Know Your Natives – Self-Heal

Self-heal (Prunella vulgaris ssp. lanceolata*) of the Mint (Lamiaceae) family is a perennial herb found in generally moist soils.  It occurs as a native species throughout the lower 48 states and Alaska.  In Arkansas, it occurs throughout the state.  The origin of the genus name is not known.  The specific epithet is from Latin for “common” as in “widespread”.  The subspecies name is from Latin, meaning “lance-like”, a reference to the leaf shape.  Other common names include lance-leaf self-heal and heal-all.  Self-heal grows well in various moist soils with full or partial sun in prairies, woodland borders and roadsides.  It was widely used by Native Americans and early settlers for medicinal purposes and it is still used as a medicinal herb.

Self Heal - Prunella vulgarisPhoto 1:  A mature plant prior to stem development.  Leaf blades, marked with glandular dots, have slight serrations on their margins.

Self-heal has shallow, light tan, long fibrous roots that radiate from a small central caudex, and the plant may also produce short roots from the lowermost portion of the stems that are in contact with soil.  Plants may have one to a half-dozen mostly erect stems with a final height from 6 to 12 inches or more.  Secondary stems may grow from upper leaf axils of main stems.  Stems are square in cross-section (a characteristic of mints) with sharp corners.  Width of stems, at about 1/16 inch, is fairly uniform from stem base to uppermost leaves.  Stems are a light green, but the lower portion may be reddish.

Self Heal - Prunella vulgarisPhoto 2:  In mid-May, this plant in a sunny site has 20 stems with minimal secondary stem growth.

Basal and stem (cauline) leaves have a similar appearance although basal leaves tend to have longer petioles in relation to leaf blade and may have more prominent glandular dots.  Cauline leaves, opposite and widely spaced, are petiolate except for the uppermost leaf pair, which have variably shaped “petioles” (see below).  Blades, broadest below mid-leaf, are broadly lanceolate to ovate with an obtuse tip and rounded to cuneate base.

Leaves of self-heal, medium green in color, may be 2 inches long and ¾ inch wide, or larger.  Margins may be entire (smooth) or have widely spaced hardly noticeable serrations and may be undulating.  Venation, recessed above and prominent below, is pinnate with a half-dozen or so secondary veins that are offset across the primary vein (midvein).  These secondary veins, extending from the primary vein, stop short of the leaf margin and fade out as they arch toward the leaf apex.  The recessed appearance of the primary vein continues down the petiole as a central groove.  Petioles may have thin wings that extend down from the leaf blade, with the wings more evident nearer the blade.

The uppermost pair of cauline leaves generally occurs immediately below the inflorescence, but a short stem segment may occur between the uppermost leaf pair and the inflorescence.  In the case of secondary stems, the uppermost pair of leaves are typically the only leaves on the stem, but occasionally another pair of leaves may be present.  Appearance of the uppermost leaf pair varies from plant to plant regarding shape of blade and petiole.  The blade outline may be similar to other cauline leaves or more linear and may be on a very short petiole or  sessile.  Alternatively, the “petiole” may be thin and widened so that its appearance is similar to that of the floral bracts (see below), with the blade size reduced and strongly linear.

The inflorescence is a rounded, cylindrical cluster at the tip of the stem, typically immediately above the final leaf pair.  Clusters have a loose, spongy feel.  Clusters, at the time of flowering, are about 1 inch long, but may be 2 inches long when flowering is completed.  Flowering occurs in late spring and lasts for about a month.  Flowers may bloom sequentially in tiers from lowermost to uppermost or flowers at anthesis may be scattered throughout the cluster.

Inflorescence clusters are composed of leafy floral bracts and flowers that are attached to a columnar peduncle.  Floral bracts occur in opposite pairs. Like the leaves, each pair is rotated 90 degrees from the pairs directly above and below it (decussate).  These cordate (heart-shaped) bracts may be up to ½ inch long and wide; they are sessile, ciliate (hairs on margins), tissue-thin, and light green with a darker green or purple rim.  Bract size decreases from the inflorescence base to the apex.  A whorl of six flowers occurs immediately above the floral bracts with fewer than six flowers per whorl near the top of the cluster.  Above the final whorl, several pairs of decreasingly-sized bracts create a flattened top.   When floral bracts have especially long pointed apices, the typically round cluster appears square.

Self Heal - Prunella vulgarisPhoto 3:  Display showing upper and lower leaf surfaces along with a stem terminating with a flower cluster.  Leaves, stems and calyxes are pubescent.

Self Heal - Prunella vulgarisPhoto 4:  Display of an expanded cluster alongside its stalk and axis.  Cluster had five whorls of flowers, each whorl sited above a pair of floral bracts.  Numbers with bracts correspond to numbers along stripped axis to show point of origin.

The tubular corollas are first seen as dark-colored nubs that appear from within calyxes.  A flower, about ¾ inch long from base of calyx to end of corolla, has an upper lobe and three smaller lower lobes.  Pedicels are very short.  The exterior of the upper lobe is typically purple (as first shown by nubs), while the lower lobes and the interior of the flower are a lighter color.  The four lobes join within the calyx to form a constricted floral tube.  The upper lobe, with a notched central ciliated ridge, is hump-backed above the notch toward the end of the lobe so that a hood (galea) is formed.  Rounded margins of upper lobes are flared upward.  Two oblong lateral lower lobes are projected forward while the larger middle lower lobe, with a broad fringed margin, bends down.

