Know Your Natives – White Snakeroot

White snakeroot (Ageratina altissima*), formerly Eupatorium rugosum, of the Aster, Sunflower, or Composite (Asteraceae ) family has a multitude of small white flowerheads late in the growing season. It occurs from Texas to eastern North Dakota, across all states to the Atlantic and Gulf Coasts, but of limited occurrence in Florida and Georgia. In Arkansas, white snakeroot, the only species of Ageratina in the state, occurs statewide. The genus name (originating from “ageratum”) is based on Greek words for “unaging”, in reference to the long-lasting color of the inflorescence. The specific epithet means “tallest”, in comparison with other species of the genus. The common name relates to use of the roots by Native Americans for treating snake bites. Habitat preference is mesic, fertile soils in riparian to upland deciduous woodlands and woodland edges and thickets. Plants are poisonous.**

White snakeroot, a perennial herbaceous plant, may have a single, foot-tall stem or several stems to 3+ feet tall. Plants are mostly glabrous (hairless), with minute pubescence near the inflorescence. Stems have well-spaced, opposite, decussate leaves with slender petioles with widened bases. Between the widened petiole bases of paired opposite leaves, an “edge” connects the bases resulting in prominent nodes along larger stems. The root system consists of small basal caudices with a mass of long-radiating, near-surface, wiry roots.

Photo 1: New spring growth arising from the caudex. Photo – March 22.
Photo 2: A small plant in a deciduous woodland setting. Leaves and branches are opposite and decussate. Photo – August 10.

Axillary branching occurs in the upper portion of plants. While shorter plants with few branches remain erect without support, larger plants that have numerous branches in their upper portion tend to lean or recline when lacking support. Stems and branches, mostly in opposite pairs, are ascending and rigid with branches rising from stems at about 45 degrees. Branches may be 16 inches long with sub-branches at their upper portion that may be 8 inches long. Overall, stems and branches are yellowish green, with lower stems and branches transitioning to reddish brown and becoming hardened. Larger stems have pith-filled centers. With heavy seed production, thick stands may develop in preferred habitats. 

Photo 3: This plant (seen from above), has characteristic opposite leaves and axillary branches growing from nodes (red arrow on left). Photo – September 23.

Leaves are ovate-lanceolate, to 6 inches long, becoming lanceolate and decreasing in length above. Blade margins are sharply serrate. 

Photo 4: Petiolate leaves vary from ovate-lanceolate lower leaves to lanceolate upper leaves. Venation and marginal serrations are prominent. Adaxial side shown on left and abaxial side on right.

White snakeroot blooms in mid-fall. A large plant may have stems that divide into a dozen or more branches, each with sub-branches. The inflorescence consists of “large cymes” (3 to 4 inches across) of composite flower heads. Flowerheads are uniformly distributed across the cymes, creating a rounded inflorescence. Flowering sequence gradually shifts from central cymes to outer cymes.

A colony of white snakeroot in early bloom with its masses of flowers of the purest white is one of the most beautiful sights in the deciduous forest. The flowerheads are composed of 1/4-inch-long, white disk florets set into 1/8-inch-long, elongate, pale green involucres. Involucres comprise a single series of clasping, linear, appressed phyllaries (bracts) with acute apexes. Each flowerhead, to ⅛ inch across, bears 12 to 30 tubular florets with white corollas surrounded by pappus bristles. At anthesis, corollas have 5 flared, acutely triangular lobes. Stamens, a slightly off-white color, form an elongate central anther ring. The style, exerted through the anther ring, presents the pollen, after which its apex divides to expose two long stigmatic surfaces. The stigmatic surfaces are thin and twisty, so that cymes, with all flowerheads facing the sunlight, have a delicate appearance.

Photo 5: This single large cyme is composed of a half dozen smaller cymes, each with 10 to 25 flowerheads. Photo – October 14.
Photo 6: Involucres are formed by a single series of linear, appressed phyllaries. Pedicels and peduncles are puberulent.

Fertilized florets produce narrowly oblong, black achenes (“cypselae” in the Aster family) topped with the radiating bristles of the pappus (morphologically, highly modified sepals). Achenes are ribbed and glabrous. Seed dispersal is by wind.

Photo 7: As elongate black achenes mature, the pappus flares to provide support for wind dispersal. Photo – October 14.

White snakeroot, retaining a fresh appearance into fall, produces masses of white flowers for a month or more. It can be quite showy in partial to full shade gardens, especially striking if other fall bloomers (goldenrods, asters) are nearby. In a garden setting, plant spreading may need to be controlled by select plant removal and deadheading to prevent aggressive self-seeding. Also, it is a nice plant for naturalized areas. A variety of insects feed on the leaves, nectar and pollen. It is not eaten by deer.

Photo 8: In natural woodlands, white snakeroot may form dense stands. Photo take on Mt. Magazine, September 26.

* Two varieties have been classified: A. altissima var. altissima (white snakeroot) and A. altissima var. roanensis (Appalachian snakeroot). Varieties are identified by slight variation in the length and shape of the phyllaries. In Arkansas, A. altissima var. altissima is the only recognized variety.

** The entire plant, containing tremetol, is poisonous. When pioneers moved west into the unsettled eastern forest, cattle were often left to graze in open woodlands, ideal habitat for white snakeroot, and people developed “milk sickness” from tainted milk and meat. Thousands died before white snakeroot was identified as the source of the toxin. With current herd and milk production practices, milk sickness is now uncommon. 

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Boott’s Goldenrod

Boott’s goldenrod (Solidago arguta subsp. caroliniana var. boottii*) of the Aster or Sunflower (Asteraceae) family is a medium-size goldenrod with large ovate-lanceolate basal leaves. The genus name is from Latin for “to make whole” or “to heal” in reference to purported health benefits derived from some species of the genus. The varietal name honors Francis Boott, 19th-century American physician and botanist. Boott’s goldenrod, with limited U.S. occurrence, ranges primarily across the western two-thirds of Arkansas, extending into nearby portions of Louisiana, Texas, Oklahoma and Missouri. Preferred habitat is mesic to dry sandy soils in partially shaded woodland openings and borders. It is one of 28 species of goldenrod that occur in Arkansas.

Photo 1: This 3-inch-long caudex bears both 3½ foot tall stems of the current season and short, leafy fall growth. The white shape extending from the base of the right stem is a new rhizome that will terminate with a subsidiary caudex. Photo – mid October.

