mm macro lens on Canon EOS 50D
Photo 1: ISO 100, 1/200 sec, f/11, MT-24EX flash w/ Sto-Fen-Puffer diffusers.
Photo 2: ISO 200, 1/200 sec, f/5.6, ambient light.
Photo 3: ISO 100, 1/60 sec, f/9, flash w/o diffusers.
Photo 4: ISO 200, 1/250 sec, f/5.6, ambient light.

Copyright © Ted C. MacRae 2010

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Great Plains Ladies’-tresses

First things first—everyone who participated in the quiz in the previous post correctly identified the orchid flower in the photo as belonging to the genus Spiranthes, and a few were on the right track with their species suggestion of S. cernua.  However, Scott Namestnik from Indiana and Doug Taron from Illinois, were the only ones who recognized it to be a close relative of that species, the recently-described S. magnicamporum.  Nice job!  The plants in these photographs were found during early October in the dry dolomite glades of White River Balds Natural Area in southwestern Missouri (part of Ruth and Paul Henning Conservation Area).  The creamy white inflorescences stood in stark contrast to the russet big bluestem (Andropogon gerardii) and rusty gold Indian grass (Sorghastrum nutans) stems that dominated the rocky landscape.

Spiranthes¹ is one of the more complex genera of North American orchids, seven of which are known to occur in Missouri (Summers 1985).  Spiranthes magnicamporum² is closely related to S. cernua and was only recently (1975) described as a distinct species.  Conclusive separation of the two species requires microscopic examination of the seeds (those of S. magnicamporum are monoembryonic, whereas a large percentage of the seeds of S. cernua are polyembryonic) (Luer 1975).  In the field, however, S. magnicamporum can generally be distinguished from S. cernua by its spreading rather than appressed lateral sepals and absence of basal leaves at the time of flowering³.  It is likely that many previous records of S. cernua in Missouri actually refer to this species, as both occur throughout much of southern Missouri and sporadically in northern Missouri (refer to the USDA Plants Database Missouri county level distributions for S. cernua and S. magnicamporum).  However, they are ecologically isolated in that S. cernua prefers wet lowlands with acidic soils, while S. magnicamporum is typically found in drier uplands with calcareous soils.  Both species are late-season bloomers, but S. magnicamporum blooms even later (mid-September into November) than S. cernua (mid-August to mid-October) and has more fragrant flowers.

¹ From the Greek speira—σπειρα,—”coil,” and anthos—ανθος,—”flower,” referring to the coiled or spiraled spike of flowers common in the genus.

² From the Latin magnus, “large,” and campus, “plain,” meaning “of the Great Plains” in reference to the primary geographic area where this species is found.

³ My identification of these plants as Spiranthes magnicamporum was confirmed by Dr. George Yatskievych, author of Steyermark’s Flora of Missouri.

Orchids as a whole exhibit highly specialized pollination biology, and species of Spiranthes are no exception, with the spiral arrangement of their flowers evidently an adaptation to pollination by long-tongued bees (e.g. bumblebees, Bombus spp., and megachilid bees) (van der Cingel 2001).  Flowers are protandrous, i.e., they are functionally male when they first open and become functionally female as they age, and open sequentially from the base, resulting in female flowers on the lower inflorescence and male flowers on the upper inflorescence.  Thus, bee pollinators tend to act as pollen donors when visiting lower flowers and pollen recipients when visiting upper flowers.  Pollinia from male flowers are attached to the bee’s proboscis as it tries to access nectar secreted into the base of the floral tube.  When visiting a plant, bees start at the bottom of the inflorescence and spiral up to the top before flying to the next plant.  The reasons for this behavior, called acropetal movement, are not fully understood but could be related to the tendency for nectar rewards to be greater in the lower flowers.  Whatever the explanation, the result is to promote outcrossing between neigboring plants.

While specific insect pollinators have been documented for a number of Spiranthes spp., apparently the only account of pollination in S. magnicamporum is documented by Jeffrey R. Hapeman, author of the website Orchids of Wisconsin:

I have seen a bumblebee (Bombus nevadensis ssp. americorum) pollinating Spiranthes magnicamporum in a prairie in southeastern Wisconsin. After visiting a number of inflorescences, the bee began to vigorously scratch at the pollinia on its proboscis, trying to remove them. The bee became so involved in trying to remove the pollinia that it fell to the ground, where it was easily captured. The specimen was determined by Steve Krauth, and is deposited in the Insect Research Collection at the University of Wisconsin-Madison. Apart from this observation, there are no published accounts of pollination of S. magnicamporum.

Photo details:
All photos: Canon 100mm macro lens on Canon EOS 50D (manual mode), ISO 100, MT-24EX flash w/ Sto-Fen diffusers.
Photo 1: 1/160 sec, f/14, flash 1/2 power.
Photo 2: 1/250 sec, f/16, flash 1/4 power.
Photo 3: 1/250 sec, f/20, flash 1/4 power.
Photo 4: w/ 36 mm extension tube, 1/250 sec, f/16, flash 1/8 power.

REFERENCES:

Luer, C. A.  1975.  The Native Orchids of the United States and Canada Excluding Florida.  The New York Botanical Garden, 361 pp. + 96 color plates.

Summers, B.  1981.  Missouri Orchids.  Missouri Department of Conservation, Natural History Series No. 1, 92 pp.

van der Cingel, N. A.  2001.  An atlas of orchid pollination: America, Africa, Asia and Australia. A. A. Balkema, Rotterdam, Netherlands, 296 pp.

Copyright © Ted C. MacRae 2009

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Friday Flower: Yes, it’s an orchid…

Photo details: Canon 100mm macro lens on Canon EOS 50D (manual mode), 36 mm extension, ISO 100, 1/250 sec, f/16, MT-24EX flash @ 1/8 power w/ Sto-Fen diffusers.

…but what kind? Identifying the genus should be relatively easy, but I suspect a species identification will be more of a challenge.  I’ll provide a little information and even a couple of literature sources that might be useful for achieving a specific determination.

  • Date of photograph: October 5, 2009.
  • Location: White River Balds Natural Area, Taney County, Missouri.
  • Habitat: Dolomitic limestone glade.

Answer and more photos will be posted shortly, so give it your best shot. Think big!

REFERENCES:

Luer, C. A.  1975.  The Native Orchids of the United States and Canada Excluding Florida.  The New York Botanical Garden, 361 pp. + 96 color plates.

Summers, B.  1981.  Missouri Orchids.  Missouri Department of Conservation, Natural History Series No. 1, 92 pp.

Copyright © Ted C. MacRae 2009

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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.

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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