Flowers have four stamens (forked at top) that are adnate (fused) to the floral tube and a bifurcated style attached at the depression in the center of the deeply 4-lobed ovary.  The four stamens are forked near the tip, with a longer pair and a shorter pair, together with the stigma, positioned under the hood of the upper lip of the corolla.

The calyx, comprising an upper and lower lobe of different structure and shape, is open and gaping before corolla nubs appear.  The upper lobe is hump-backed with a flattened upper side and two lateral rounded sides.  The lower lobe, more narrow and planar, is positioned between the sides of the upper lobe.  While the top of the upper lobe is mostly featureless, the sides of the upper lobe and the lower lobe have longitudinal ridges.  The ridges on the lower lobe terminate as two pointed teeth.  The upper lobe has a rounded to truncated, notched (emarginate) apex with three tiny spines.

Self Heal - Prunella vulgarisPhoto 5:  Ciliated ridge of hump-backed upper corolla lobes and fringe on middle lower lobe can be seen.  Top of cluster is closed by floral bracts.

Self Heal - Prunella vulgarisPhoto 6:  Pairs of rounded floral bracts separate whorls of flowers set in spiky calyxes.  Lowermost bract is indicated by an arrow.

Self-heal is typically pubescent.  Stems are densely covered with stiff hairs (hispid) which continue along petioles and, to a lesser degree, onto leaf blades.  The lower leaf surface is more pubescent than the upper surface, especially along veins.  Sides of the upper calyx lobe are hispid while the dorsal surface is glabrous.  The lower calyx lobe is also hispid.

A fertilized ovary produces a brown cluster of four tiny, smooth, brown and egg-shaped nutlets that are retained in an upright position into late summer.  As rain drops strike the fruiting clusters, nutlets can be flipped out of the calyxes for dispersal by flowing water.

Self Heal - Prunella vulgarisPhoto 7:  In this early July photo, fruiting clusters have dried and stems are deteriorating.

Self-heal is a “pleasant-looking” small perennial that may work well in a moist sunny garden.  It has attractive leaves, flowers and flower clusters–good plant for bees and butterflies.  This native mint does not seem to spread aggressively by seed although it does spread by rhizomes.  If reseeding is an issue, bloomed-out flower clusters can be easily removed.  Flower clusters may be eaten by deer which would encourage an extended bloom period via lateral branching.  Being herbaceous (non-woody), the above-ground portion of this perennial plant disappears in winter.

  • The non-native self-heal (Prunella vulgaris ssp. vulgaris) is reported from scattered areas across the U.S., but is not currently known to occur in Arkansas.  Whereas stems of Prunella vulgaris ssp. lanceolata are mostly erect and with mid-stem leaves that are three times as long as wide, stems of Prunella vulgaris ssp. vulgaris, a shorter species, generally recline on the ground, with roots at leaf nodes and mid-stem leaves twice as long as wide.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – New Jersey Tea

New Jersey tea (Ceanothus americanus) of the Buckthorn (Rhamnaceae) Family is a small, deciduous, thorn-free shrub.  The genus name originates from a Greek word for “spiny plant” or a Latin word for “thistle”.  The specific epithet relates to its occurrence in the Americas.  In the U.S., it is found from Texas to Nebraska to Minnesota thence east and south to the coasts.  In Arkansas, the shrub occurs statewide.  Another common name is wild snowball.  Habitat preference is variable, but the species is typically found in dry to well-drained soils of sandy, sunny prairies or rocky slopes and open woodlands.  Leaves were used as a tea substitute during the American Revolution.

New Jersey Tea - Ceanothus americanusPhoto 1:  In mid-May, this young plant exhibits rapid apical stem growth that will not produce inflorescences in the current year.

New Jersey tea, maturing with a dense rounded shape at 2 to 3 feet tall, has erect to ascending stems and branches.  Plant height is limited because apical buds of mature plants do not develop and upper portions of stems and branches die over winter.  In spring, multiple new branches grow from the previous year’s leaf axil buds on surviving portions of stems and branches.  Also, new stems may arise from near ground level, which produce inflorescences the first year.  Current year’s stems and branches, slender and pubescent with short fine hairs, become rigid and woody during the year.

New Jersey Tea - Ceanothus americanusPhoto 2:  Also in mid-May, this mature plant approaches maximum bloom.

Current year’s stems and branches are light yellowish-green, with similar color retained on those that survive for a second year, although slightly darker and without pubescence.  Older surviving stems and branches become trunk-like and brown and roughened.  Stems, branches and trunks are round (terete).  With a stout, deep taproot and excellent regrowth abilities, New Jersey tea is resistant to drought, fire and deer.  In conjunction with bacteria, New Jersey tea “fixes” nitrogen in the soil much like many legumes.