One to a half-dozen 3 to 4-foot stems with base-diameter of ⅛ to ¼ inch arise from a stout, branched caudex. They are erect to ascending, longitudinally ribbed, and hard and tough with age, with multiple branches, to 2-3 ft long, above. The sunny side of the light green stems and branches becomes reddish brown. Stems are usually glabrous below, becoming appressed pubescent into the inflorescence.

Leaves are borne predominantly toward the base of the plant and are usually present at flowering. Larger leaves below have ovate-elliptic blades to 5½ inches long and 3 inches wide on winged petioles to 5¼ inches long. Leaf margins are sharply toothed. Leaf pubescence is somewhat variable, but upper leaf surface tends to be glabrous while lower surface has short, erect hairs, especially along veins.

Photo 2: This early April photo shows leaves with ovate-elliptic blades and long petioles. Petioles are winged to their base by narrow bands of decurrent blade tissue.
Photo 3: Upper (above) and lower surfaces of larger leaves have irregular pinnate venation.

Median and upper stem leaves gradually become smaller and lanceolate, marginal serrations become less prominent, and petiole length decreases. Leaves near the branch tips are less than ¼ inch long and ⅛ inch wide, sessile, and with entire margins. 

The inflorescence, in September and October, occurs mostly as a panicle with long recurved branches and heads oriented upward. The inflorescence is usually broad and open, less often elongate and narrow. Flowering sequence along the branches is from tip to base.

Photo 4: Flowerheads are secund, i.e., oriented upwards. Racemes and individual flowerheads are subtended by a leafy bract. Note strigose pubescence and small elliptic leaves with entire margins. Photo – early September.

Flowerheads, about ⅜ inch long, are set in elongate, cup-like involucres comprising 3-4 series of spirally arranged, overlapping bracts or “phyllaries”. Inner phyllaries are longer than the outer and transition to bracts along the pedicels. Involucres are nearly glabrous; pedicels and rachises are covered with short strigose pubescence.

Photo 5: The main raceme (left to right across photo) bears short, secondary racemes composed of two flowerheads each, along with a single flowerhead at far right. Phyllaries of the involucres transition to bracts along the pedicels. Photo – early September.

The bright yellow flowerheads have 2-8 pistillate ray florets (with pistils only) which surround up to about a dozen perfect disk florets (with pistils and stamens). Ray florets have strap-shaped corollas with rounded, notched tips and slightly exserted styles with bifurcated stigmas. Disk florets are tubular with 5-lobed corollas and 5 stamens with thin filaments topped with elongate anthers. Anthers are fused into a ring that surrounds the style, which elongates through the anther ring, carrying the pollen above the corolla where it is available to pollinating insects. Once fully exserted, the pair of linear stigmatic surfaces divides and becomes receptive to pollen. Corollas are surrounded by hair-like modified sepals (pappus) which attach to the tip of flat-topped, elongate ovaries.

Photo 6: Flowerheads here have 3-4 strap-shaped ray florets and up to a dozen or so tubular disk florets. The linear stigmatic surfaces of both ray and disk florets are divided. Pistils of disk florets are significantly larger than those of ray florets.

Fertilized florets produce flattened, 1/16 inch long, minutely pubescent, elongate achenes topped by bristly hairs (pappus). The hairs allow achenes to be dispersed widely by wind.

Photo 7: In this late November photo, the gradual decrease of leaf size up-stem into the branches can be seen. Achenes are ready for dispersal on next windy day.

For a garden or natural area, Boott’s goldenrod can add height and bright yellow fall color. This species has a smoother appearance than many of the other Arkansas goldenrods. It is not especially aggressive by seed or caudexes. It is not favored by deer. Flowerheads and leaves are an insect magnet.

Boott’s goldenrod has several general characteristics which help separate it (with difficulty) from the 26 other Solidago species that have yellow flowerheads: 1) one to several tall stems with widely separated leaves, 2) large ovate-elliptic basally disposed leaves, 3) an inflorescence with widely separated straight to recurved branches and sub-branches, 4) secund flowerheads, and 5) overall smooth appearance of the plant.

*Boott’s goldenrod has been identified by Arkansas authorities as Solidago arguta subsp. caroliniana var. boottii. Other authorities have identified it as Solidago arguta var. boottii or Solidago arguta subsp. bootii.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Carolina Elephant’s-Foot

Carolina elephant’s foot (Elephantopus carolinianus), of the Sunflower or Aster or Composite (Asteraceae) family, is a herbaceous perennial with leafy stems and small lavender flowers. In the U.S., the species occurs from eastern Texas to southeastern Kansas, east to Pennsylvania and New Jersey, and southward to the Atlantic and Gulf Coasts. In Arkansas, it occurs statewide. The genus name and common name refer to the large, flat, prostrate basal leaves of some species of the genus (though not this species). The specific epithet and common name refer to the plant’s original recognition and description from the Carolinas. Preferred habitat is well drained yet moist soils of partially sunny lowlands or open woods and along woodland edges and streams. The plant has a small knobby root crown with long, descending, tough and ropy fibrous roots.

Photo 1: Root crowns are relatively small in comparison to the ropy roots. Stubs of previous years’ stems are at center of this crown. Rings on stems result from clasping petioles of dropped leaves. Diameter of upper stem is 3/8 inch. Photo – September 16.

Plants generally have a single stem 1 to 2 feet tall, but may reach a height of 4 feet. Stem leaves at the base are closely spaced, to ¼ inch apart, becoming widely separated (and clearly alternate) above. From lower to mid-stem, clasping leaves form a sheath around the stem. Stems are densely hirsute below, with hairs becoming sparser and somewhat more appressed in the inflorescence.

Photo 2: Clasping lower leaves and densely pubescent lower stem. Photo – July 5.
Photo 3: Leaves are widely spaced along terete stems. Branches diverge from stems at leaf bases. Photo – August 16.

Leaves, to 8 inches long and 3½ inches wide, are dark green above and medium green beneath. Below the inflorescence, they are clasping, with margins coarsely serrate to scalloped; within the inflorescence, smaller leaves are sessile and subentire. Upper and lower surfaces are sparsely hirsute, with longer and more dense pubescence along the veins beneath.

Photo 4: Stems continue and branches originate from sheathed bases of petioles. Large leaf on upper left is 6¼ inches long and 2½ inches wide.