Buds occur at all leaf axils of current year’s stems and branches.  The uppermost two to eight axillary buds produce floral branches (see below) in the current year while lower axillary buds remain dormant to produce branches with inflorescences the next year.

Alternate leaves are irregularly spaced from about ½ to 2 inches along current year’s stems and branches.  Leaf size, too, is irregular, but generally, the largest leaves are in the mid-section of stems and branches.  Largest leaves are about 2¼ inches long and 1 inch wide with ¼ inch petioles that may be slightly reddish.  Leaves are elliptic, with a rounded base and broadly acute tip, and they appear slightly rough.  Upper and lower leaf surfaces have short pubescence so that leaves, when carefully handled, feel like flannel.  Upper leaf surface is a medium green while lower surface is more yellowish-green and dull, with prominent pale yellow veins.  Veins are slightly recessed above and expressed below.  Uniformly sized and shaped serrate margins extend from the lower one-fourth of the leaf blade to the apex.

Leaves have unusual and distinctive venation.  Three primary veins originate at the petiole, with the two arcuate (curved) laterals terminating at the leaf margin about a fifth of the leaf-length from the apex.  All secondary veins are arcuate.

Inflorescences occur as multiple elongate globular clusters of tiny flowers at the ends of floral branches that grow from upper leaf axils of current year’s stems.  The floral branches, which may be 2½ inches long and tend to be twisty, are leafless or occasionally bear one or two small (to ¼ inch long) leaves.  Floral branches and peduncles have the same color and pubescence as current year’s stems and branches.

Flower clusters, which may be 1 inch or more long and ½ inch or more in diameter, consist of many tiny white flowers in small panicles around a central axis.  Within the panicles, multiple flowers grow from common points.  Clusters, which may be held upright or oriented in various directions, can have 200 flowers.  Flowers in each cluster bloom at about the same time, with the clusters originating from lower on the stem or branch blooming first.  Flowers are slightly fragrant.

New Jersey Tea - Ceanothus americanusPhoto 3:  Display of a flowering branch separated into sections.  Note unusual leaf venation and long twisty, leafless peduncles.  Lower side of a leaf is shown at left-center.

Flower buds nearing anthesis have five rounded-triangular sepals that tightly cover five enclosed petals that cover five large anthers (one anther per stamen).  Buds, when viewed from above, are star-shaped with five rounded “knobs” created by the enclosed anthers.  With anthesis, sepals separate slightly, but remain in the same position as in the bud.  The five fragile-looking petals, with broad cupped apices and long narrow bases (ladle shape), extend through the opening between sepals and flare wide.  The large yellow pollen-bearing anthers are at first within the bowl of the ladle, but later, anthers become dark (pollen having been discharged) and filaments project anthers above the corolla.  Flowers, 3/16-inch-wide from petal tip to petal tip, are positioned in a flat plane well above slender white pedicels that are as long as the flowers are wide.

New Jersey Tea - Ceanothus americanusPhoto 4: Flower buds are knobby due to size of enclosed anthers.  Several flowers are shown at anthesis such as the one at lower-center.  Note long white pedicels.

New Jersey Tea - Ceanothus americanusPhoto 5: Inset of a single flower shows pollen-bearing anthers shifting out of bowl of ladle. Several more mature flowers within the panicle, also shown, have dark pollen-free exserted anthers.

Flowers, with fertilization, produce triangular, 3-chambered, rough, light-green capsules about ¼ inch wide on green stems (previously white pedicels).  Capsules become black at maturity and the upper portion splits to release a single smooth, brown, ovoid seed from each chamber.

New Jersey Tea - Ceanothus americanusPhoto 6:  In this mid-July photo, some of the triangular seed capsules have matured (black ones) and split to release seeds.  Previously white pedicels are now green.

A second species in the genus occurs in Arkansas; namely, “inland New Jersey tea”, also called “redroot”, (Ceanothus herbaceus), which is known from a number of western and central counties.  Although also a woody shrub, it was given the epithet “herbaceus” when it was described because it was mistakenly thought to be an herbaceous perennial.  This species has narrower, glabrous leaves that are tapered at both ends.  Smaller and more rounded or dome-shaped flower clusters occur terminally from the uppermost leaf axil of each stem.  Inland New Jersey tea generally blooms a few weeks earlier than New Jersey tea.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Western Daisy

Western Daisy (Astranthium ciliatum) (formerly Astranthium integrifolium*), of the Aster (Asteraceae) family, is an annual species with daisy-like flower heads. In the U.S., it is found in Arkansas, Oklahoma, Texas, southern Nebraska and southwestern Missouri with greatest concentrations in eastern Oklahoma and western Arkansas. The genus Astranthium comprises about a dozen species from the southern U.S. and Mexico, with only a single species in Arkansas, reported from the Ozarks, the Arkansas River Valley and the Ouachitas. Another common name is Comanche western daisy. The genus name is from the Greek for “star” referring to the star-like flower heads as seen from above. The specific epithet, also of Greek origin, means “fringed with hairs” in reference to the plant’s pubescence. Western daisy is found primarily in full or partial sun in sandy, clay or loam soils of grasslands, glades, and open deciduous woods mostly in lowlands. Plants have short, tough tap roots or multiple, radiating, near-surface roots up to 2 inches long.