In most composites, the so-called flower is actually a head, an inflorescence comprising a group of tightly clustered, small flowers that mimic a single flower. In the genus Elephantopis, these primary heads are further clustered into secondary heads. In E. carolinianus, 3-20 such heads are grouped together, at the tip of 1-4 inch peduncles, into secondary heads about ⅜ inch high and ⅝ inch broad, subtended by 3 leaflike bracts. Each primary head is itself surrounded by an involucre of about 8 phyllaries (bracts), with the 4 inner phyllaries longer than the outer. These thin, lanceolate to linear phyllaries are whitish proximally and green distally. Each involucre surrounds 4 flowers.

Photo 5: The 1 inch peduncle (as shown) was separated from three floral bracts and about a dozen primary heads with involucres. All flowers are past anthesis. Developing fruit can be seen at top of photo. Photo – September 22.

Blooming occurs over a month in September-October with individual flowers lasting for one day. Flowers are ¼ inch long, with lavender (sometimes white), 5-lobed corollas. The lobes are positioned to one side of the flower, creating bilaterally (rather than radially) symmetrical corollas. All 4 flowers within a single involucre bloom on the same day. And because they are positioned radially around the involucre’s center, each primary head of 4 corollas appears to form a single 4-petaled, 20-lobed flower! With only a few involucres blooming at one time, a plant may continue to produce “flowers” for a month or more.

Lobes of the corolla join abruptly to form a slender tube twice as long as the lobes. The corolla tube arises from the summit of a bullet-shaped ovary, each ovary bearing, just outside the corolla tube, 5 awnlike scales that taper to bristle-like tips. These awns, which are modified sepals, constitute the so-called pappus, a structure in the Composite family typically involved in dispersing the 1-seeded fruits.

Photo 6: Leafy bracts subtend a cluster of involucres. As shown, flowers of one involucre are in bloom. Photo – September 23.

Flowers have 5 stamens and 1 pistil. White stamens have thread-like filaments bearing relatively large elongate anthers. The anthers are fused to each other to form a tube through which the style passes at anthesis. As the style extends like a plunger through the anther tube, clingy white pollen is pushed above the corolla and becomes available to pollinating insects. Thereafter, the style bifurcates to expose the elongate stigmatic surfaces to pollen, typically from the flowers of other heads.

Photo 7: This single involucre was separated to show four flowers with ovaries, styles with pollen-covered, bifurcated stigmas, and modified sepals (the pappus of seed-dispersing awns).

With fertilization, ovaries mature into elongate, flattened, 1-seeded, brown achenes. Achenes are ribbed and have very short hispid pubescence. They are tipped with 5 awns slightly longer than the achenes. With drying, awns become straight and stiff, so that, together with the pubescence, dispersal may be effected by passing mammals.

Photo 8: The bullet-shaped, brown, ribbed achenes have stiff awns and hispid pubescence. Squares equal ¼ inch. Photo – November 17.

Carolina elephant’s foot has an interesting open structure and flower heads that are unusual for the Aster family. Plants would be an interesting addition in a garden setting, but due to aggressive self-seeding, this species is perhaps better suited for a natural area. It does well in light shade and can survive temporary dry periods. It is nibbled by deer.

In addition to Carolina elephant’s foot, two other species of Elephantopis occur in Arkansas: smooth elephant’s foot (E. nudatus) and hairy elephant’s foot or devil’s grandmother (E. tomentosus). Carolina elephant’s foot can be distinguished from the other two by its large, well-developed stem leaves. Smooth elephant’s foot and hairy elephant’s foot have leaves that are primarily basal––they lie flat on the ground, like feet.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Jumpseed

Jumpseed (Persicaria virginiana), formerly Polygonum virginianum, of the Buckwheat (Polygonaceae) family has slender knobby terminal stems that bear long racemes of tiny white flowers. In the U.S., it occurs throughout the East, from eastern Texas north into eastern Nebraska and southern Minnesota, and east to the Atlantic and Gulf coasts. In Arkansas, it occurs statewide. The genus name originates from Greek words for “arrow-shaped,” a description of the leaf-shape of many Persicaria species. The specific epithet refers to the species’ original discovery in Virginia. Another common name is painter’s palette, based on a splash of color across some of the larger leaves. The name “jumpseed” describes the kinetic dispersal of the seeds. Preferred habitats are areas of rich, relatively moist, woodland soils in shade to partial shade.

Photo 1: Jumpseed has large lower leaves that may have a splash of darker color at mid-leaf. Other plants shown: lady fern (Athyrium filix-femina), spotted jewel weed (Impatiens capensis), and aniseroot (Osmorhiza longistylis).

Jumpseed is an herbaceous perennial with shallow reddish, branching rhizomes and long fibrous roots. Current year’s stems grow from tips of dominant rhizomes, and new rhizomes extend from near the same tips. Older portions of the rhizomes remain viable.

Plants may be 3 to 5 feet long, but plant height is shorter due to the somewhat reclining stems. The sturdy terete stems are divided by several prominent, hairy, widely spaced, leaf-bearing nodes. Main stems of mature plants are mostly glabrous below and pubescent in the inflorescence with short ascending hairs. Stems are a light green early in the growing season, but become darker green with reddish shading. In open conditions, plants can bear axillary branches up to 2 ft long from the upper nodes.

Photo 2: Two stems from the same rhizome display widely spaced main-stem nodes and axillary branching. Photo – September 6.

Nodes along the main stem, especially earlier in the growing season, bear tubular sheaths to ¾ inch long that are heavily covered with glassy hairs. Such sheaths, termed ocreae (singular ocrea), are modified stipules attached like a collar at the petiole base and characteristic of several genera in the buckwheat family. Jumpseed ocreae are truncate at the apex. Early in the growing season, they are the same color as the stems, but become dry and brown by the time of flowering. 

Jumpseed has simple, alternate leaves. Lowermost stem leaves (no basal leaves), to 7½ inches long and 4 inches wide, are ovate-elliptical, becoming smaller and more elliptical along the upper portion of the main stem and lower portion of some axillary stems. Larger leaves tend to have ascending auricles (ear-shaped projections) where the blade joins the pedicel. Leaf color is medium green above and a silvery lighter green beneath; larger leaves often have a dark chevron-shaped overlay at midleaf, oriented toward the leaf apex. Leaf margins are entire. Venation is pinnate, with secondary veins that arch toward the leaf apex. Petiole length is about 1 inch long, becoming shorter above. Petiole bases clasp the stem. Short, hooked, hispid pubescence covers upper and lower leaf surfaces, with the upper surface bearing longer hairs and feeling rougher than the lower. Short, hispid pubescence may also be scattered along the petioles and upper stems.