Western daisy is a small plant that germinates in the fall into winter to form a flat-lying rosette of basal leaves, up to several inches across. Basal leaves, which have entire (smooth) margins, have an oval shape with a gradually narrowed base (spatulate) and long hairs on the upper surface. In late winter, ascending leaves, attached to a rudimentary main stem, appear from the center of the basal leaves as the basal leaves begin to fade.

Western Daisy - Astranthum ciliatumPhoto 1: In mid-March, new leaves appear as the winter leaves fade. Note pubescence.

Stems, which are light green, are slender, with three to four slight ridges that connect with petiole-stem junctions. Faint reddish shading may be present on stems, especially on ridges. The single main stem typically has secondary stems that arise from lower leaf axils at about 35 degrees off the main stem.

In less desirable sites (excessive shading, crowding, etc), plants may remain small with no or few secondary stems. In more desirable sites, secondary stems become dominant and, in turn, may bear stems that arise from their upper leaf axils. Secondary stems, which are fairly straight, are erect on smaller plants and spreading to ascending on larger plants. Stems are covered by soft hairs.

Western Daisy - Astranthum ciliatumPhoto 2: In less desirable sites, plants remain small with limited secondary stem growth. White flowers shown with the western daisies are of long-flower cornsalad (Valerianella longiflora).

Alternate stem (cauline) leaves have the same light green color on upper and lower sides as the stems; however, lower sides are slightly shinny. Blades of lower cauline leaves, to 1¼ inch long and ½ inch wide, have an elongate oval shape that is entire and widest at mid-blade, becoming gradually more narrow toward their sessile bases (oblanceolate). Blades of upper cauline leaves become increasingly smaller and more narrow up-stem, with more pointed apices (lanceolate) and while remaining sessile. Lowermost leaves have relatively long pubescence on upper blade surfaces and margins (ciliate) and have glabrous lower surfaces. Spacing of cauline leaves varies with the greatest spacing being at the lowermost portions of stems where spacing may be up to 1¾ inches while other leaves may be spaced at ¼ inch. Leaf-stem junctions are at about 35 degrees from which point lower leaves recurve downward while upper leaves extend mostly outward.

The primary inflorescence of western daisy, at apices of stems, consists of single composite flower heads on long peduncles. Peduncles, measured from the base of the flower head to the uppermost cauline leaf, vary from short (1 inch or less) to long (to 3 inches). Peduncles have the same appearance as stems. As peduncles grow and strengthen, flower head buds are at first upright, then droop and then become upright again at anthesis. A plant’s first flower head to reach anthesis, regardless of plant size, is that single head at the apex of the main stem.

Western Daisy - Astranthum ciliatumPhoto 3: In more desirable sites, as shown by this single plant, secondary stems become dominant. Flower head at apex of main stem can be seen mostly hidden at center of plant. Note drooping flower head buds.

Western Daisy - Astranthum ciliatumPhoto 4: Display of flower heads from buds (lower left) to past-anthesis (lower center).

Flower heads, up to about an inch wide, have up to about 20 pistillate ray florets and numerous bisexual disk florets (radiate flower heads). Flower heads have a slightly domed center and rounded (hemispheric) involucres. Strap-like ligules of the ray florets, about ¼ inch long, have rounded to notched apices and constricted bases. Ligules typically overlap, but may also be spaced slightly apart. The constricted bases of the ligules are not especially visible except when ligules are spaced apart. Ligules are typically light lavender, but may be white. Lavender ligules may or may not have white coloration near their bases, with the color change being gradual or sharp. Tiny disk florets, yellow with yellow pollen, have five triangular spreading lobes on the rim of a short corolla tube set on a green ovary. Exserted anthers, cohering into a short tube, make pollen available to insects when the style, like a plunger, pushes through their tube, depositing the pollen on the surface of the disk. Involucres, composed of about 20 thin, lanceolate, green, equal-length and appressed phyllaries, are slightly overlapped along their translucent edges.

Western Daisy - Astranthum ciliatumPhoto 5: Disk florets have five lobes on their corollas and anthers fused into a tube, as shown by the outer ring of florets. Ligules have constricted bases and rounded to notched apices. The disks of composite heads typically flower from the outside in toward the center. Here the head is just getting started. Notice how immature the central disk floret buds are compared to those near the perimeter.

Western Daisy - Astranthum ciliatumPhoto 6: Involucre composed of lanceolate, slightly overlapping phyllaries. Peduncle, slightly ridged and pubescent, has same appearance as stems. Spiders and pollinating insects often have encounters on flower heads.

With fertilization, each floret produces a single one-seeded achene of two fused carpels (the two halves of a typical sunflower seed wall), surmounted by a ring of hairs at the top (pappus). With fruit (achene) dispersal, a conic, pitted receptacle remains.

  • Western daisy (Astranthium ciliatum) has been assigned to various species and subspecies over time. The most recent reassessment defines western daisy (Astranthium ciliatum) as being found west of the Mississippi River and eastern daisy (Astranthium integrifolium) as being found east of the Mississippi River.

Footnote: There are several other species of composites in Arkansas, representing various genera, that may also have “daisy” in their common names.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Hairy Phacelia

Hairy phacelia (Phacelia hirsuta) of the Borage (Boraginaceae) family [formerly of the Waterleaf (Hydrophyllaceae) family] is a beautiful annual forb with blue flowers.  In the U.S., it is found naturally in Arkansas, Louisiana, Texas, Oklahoma, Kansas, and Missouri, as well as introduced in Kentucky and Pennsylvania.   In Arkansas, it is found throughout much of the state but sparser in the Mississippi Alluvial Plain.  Its habitats include rocky to sandy, moist soils in open woods and woodland margins, as well as in prairies, glades, and along roadsides.  The genus name is from Greek for “bundle”.  The specific epithet is from Latin for “hairy”.  Another common name is “hairy scorpion-weed”.

Hairy phacelia, whose basal leaves appear in late winter, develops a main stem and several long to short secondary stems.  The longest secondary stems grow from lower leaf axils of the main stem, with shorter stems growing from mid-stem axils.  Plants have a tap root and a few fibrous roots.  The main stem and secondary stems reach a similar maximum height of up to 1 to 1½ feet.  Stems, which are weakly erect, are yellowish-green with the lower portion having a reddish cast.  Stems are very hairy (hirsute).

Hairy Phacelia - Phacelia hirsutaPhoto 1:  In this April 2nd photo, several secondary stems have developed and developing inflorescences can be seen at the ends of stems.

Alternate hirsute leaves, with three to seven leaflets or lobes, are widely spaced on a mature plant and are medium green on upper surfaces and light green on lower surfaces.  Leaves feel soft and venation is not especially distinct.

Lower leaves, which have long, grooved petioles, have blades from ¾ inch to 1¼ inch long, with maximum width of about ¾ inch.  These leaves typically have a lower lateral pair of opposite, oval leaflets and, higher up the leaf blade, one to three pairs of mostly opposite, lateral leaflets that may have widened bases, becoming lobes.  Additionally, a large, broad terminal lobe is partially cleft on both sides so that it is three-lobed.  Lower leaflets (or lobes) are set perpendicular to the rachis while upper lobes (including those of the terminal lobe) are angled toward the apex.   Lobes of these lower leaves may be wide or narrow with rounded tips.

Upper leaves (from mid-stem into the inflorescence), have blades that decrease in size up-stem to uppermost leaves that may be ¾ inch long and 3/8 inch wide.  Transitioning up-stem,  lobed leaves gradually change shape from petiolate to sessile and lobe blade tissue increasingly extends along the rachis.  The uppermost leaves have two or three pairs of lateral lobes that may be either narrow-rounded and angled toward the leaf apex or narrow-acuminate and out-flared.   The shape of terminal lobes is similar to lateral lobes.  Petioles of mid-stem leaves (when present) are wide and broadly grooved.  Smaller leaves are sessile.

Hairy Phacelia - Phacelia hirsutaPhoto 2:  Leaf display shows changing leaf shape from basal to upper stem leaves (left to right).  Note that leaves change from petiolate to sessile and from having some leaflets and lobes to having all lobes.

Inflorescences, in mid-spring, grow from uppermost leaf axils of main and secondary stems.  Appearing first as round clusters (or “bundles”) of compacted sepals, with growth, the coiled nature of the inflorescence becomes apparent.  Coils consist of a peduncle with up to twenty or more flowers alternately arranged along the upper side of the inflorescence axis (rachis).*  Flowers are each on a short pedicel.  Peduncles and pedicels have short hirsute pubescence.  As the coil straightens, flowers reach anthesis from lowermost to uppermost.  When fully straightened and the last flower has bloomed, the inflorescence may be 3 inches long.   The entire plant continues to stretch out (grow) until the final flowers have bloomed.

Hairy Phacelia - Phacelia hirsutaPhoto 3:  Developing inflorescence appears as rounded clusters of loose, hirsute sepals at ends of stems.

Flowers, about ½ inch in diameter, have a deeply divided calyx with five long, narrow, spreading lobes with hirsute exteriors, and a bowl-shaped corolla notched to create five broadly-triangular but rounded lobes.  The exterior of the corolla is pubescent.  Corollas are typically a light blue to lavender overall, but may have a lighter or white center.  Just below mid-corolla, ten dark purple, round to three-sided spots encircle the flower’s center, two spots below each lobe.  These purple spots are surrounded by a haze of lighter color.  Five spreading stamens have white filaments, with anthers at first bearing light yellow pollen that become black.  Filaments, adnate at the base of the corolla, are covered by long radiating white hairs.  The pistil, also white, has a forked and pointed style whose stigma does not become receptive to pollen until after the pollen from the same flower has been shed (protandry–an adaptation that reduces self-pollination).   Stamens and pistils, all about the same length, extend slightly beyond the rim of the corolla.  A light green, superior ovary, also hirsute, has the shape of an elongated, round cone.

After producing seed capsules containing a small number of brown seeds, these annual plants quickly die.