Photo 3: Hairy truncated sheaths or ocreae extend from nodes. At right, several sheaths are in contact before becoming separated as the stem elongates. Note clasping petioles. Photo – April 20.

Photo 4: The lower ovate-elliptical leaves often have a dark chevron-shaped overlay. Venation is pinnate. Midveins of upper (left) and lower (right) leaf surfaces are expressed. Leaf on left has auricles. Photo – April 20.

Racemes, from 3 inches to 2 feet long, bear well spaced, easily discernable clusters of one to three flowers. Nodes, sheaths and rachis of racemes are clothed with scattered, straight, stiff, ascending hairs. Lengths of terminal and axillary racemes may equal or exceed length of the main stem.

Photo 5: Upper portion of main stems is arching, while axillary branches and their inflorescences tend to spread downward. These upper leaves are elliptical. Along racemes, spacing of sheaths is regular. Photo – August 21.

In mid-summer, tiny, dangling flowers, about 1/8 inch long, bloom from the base of the racemes upward. Short pedicels are hidden within the sheaths. At fruiting, they are seen to be about half as long as the flower.

Flowers have four greenish white to white (occasionally pinkish), acute, flaring sepals (petals are absent), four or five stamens, and a pistil with a deeply divided style. Stamens have slender, white filaments topped with ball-like white anthers. They project slightly above the perianth, while the styles extend slightly past the stamens.

Photo 6: Flowers have five stamens and one divided style. At bloom stage, pedicels are hidden within protective sheaths. Sepals and styles persist as seeds mature.

With fertilization, pedicels reach their maximum length, about 1/4 inch, and sepals close around the developing fruits. Mature, spindle-shaped, pointy, 1-seeded fruits (achenes) are bent downward on reflexed pedicels. When touched, for example, by a passing animal, tension stored in the pedicels causes achenes to be forcefully thrown (“jumpseed”) to several feet. Seeds can also be dispersed when the hooked styles latch onto a passing animal.

Photo 7: Ovoid 1-seeded achenes, covered by dried sepals, are ¼ inch long from base to tip of styles. Tension in the pedicels, now exposed, causes achenes to jump from the raceme when touched. Lower side of a leaf is shown in background. Photo – October 18.

Jumpseed may be appropriate as an accent plant in a partially shady, low-wind garden with rich mesic soil. Its rhizomes seem to extend slowly. For a larger natural area, it may be effective as a groundcover (new plants can be established from root division, and plants self-seed readily). Plants provide coarse texture and their oddly proportioned leaves and inflorescences add interest. They are not bothered by deer or rabbits.

In addition to Persicaria virginiana, 13 other species of the genus occur in Arkansas, of which three are not native. All 14 species have sheathed (ocrea-bearing) stems and small flowers in racemes. Persicaria virginiana can be distinguished by its woodland habitat; large, more ovate leaves; exceptionally long and slender spike-like racemes; well-separated flower clusters; and exserted styles.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Tall Coreopsis

Tall coreopsis or tall tickseed (Coreopsis tripteris) of the aster or sunflower family (Asteraceae) is the tallest of eight coreopsis species native to Arkansas. It occurs primarily from eastern Texas north to Iowa and Wisconsin and east to Pennsylvania and the Florida panhandle. In Arkansas, it grows throughout much of the state except for lower areas of the Mississippi Alluvial Plain and West Gulf Coastal Plain. The genus name originates from Greek words meaning “like a bug,” in reference to the bug-like appearance of the fruit. The specific epithet is from Greek words for “three” and “wings” in reference to the plant’s trifoliate leaves. Preferred habitats include sunny to partially sunny sites in various well-drained yet mesic soils in prairies, rights-of-way, thickets, open woodlands, and wood margins, often along creeks.

Tall coreopsis is a herbaceous perennial that develops a tough caudex from which long ropy roots and rhizomes radiate. Rhizomes are segmented; each segment is subtended by an encircling growth-ring marked by a brown triangular bract. New clonal plants form at the ends of rhizomes. Caudices support one to several erect to leaning stems up to eight or more feet tall, unbranched except above in the inflorescence. Stems are slender, round, and hollow, glaucous when young, light green and glabrous, with slight rounded ridges. They bear widely spaced, opposite, decussate pairs of mostly compound, trifoliate leaves, about 2-3 inches apart. Petioles of the paired leaves have broadened bases which extend to form a ring around the stem.

Photo 1: This clonal rootstock has a “parent-rhizome” extending to lower left and a single stem extending to upper right. Rootstock has ropy roots and segmented new rhizomes and stems.

Trifoliate leaves measure about 5 inches long (including 1+ inch petioles) and 3¾ inches wide, with a central leaflet and two (sometimes four) opposite wide-spreading lateral leaflets. Leaflets are lanceolate, mucronate at the tips, the central leaflet to 3½ inches long, the laterals about 1 inch shorter. Lateral leaflets are sessile; central leaflets are stalked on petiolules* to ¼ inch long. Minute, soft pubescence (visible with magnification) covers upper and lower surfaces of leaf blades. Simple leaves occur in pairs at the base of floral branches, decreasing in size to become the bracts and bracteoles of the inflorescence. Crushed leaves have an anise scent.

Photo 2: In this late June photo, glaucous stems bear compound, trifoliate leaves with lanceolate leaflets. Stems have slight longitudinal ridges.
Photo 3: Compound leaves occur along the stem with simple leaves at and within the inflorescence. Secondary and tertiary veins are obscure. Photo – July 26.

The inflorescence, at the upper portion of the stem, consists of flowerheads supported by erect glabrous pedicels (to 1 inch), peduncles (to 2½ inches), and branches (to 6 inches). The lower branches tend to be longest. Blooming extends over a month in mid-summer. 

Photo 4: In this early August photo, the tall stems tend to lean. Flowerheads have a dark central disk.

At anthesis, flowerheads have a bowl-shaped, double involucre: an inner series of 6-10 yellow-green, broader, longer phyllaries, reflexed at their tips, and an outer series of narrower, bright green, ascending phyllaries.