Hairy Phacelia - Phacelia hirsutaPhoto 4:  Flowers reach anthesis at the top of a coil in sequence (from left [bottom of coil] to right [top of coil] in this photo) as coil straightens.  “Newest” flower at right bears pollen.  Forked styles can be clearly seen in the lower two flowers.

Hairy Phacelia - Phacelia hirsutaPhoto 5:  In this May 1st photo, plant is nearing the end of its life cycle.  Plant being collected by botanist Eric Sundell (a reviewer of these articles), accompanied by Milanne Sundell.

In addition to hairy phacelia, six other native phacelia species occur in Arkansas.  Hairy phacelia is the most common and widespread.  Three of the others are rare to very rare in the state and are tracked by the Arkansas Natural Heritage Commission.  Hairy phacelia can be distinguished from the other six by several characteristics, including 1) high degree of pubescence and 2) light blue to lavender, bowl-shaped, 10-spotted corollas, 3) stamens and pistils that are only slightly exserted from corolla, and 4) rounded, non-fringed corolla lobes.

Another species in the borage family that can occur in the same habitats as hairy phacelia and with some similar characteristics is “large-flower baby-blue-eyes” (Nemophila phacelioides).  It occurs in the west-central part of the state.

  • This inflorescence style is referred to as a scorpioid (resembling the coil of a scorpion’s tail) or helicoid cyme which is typical of the borage family and the source of one of the general common names, “scorpion-weed”, for some members of the genus Phacelia.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Wild Geranium

Wild geranium (Geranium maculatum) of the Geranium (Geraniaceae) family is an herbaceous woodland perennial that blooms in early spring.  In the U.S., this species is found from Louisiana and Oklahoma to Florida and north to North Dakota and Maine.  In Arkansas, it is found primarily in the highlands of the northwestern half of the state.  The genus name is based on a Greek word meaning “crane” in reference to the shape of the plant’s fruit which resembles a crane’s bill.  The specific epithet means “spotted” in reference to leaf spots that may or may not be present.  Other common names include spotted geranium, spotted crane’s-bill and wood geranium.   Preferred habitats are wind-protected, lightly shaded, mesic deciduous woodlands and borders with various well-drained soils, but it may be found in sunny sites with consistent moisture.  Plants may wilt on hot, windy spring days and may enter early dormancy if drier conditions persist.

Wild geranium, a long-lived plant, has stout, dark-colored, round, shallow, rough rhizomes with a light orange interior.  Nubs develop along the sides of rhizomes that may grow into rhizome branches.  Fibrous roots grow from the lower sides of rhizomes.  Leaf and stem growth originates at the tips of rhizomes.  With mature plants having many growth tips on relatively short rhizome segments, a mature plant develops a rounded mound of leaves.  A plant may have a spread of 2 feet or more and reach a height of 1½ feet tall.

Wild Geranium - Geranium maculatumPhoto 1:  A partial rhizome and plant (detached from main rootstock at light orange area).  This rhizome segment has several branches, including a small branch on the right side.

The first leaves to appear in spring are basal leaves.  The largest basal leaves, on the periphery of the leaf mound, may be 6 or more inches wide with a petiole that is 7 or more inches long.  Leaves, medium green on upper surface with a similar dull green lower surface, are finely pubescent on the upper surfaces as well as along the veins of lower surfaces.  Basal leaves have a terminal lobe and two lateral lobes on each side, arranged in palmate fashion.  The five lobes have wedge-shaped bases with wider mid-sections which gradually narrow in width to acutely rounded apices.  Margins, which are entire near lobe bases, have short to long crenulations from mid-leaf to the leaf apex.  The five lobes join together shortly above the petiole.  The petiole is attached to the underside of the leaf blade at the leaf margin.  Individual leaves and petioles are weak, but remain upright due to support from nearby leaves.  The interior of the leaf mound is filled with smaller leaves.  Basal leaves remain into late fall.

Wild Geranium - Geranium maculatumPhoto 2:  A relatively small young plant.  Note wide uncleft area at center of leaves as well as flower buds making their appearance (see top center and lower right).  White flowers in upper left are those of rue anemone (Thalictrum thalictroides).

Shortly after the leaf mound has formed in late winter to early spring, clusters of football-shaped flower buds become visible across the leaf mound, as stout, unbranched and heavily pubescent floral stems rise above the basal leaves.  Stems, five sided (more pronounced on larger stems) with downward-trending prominent pubescence and one or more pairs of cauline (stem) leaves, grow from rhizome tips surrounded by basal leaves.  Stems continue to grow until buds are well above basal leaves and the lowermost pair of cauline (stem) leaves is positioned just above the basal leaf mound.  As flowers reach anthesis, stems (with inflorescence) may have a total height of 16 inches with 10 of those inches below the lowermost cauline leaf pair.  The lowermost pair of cauline leaves subtends two or three erect, light green to reddish secondary stems that may be 5 inches long.  These secondary stems, in turn, support one to three peduncles (stalks of inflorescences) from which several pedicels (stalks of flowers) grow that are also subtended by leaf pairs.  Leaves of the lowermost cauline leaf pair have the same appearance and size as basal leaves, but with petioles that are from ¾ inch to 4 inches long.  Leaves that subtend pedicels have the same texture as other leaves, but appear three lobed at about 2 inches wide, with short petioles or perhaps sessile.