Flowerheads are 1+ inch wide, with 6-10 bright yellow ray florets and 40-50+ disk florets. The showy ray florets are sterile, serving only to attract pollinators. Disk florets have reddish brown, tubular corollas (with five short, triangular, recurved lobes) and an equal number of stamens that become strongly exserted. Anthers are fused into a tube through which the style emerges, carrying and exposing the yellow pollen to pollinating insects. With pollen of the disk florets dispersed, the style divides and recurves to expose two elongate stigmatic surfaces. The central disk is flat before disk florets open, but becomes “prickly” as florets bloom sequentially from perimeter to center.

Photo 5: Pubescent inner phyllaries and linear-oblong outer phyllaries (brown tipped) can be seen on flowerhead at right. At left, corollas of sterile ray florets are fully extended while some disk florets are releasing pollen. One disk floret (upper left) has matured to show its bifurcated stigma.
Photo 6: Lower surface of flowerhead, right, shows dull yellow tips of the inner phyllaries at the base of the bright yellow ray florets as well as the green, ascending outer phyllaries. Flowerheads have been removed from pedicel/peduncle at left.

Disk florets that are fertilized form achenes (“cypselae” in the aster family) that vary in appearance depending on position within the central disk. Achenes at and near the perimeter are flattened, thin, and ovoid with narrow lateral wings as well as a concave and a convex side. Achenes at and near the disk center are elongate and wingless. All achenes are dark brown to black at maturity and truncate at apex and base. Seed dispersal is by wind and gravity, and sometimes subsequently by water.

Photo 7: Three disk florets are shown at left. On the flowerhead, three disk florets have exserted anther tubes and pollen has been released. Three maturing winged achenes (from a different flowerhead) are shown at lower left.

In a garden setting, tall coreopsis would be best suited for a natural area, due to its ability to develop colonies from rhizomes and self-seeding. Foliage and flowers are both attractive. Plants lean when bearing flowerheads due to the height of their slender stems. It is not bothered by deer.

Tall coreopsis, one of eight native Arkansas species in the genus, can be distinguished from the other species by a combination of its 1) height, 2) trifoliate compound leaves composed of lanceolate leaflets, 3) late period of bloom, and 4) reddish-brown central disk.

*Petioles are stalks of leaves. Petiolules are stalks of leaflets. Pedicels are stalks of a single flower or flowerhead. Peduncles are stalks of cluster of pedicels.

Article and photographs by ANPS member Sid Vogelpohl

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ANPS Fall 2019 Meeting Reminder

⏰HURRY – Only three days left!

Make your reservations now for ANPS Fall Meeting

Arkadelphia – Caddo Valley

Hotel Location:

Hampton Inn
108 Malvern Road
Arkadelphia, AR 71923
870-403-0800

Meeting Location:

Parks & Recreation Center, Feaster Park
2555 Twin Rivers Drive
Arkadelphia, AR 71923
870-246-5499

We have 15 double queens at $109 per night plus tax and 5 single queens for $89 per night plus tax. Breakfast is included. Be sure to mention that you are with ANPS when making your reservation. More rooms can be had at same rate if available.

Both Live Auction and Silent Auction on Friday Night – bring your wares!

Directions and Field Trip Details to follow next week.

In the interim, see the recent Fall issue of The Claytonia for more information or click here.

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Know Your Natives – Missouri Ironweed

Missouri ironweed (Vernonia missurica) of the Sunflower or Aster (Asteraceae) family has vivid purple to magenta composite flowerheads in midsummer. The primary area of occurrence extends from western Florida to eastern Texas, north and east to Indiana and southern Michigan. In Arkansas, the species occurs throughout most of the state, though apparently absent from portions of the Ozark Mountains. The genus name commemorates William Vernon, an English botanist who worked in Maryland in the late 1600s. The specific epithet means ‘of Missouri’, in reference to where the species was originally described. Preferred habitat consists of various moist to mesic loamy to sandy soils found in low lying areas of prairies, woodlands, and drainage ditches.

This sturdy herbaceous perennial has a root crown with a mass of whitish ropy roots. New stems sprout from the periphery of the crown as central old stems die and form a tough, hard rootstock. Rhizomes, to a foot long, may extend from this rootstock and terminate with an offset clonal plant.

Photo 1: Current-year stems grow from the root-crown (as shown) as well as from short rhizomes. Circular scars are remnants of old stems.

Missouri ironweed is an erect herbaceous perennial with one or more tough, erect, round, ridged stems 3-5(-6) feet tall. Stems are unbranched except at the upper nodes, where lateral branching terminates in flowerheads (see below), producing a large, showy inflorescence. Stems and branches bear a dense coat of short, soft, white, tangled hairs, especially along ridges, that continue onto peduncles (ca 1 inch long) and pedicels (ca ½ inch long).

Photo 2: Erect stems have leaves that are fairly evenly spaced and of similar size. Tomentose pubescence gives stems a whitish cast. Photo – May 16.
Photo 3: Branches, growing from upper leaf axils, attain a final height that is greater than the height of the short branches that terminate the main stem. Pubescence extends into the inflorescence. Photo – July 11.

The alternate, mostly lanceolate-elliptic leaves are widest at the middle. Larger stem leaves measure about 6 inches long and 1½ inches wide. Apexes are acuminate (gradually tapering); sessile bases (without petioles) to subsessile bases (very short petioles) are cuneate (wedge shaped). Bases may have a light purplish shading. Leaf margins have weak serrations with tiny white tips; serrations decrease toward base and apex. Upper leaf surface is medium green with a glabrous (hairless), whitish midrib, especially along the lower portion. Lower leaf surface is light to whitish green. The upper leaf surface feels slightly rough (the hairs are short and stiff but not readily seen), while the lower surface feels downy from the tomentose pubescence.

Photo 4: Leaves, with white-tipped serrations, have an acuminate apex and cuneate base. Lower surface (right) is densely soft pubescent which highlights venation.

Missouri ironweed’s inflorescence consist of flowerheads comprising up to 45 or more disk florets. Flowerheads are organized into small, flat-topped, cymose arrays terminating branches growing from the main stem and upper leaf axils. Flowering begins at the terminal clusters of the main stem and gradually spreads outward. A single stem may support 200 flowerheads. The overall inflorescence is flat-topped.

The heads’ urn-shaped involucres, ¼ inch long and wide, are composed of five to seven series of imbricated bracts, called phyllaries in the composite family. Phyllaries are tightly appressed their entire length, broadly ovate, and ciliate along the margins. They are dark green in the center, with lighter-colored margins that turn purplish when flowerheads are in bloom.

As heads start to flower, a thick broom-like cluster of hairs or bristles appears above the involucre. The “broom” constitutes the “pappus,”* a cluster of 12-20 hairs attached to the top of each ovary in the head. Purple to magenta tubular corollas soon push out of the encircling bristles, forming a showy, frilly flowerhead. Corollas have five sharply tipped recurving lobes, five stamens and a single pistil with an inferior ovary. The thin elongate anthers of the stamens form a tube around the style. As the style elongates through the anthers, it carries their pollen above the corolla, where it can be picked up by pollinating insects. Stamens soon wilt, after which the upper portion of style divides and coils backward to expose two elongate, now-receptive, stigmatic surfaces. Flowerheads are up to 1 inch across and ¼ inch high.

Photo 5: Flowering, that begins with the terminal cluster of the main stem, as well as at the center of individual arrays, gradually spreads outward. Lower portion of lateral branches are leafless. Photo – July 30.
Photo 6: A complete flowerhead is shown at center and a dissected flowerhead to left and right. A subtending leaf and a single floret are also shown. Note the distinctive, ciliate phyllaries and the brown pappus.

Shortly after florets pass anthesis, the corolla, stamens, and style drop off. The ovaries then dry to become flattened, elongate, ridged achenes (often referred to as cypselae in the sunflower family). Pappus fluffs up to form a circle of radiating hairs. As achenes dry, they are dispersed by breezes.

For a moist to wet natural area or native plant garden, Missouri ironweed provides good vertical structure and showy flowers that attract butterflies and other insects. It can do well in partial or full sun, as determined by its dependence on good soil moisture. It is not eaten by rabbits or deer. Once plants are established, control may be needed to restrict spread (dead heading and pulling clonal plants).

Missouri ironweed is one of seven ironweeds that occur in the state. Species with similar characteristics and that are also widespread in Arkansas are tall ironweed (Vernonia gigantea) and Baldwin’s ironweed (Vernonia baldwinii). Characteristics to distinguish Missouri ironweed, considering the somewhat variable leaf shape and pubescence of the three species, are 1) tomentose stems, branches, lower leaf surfaces and 2) appressed, pointed phyllaries with purplish margins, and 3) relatively large number of florets per flowerhead. Tall ironweed has blunt, appressed phyllaries, and Baldwin’s ironweed has appressed phyllaries with pointed recurved tips (involucres appearing bristly). Both have fewer florets per flowerhead on average than Missouri ironweed. Additionally, tall ironweed leaves are not tomentose on the undersides.  Where ranges overlap, ironweeds may hybridize.

*The pappus is a modified calyx, distinctive for the sunflower family. It takes a variety of forms, for example, bristles, scales, and awns. The pappus is often the agent of seed dispersal.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Baldwin’s Ironweed

Baldwin’s ironweed (Vernonia baldwinii) of the Sunflower or Aster (Asteraceae) family is a tall plant with a vivid purple inflorescence. In the U.S., it occurs principally from central Texas north to Nebraska and Iowa and east to Illinois and Arkansas. In Arkansas, it is found in the northwestern two-thirds of the state, primarily within the Interior Highlands. The genus name recognizes William Vernon, an English botanist who worked in Maryland in the late 1600s. The specific epithet recognizes William Baldwin, the original collector of the plant. Habitats have somewhat dry to slightly moist soils in open upland-woods, prairies and rights-of-way.

This sturdy herbaceous perennial has a whitish root-crown from which numerous ropy roots extend in all directions. As central stems die at the end of the growing season, short (less than an inch) rhizomes extend from the old rootstock and produce new stems immediately adjacent to the dead stems. The dead woody core of the older rootstock persists.

Photo 1: Dead stems at center of rootstock remain from previous years’ growth. Roots may be several feet long.

Stems bearing current year’s growth are erect, straight, tough, and brittle, typically 3 to 4 feet tall, but occasionally taller in preferred habitats. Stems are unbranched below the inflorescence. Floral branches, up to 10 inches long, spread widely from the main stem at the top of the plant. Stems and branches are typically tomentose (with short, soft, dense hairs), however pubescence is variable from plant to plant. Stems are round in cross-section.

Photo 2:  New growth of Baldwin’s ironweed is conspicuous in early spring. Photo – May 4.
Photo 3: Erect stems and branches are typically tomentose. Lower floral branches are shown. Photo – July 13.

Plants bear alternate, lanceolate to lance-ovate leaves with acuminate apexes and rounded tapered bases. Leaves have a slightly shiny, medium to dark green adaxial surface and a duller lighter green abaxial surface. Leaves may be short-petiolate or sessile. Pubescence of stems and branches extends onto the petioles, as well as to the midrib and lower surface of the leaf. The upper surface of the leaf blade may be scabrous (rough) to nearly glabrous. Leaves are large from plant base to floral branches, up to 8 inches long and 2 inches wide. Larger leaves are serrate along most of their length, with minute bristly yet soft teeth where veinlets terminate.

Photo 4: Leaf shape varies from lanceolate to lance-ovate. Display shows adaxial side of leaves except leaf at right. Leaf at right is 8 about inches long. Note serrate margins. Photo – July 14.

Flowerheads of Baldwin’s ironweed consist of disk flowers only (no ray flowers). The showy, somewhat flat-topped inflorescence measures a foot or more across and comprises a complex, branching arrangement of the heads in cymose arrays. Pubescence of the stems and branches continues onto the peduncles (stalks of groups of flowerheads) and pedicels (stalks of individual flowerheads). Flowering sequence is from the center of arrays outward, with those that terminate the main stem blooming first.

Photo 5: Inflorescence consists of numerous flat-topped cymose arrays of discoid flowerheads. Photo – July 13.

Flowering begins in mid-July and continues for several weeks. Heads consists of a compact group of 15 to 35 (typically 20 to 30) disk flowers, set in a campanulate (bell-shaped) to spherical involucre. The involucre comprises 5 to 6 series (layers) of tightly imbricated (overlapping) phyllaries (bracts). Phyllaries typically bear acute, recurved tips, giving the involucre a bristly appearance. 