Wild Geranium - Geranium maculatumPhoto 3:  A single pair of cauline leaves subtends an upper secondary stem and two lower peduncles.  Note the small leaves on the secondary stem and bracts at bases of stems, peduncles and pedicels.

At anthesis in early spring, 1½-inch-diameter, upward-facing flowers are light pink to lavender with whitish centers.  Flowers have five obovate overlapping petals and five light green pubescent, slightly ridged and elongate sepals.  Sepals have acute apices that terminate in needle-like tips (cuspidate).  Petals, with broadly rounded upper margins, surround ten stamens and a pistil.  Stamens, connecting to the flower’s axis below a hairy ovary (superior ovary) of five carpels, have white slender filaments with elongate yellow anthers that bear yellow pollen.  Pistils have a long slender whitish style that branches into stigmas, spread wide and curved backwards.  Styles attach to the smooth, round-elongate, green sections of the ovary.  Individual flowers, up to about ten to fifteen per stem in pairs or in loose clusters, are in bloom for one to two days.  Total bloom time for a plant extends over about three weeks.  Cauline leaves wither in late summer, well after seeds have dispersed.

Wild Geranium - Geranium maculatumPhoto 4:  Flowers, changing from pink to lavender, have widely flared, overlapping, obovate petals.

Wild Geranium - Geranium maculatumPhoto 5:  Display showing flower buds (lower right), front and back of flowers and a flower past bloom (lower left).  Note five-branched, back-curved stigmas.

After fertilization of a flower, the lower portion of the style quickly elongates and the five seeds–one in each ovary chamber–enlarge so that the ovary becomes knobby. The stigmas wither so that a crane’s-bill-like fruit is formed.  The fruit, an inch or more long, matures and dries to a dull dark brown rigid capsule.  Five dark brown oval seeds, resting just above the drying sepals, are tightly held in place, capped by the ovary wall, which bears five long thin arms (springs) aligned along the style and firmly attached just below the dried stigma.  When the seed-bearing structure is dry, each cap and associated spring-loaded arm snaps upward.  With this instantaneous snap, the seed is catapulted a number of feet away from the plant.   Caps and arms, now curled backwards, remain attached at their upper end to a dried woody style.

Wild Geranium - Geranium maculatumPhoto 6:  The flower at lower position in display (sepals and anthers removed) exhibits a prominent pistil prior to spreading of stigma.  The two fruits (sepals removed) show the closing of the stigma (more mature fruit at top of display), strong growth of their “bills” and enlargement of ovules.

Wild Geranium - Geranium maculatumPhoto 7:  In this display, the central stem bears a green immature fruit and a brown fruit that is poised to launch its five seeds.  Seeds (examples on left) of the other two stems have been launched with caps and arms curled backwards.

For a partially sunny woodland garden, wild geranium is effective as a long-lived plant that is easy to cultivate.  It has attractive form and foliage and showy flowers and fruit so that it is appropriate for specimen plants or as a groundcover.  It is easily propagated by division.
The only other widely occurring native species of the genus found in Arkansas is Carolina crane’s-bill (Geranium carolinianum var. carolinianum).  This is a small weedy annual/biennial species with small palmate leaves and purplish pink flowers.  Another Arkansas native species known from only Miller County is Texas crane’s-bill (Geranium texanum).  Four non-native species of the genus are also found in the state.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Fire Pink

Fire pink (Silene virginica) of the Pink or Carnation (Caryophyllaceae) family is an herbaceous woodland perennial with bright red flowers.  In the U.S., fire pink is found from Florida to New York and westward to northeast Texas, Kansas and Minnesota.  In Arkansas, it occurs across the highlands of the northwestern half of the state.  Its habitat includes partially sunny sites of open woods with various dry to mesic soils where ground vegetation is sparse.

This species occurs as single plants or small colonies of scattered plants.  The base of the plant comprises a woody branched caudex that rests directly on the soil surface, along with a branched tap root, and white fibrous roots growing directly from the caudex.  The caudex of a mature plant has multiple growth points.

Appearance of the basal leaves varies depending on the plant’s age and the season.  New plants have egg-shaped leaves with a constricted base (obovate).  Mature plants have rosettes of basal leaves that are spoon-shaped (spatulate) with narrow elongated bases.  Fresh basal leaves have a dark green upper surface and a lighter green lower surface.  Margins are entire (without lobes or teeth), but may be crinkled.  Although leaf surfaces are glabrous (hairless), hairs occur along the margins (ciliated margins), but only from midleaf (short hairs) to leaf base (long hairs).  Basal leaves that survive over winter assume a reddish color.

Fire Pink - Silene virginicaPhoto 1:  These multiple leaf rosettes are growing from a single caudex.  Flowering stems are poised to grow as can be seem on the left side.  Photo in late February.