As anthesis approaches, heads develop a rusty-brown, dense, broom-like cluster of bristles. (This is the “pappus,” a modified calyx and a distinctive character of the sunflower family that takes a variety of forms, for example, in addition to bristles, scales and awns. The pappus often is the agent of seed dispersal.) Each disk flower produces a purple tubular corolla (occasionally pink and rarely white), with five acutely tipped, recurved lobes, which pushes up through the pappus and, en mass, forms an attractive flowerhead. Anthers of the 5 stamens are fused into a ring around the style. As the style elongates through the anthers, it carries the pollen above the corolla, where it can be picked up by pollinating insects. Stamens soon wilt, after which the upper portion of the style divides and coils backward to expose two elongate stigmatic surfaces.

Photo 6: Immature flowerheads around the perimeter of photo show a brown, broom-like pappus prior to appearance of corollas. In addition to the typical purple, flowerheads may be pink or, rarely, white. Note bristly involucres.
Photo 7: As corollas open, their lobes recurve and anther-tubes and styles emerge. White pollen is pushed upward from the anthers and displayed by the emerging styles. Styles then split to expose stigmatic surfaces.

Shortly after anthesis, the corolla, stamens and style fall from the flower, and the ovary matures to become a flattened, elongate, 1-seeded achene (often referred to as a cypsela in the Aster family)–the familiar sunflower “seed” is one such achene. The pappus, attached at the tip of the developing achene, fluffs-up to form a circle of radiating, ascending bristles. As achenes dry, they separate from the receptacle and are dispersed by breezes.

Baldwin’s ironweed may be appropriate in a larger garden–at 3-4 feet tall, probably best suited for a naturalistic garden or natural area. It readily grows in various well drained soils. Its purple flowers are showy and attract butterflies and other insects. It is not eaten by deer.

Baldwin’s ironweed is one of seven ironweeds known to occur in the state. Species that have similar characteristics and are also widespread in the state are tall ironweed (Vernonia gigantea) and Missouri ironweed (Vernonia missurica). The best characteristic to identify Baldwin’s ironweed, considering the somewhat variable leaf shape and pubescence of the three species, is its recurved phyllaries which cause involucres to appear bristly. Tall ironweed has blunt appressed phyllaries, and Missouri ironweed has appressed pointed phyllaries. Additionally, tall ironweed leaves are not tomentose on the undersides and Missouri ironweed flowerheads consistently have over 30 florets (sometimes upwards of 50).  Where ranges overlap, ironweeds may hybridize. 

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Shrubby St. John’s-wort

Shrubby St. John’s-wort (Hypericum prolificum) of the St. John’s-wort (Hypericaceae) family is a short-lived, deciduous shrub with spectacular bright yellow flowers. The genus name originates from Greek words for “above” and “picture,” from the practice of placing flowers above a wall-mounted picture to discourage evil spirits on St. John’s feast day. The specific epithet, meaning “many,” is perhaps a reference to the plant’s numerous stamens. The species occurs from eastern Texas to Minnesota, thence east to the Atlantic and Gulf Coasts, although rarely encountered along the outer coastal plains. In Arkansas, it occurs throughout most of the state, except for low-lying areas of the Mississippi Alluvial Plain and the West Gulf Coastal Plain. Habitats vary from partial to full sun in well drained dry to mesic soils of upland areas, from rocky outcrops to stream banks, fields, and woodlands.

Shrubby St. John’s-wort, to 4 feet tall and broad, has one to several ascending branches which divide repeatedly into a myriad of twigs. Shrubs are dense and rounded on sunnier sites but more open in partial shade. Older trunks and branches bear loose, papery bark that exfoliates to expose underlying smooth reddish to orangish-brown bark. 

Photo 1: The thin smooth bark exfoliates with age.
Photo 2: With bright light, the shrub develops a dense rounded shape.

Twigs become brown and woody during their first year of growth. They bear evenly spaced pairs of subsessile, opposite leaves. Low ridges extending between leaf nodes become indistinct in the second year of twig growth, due to bark exfoliation. Twigs may grow 6 to 12 inches or more per year, with greater growth on young plants and sterile shoots.

Leaves are linear-oblong with a cuneate (wedge-shaped) base and a mucronulate (short-tipped) apex, ranging from tiny to 2+ inches long and ⅜ inch broad. Surfaces are glabrous (without pubescence), shiny dark green above, dull medium green beneath. Margins are entire (without teeth), slightly wavy and revolute (rolled under). Axillary buds of the primary leaves (below flower-bearing portion of twigs) produce clusters of tiny secondary leaves. 

Photo 3: New twigs grow from previous year’s twig (on right). Note primary and secondary (axillary) leaves. New twigs have two slight ridges between primary leaf nodes.

Showy, yellow flowers bloom in early summer. They occur singly and in groups of three at and near the stem tips, typically a terminal flower and a pair of lateral, axillary flowers. Individual flowers (to 1 inch wide and ¾ inch high) remain at anthesis for one day, but with many clusters, shrubs may bear flowers for three weeks.

Photo 4: Flowers follow a pattern-of-three. As shown, the apical cluster has a terminal flower and two lateral clusters of three flowers. Note leaves subtending calyxes. (Larger leaves removed for photo.)

Smooth shiny flower buds mature from a pale green to yellow. With anthesis, five bright yellow petals (to ½ inch long and ¼ inch wide) reflex back onto the smaller sepals to expose a large, circular mass of closely spaced, golden yellow stamens. Straight, slender yellow filaments are tipped with round, golden yellow anthers. The androecium (stamens as a unit) encircles a prominent cone-like pistil, the ovary of which tapers to three stigmas formed by three united styles. As flowers fade, petals become golden brown before dropping off.

Photo 5: A dense androecium surrounds a conical pistil. Ovary consists of three compartments terminated by three united styles. Note leaves subtending calyxes.
Photo 6: Display shows petals (1), a small number of stamens (2), pistil (3), leaves that subtend calyx (4), and calyx with stamen stubs (5).

As pistils mature in late fall, their apexes and sides split and the ovaries become rusty brown capsules with three sharp points (styles). Black, flattened, oblong seeds are released as the flower-bearing portion of twigs dies.

In a garden or natural setting, shrubby St. John’s-wort would be an asset either as a specimen plant or in a loose group. As it tends to be a prolific seeder (perhaps another possible source of the its specific epithet?), seedlings may need to be removed; on the other hand, seedlings do provide replacements for short-lived shrubs. Seedlings may produce flowers by the third year. Bees (especially native bumblebees) and other insects are attracted to flowers for their copious supply of pollen; flowers produce no nectar. Not favored by deer.

Including the marsh St. John’s-worts, sometimes treated in the separate genus Triadenum, nearly twenty species of Hypericum occur in Arkansas, most with yellow flowers. Shrubby St. John’s-wort can be distinguished by a combination of its woody habit, large (as compared to the other Arkansas species) five-petaled flowers, densely packed androecium, and rounded-in-cross-section capsule. The species most likely to be confused with shrubby St. John’s-wort is five-lobe St. John’s-wort (Hypericum lobocarpum). Also a woody shrub, five-lobe St. John’s-wort has slightly smaller, more densely arranged flowers and capsules that are five-lobed in cross-section (hence both its common and scientific names).

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Royal Fern

Royal fern (Osmunda regalis var. spectabilis*) of the Royal Fern (Osmundaceae) family is a large herbaceous fern with a crown of clustered spore cases (sporangia) on the fertile fronds (leaves). In the U.S., it occurs from eastern Texas to Minnesota and thence to the Atlantic and Gulf Coasts. In Arkansas, it occurs pretty much statewide. The genus name, according to some writers, originates from “Osmunder,” a Saxon name for Thor, the god of thunder. The specific epithet is from the Latin for “regal” or “royal,” in reference to the fern’s size and majestic appearance. The variety name is from Latin for “admirable” and “spectacular”. It is also known as regal fern and American royal fern (this being the only variety in North America). Habitat preference is partly to mostly sunny areas with consistently wet to occasionally flooded soils, such as shallow, slow-moving drainages, bogs, marshes, seeps, moist bluffs, and open wetlands.

Royal fern has a stout, elongate rootstock that may be oriented vertically (in less-wet habitats) or horizontally (in wetter habitats). Dense, black, wiry and tough fibrous roots completely shroud the rootstock. New fronds (leaves) emerge at rootstock’s apex, surrounded by stubs of previous years’ leaves. Apices of vertically positioned rootstocks may become elevated above duff level while apices of horizontally oriented rootstocks remain at duff level, in rhizome-like fashion.

Photo 1: This 2½ foot-tall plant has matted fibrous roots which enshroud a horizontally oriented rootstock that is 9½ inches long and 1¾ inches wide.

In early spring, new fronds appear as large fiddleheads which unfurl into smaller fiddleheads that unfurl into pinnae (leaflets). The large fiddleheads are initially covered with long, matted pubescence which drops off as the stipe (leaf stalk) and rachis (midrib) elongate. Unfurled leaves are bipinnate or twice-cut into pinnae (leaflets) and pinnules (secondary leaflets). Depending on the habitat and season, fronds may be erect, ascending or arching. Fronds gradually decline in the fall and, except for frond stubs, disappear over winter.

Photo 2: In early April, newly emergent fiddleheads are temporarily covered with matted pubescence.
Photo 3: Fiddleheads remain mostly closed as stipes elongate to several feet tall. Fern at lower right is lady fern (Athyrium filix-femina).

Fronds are typically 2 to 4 feet long (maximum about 6 feet) and about half as wide, with widely spaced, opposite pinnae that elongate to 12 inches. Fronds are sterile or fertile, both having the same vegetative structure. However the upper pinnae and pinnules of fertile fronds lack chlorophyll and are modified into clusters of spore-producing sporangia. Below the sporangia, fertile fronds have the same appearance as sterile fronds. 

Photo 4: Sporangia of plants on left are green while those of plants on right have become brown after release of spores. Lady fern (Athyrium filix-femina) at lower right and Christmas fern (Polystichum acrostichoides) at lower left.

Pinnae of both sterile and fertile fronds bear six to twelve pinnules to 2 inches long and ⅜ inch wide. Pinnules are alternate to offset-opposite, mostly oblong, with rounded apexes and rounded to truncate-oblique bases. Margins are slightly undulating and minutely serrate. Fertile pinnae are greatly reduced in size. Spherical sporangia, maturing from frond apex downward, occur in small, naked, short-stalked clusters. Initially dark green, due to the chlorophyll-bearing spores within, sporangia turn lighter green to golden brown as they mature and then split in half across the top to release the spores. Sporangia then wither, and in late spring, the entire upper spore-bearing portion of the fertile fronds quickly shrivels. The lower portion of fertile fronds remains green and photosynthetic.

Photo 5: An 18-inch long section of a 4-foot long fertile frond. Fertile (sporangium-bearing) pinnae are significantly smaller than sterile, photosynthetic pinnae.
Photo 6: Clusters of sporangia replace leafy pinnules at tips of fertile fronds. Sporangia split across their tops for spore dispersal.

With spores dispersed, the reproductive activity of the “sporophyte phase” of the fern’s life cycle concludes. In the soil, spores germinate to produce a prothallus, the “gametophyte phase.” The tiny, ephemeral prothallus produces not spores but gametes, sperm and egg. Mobile sperm swim through ground moisture to fertilize eggs that have remained attached to their prothallus. Fertilization then produces a zygote that in turn develops into a large, perennial sporophyte plant. Readers who have taken a general botany course may remember this complex, elaborate life cycle being described as an “alternation of generations.”

Royal fern would be an excellent choice for mesic (in shade) to hydric (in sun) areas of a native plant or fern garden. Like most temperate zone ferns, it is herbaceous, dying to the ground each winter. But from spring to fall, with its large, bold structure, yet elegant and graceful fronds, it is a striking addition to the garden, either as a single plant or as a colony.

Photo 7: In fall, fronds recline and become more yellowish. Photo–October 23.

In addition to royal fern, two other members of the Osmundaceae family occur in Arkansas: interrupted fern (Osmunda claytoniana) and cinnamon fern (Osmundastrum cinnamomeum). While royal fern has bipinnate leaves (pinnae fully divided into pinnules), the other two ferns have pinnate-pinnatifid leaves (pinnae deeply lobed but not fully divided). Also, sporangia of interrupted fern, unlike those of royal fern, are located on fertile pinnae below the apex of fertile fronds (about mid-frond, with sterile pinnae above and below). Cinnamon fern has entirely separate fertile fronds that bear sporangia only.

*Recent genetic studies suggest that the royal fern of North America is a separate species (Osmunda spectabilis), but this reclassification has not been fully accepted by the botanical community.

Article and photographs by ANPS member Sid Vogelpohl

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