Flowering stems grow from centers of leaf rosettes in late winter to early spring.  These main stems, which may grow to a foot or more long, have dense short pubescence.  Initially, stems grow outward, nearly parallel to the ground.  As they mature, stems arch upward so that the plant in full bloom exhibits an open structure with long, slender, wide-spreading to erect stems.  Main stems bear two to six pairs of opposite, clasping leaves (the cauline leaves) with leaves of lowermost pairs being similar to basal leaves, but with wider bases.  Lower leaf pairs are spaced up to about 4 inches apart.  Upper leaf pairs, which become closer together and much smaller toward top of stems, are somewhat ovate with gently acuminate tips.   Lowermost cauline leaves may be five inches long while upper leaves may be only ¼ inch long.  Stems, round in cross-section (terete) and hollow (fistulose), may have a mature length of about 1½ feet.   Stems tend to be reddish on their sunny side and green on shaded side.

Fire pink’s stems have a growth pattern which could be called “pattern-of-three”, that is, leaf pairs at about mid-stem typically subtend a combination of secondary stems and flowers that total “three”.  At about mid-stem, leaf pairs subtend two secondary stems and one flower located between the two stems (referred to herein as “axillary flower”).  When this “pattern-of-three” includes two secondary stems, one of the stems is dominant and it may produce an additional stem.  All stems terminate with a group of three flowers.  On a main stem or secondary stem, the first flower to reach anthesis is the lowermost axillary flower, followed by terminal flowers of secondary stems.  Up to about 30 flowers may occur on one main stem.

Fire Pink - Silene virginicaPhoto 2:  A flowering stem with widely spaced pairs of leaves on its lower portion.  First flowers to open are axillary flowers between two secondary stems, as shown.  Note additional stems just appearing.

Flowers, occurring on separate pedicels, have calyxes composed of five fused sepals that are reddish to purplish on their sunny side while flowers are in bloom, but changing to medium green as flowers fade.  Individual flowers, which may remain showy for a week or more, are up to 1 inch long (calyx included) with a flat-faced corolla that has a width up to 1½ inches.  The corolla comprises five vibrant red petals that are evenly spaced in star-fashion.  Petals, as seen from corolla face, are elongate, each with a characteristic deep terminal notch* and often with short side wings angled toward the tip.  Petals have a sharp flexure where they transition from the corolla face to a tight floral tube formed by the overlapping of narrow petal bases.  Most of the floral tube is within the inch-long calyx tube, but about a fourth of its length is outside.  At the petals’ flexure points, they have two short red upward-extending flanges, so that five petals, as a unit, produce a small corona that encircles the floral tube.  The outside of the petals and exterior of the floral tube are a duller shade of red.  Flowers have ten light red stamens that are adnate to the short stalk of the elongate, yellow-green, cylindrical ovary hidden deep in the calyx.  Stubby, elongate, two-lobed anthers, balanced lengthwise at the tips of filaments, are at first light yellow but become grayish as pollen is released.  Three light red slender styles, arising from the apex of the ovary, have a sloped stigmatic surface.  The calyx tube, with five pointed teeth, is marked with 10 darker longitudinal ridges.  Within the confines of the calyx, bases of petals, stamens and pistils are white.

Stamens and pistils become exserted in sequence: First, the five stamens adnate to the petal bases; second, the five stamens attached in between the petal bases; and third, all three styles.  By the time the second set of stamens becomes exserted, stamens in the first set have lost their anthers and filaments have become back-flared and wilted.  Similarly, by the time the styles become exserted, stamens of the second set have also declined. This sequence of anther and style/stigma development lessens chances for self-pollination.

Fire Pink - Silene virginicaPhoto 3:  Even with a dozen main stems, the inflorescence is very open.

Fire Pink - Silene virginicaPhoto 4:  The corolla and corona are the same vibrant red.  As shown, all ten stamens have emptied their anthers of pollen and twisted backwards as stigmas become receptive to receiving pollen from a different flower.  Note the enlarged calyx behind the corolla.

Fire Pink - Silene virginicaPhoto 5:  In the flower shown with petals, two sets of five anthers are exserted while styles have not yet appeared.  The two flowers without petals show styles emerging with stamens in decline (lower center) and styles fully exerted with receptive stigmas, while stamens have wilted (upper left – note enlarging ovary).  A ridged calyx (center) is also shown.

Upper portions of flowering stems have short, glandular hairs that cause the upper stem, upper cauline leaves and caudex exterior to feel sticky (viscid).  The greatest degree of stickiness occurs nearest the flowers.  Small flying insects are often caught on the sticky surfaces and ants cannot reach the nectar.

Upon completion of bloom, calyxes become swollen, point downward and dry to a light tan.  When dry, calyx teeth roll well back so that the upper portion (now hanging down) of the calyx has a gaping hole.  The placenta of the fruit within the calyx disintegrates and the seeds readily fall out.  The small round, tan to brown seeds, with a tight C-shape, are covered with minute crowded knobs.  With flowering completed, stems gradually disintegrate, with basal leaves remaining through the summer into winter.

Fire Pink - Silene virginicaPhoto 6:  As calyxes dry, teeth roll back and seeds easily drop out.

For a garden or natural area with partial sunlight and good drainage, fire pink is an excellent year-round, well-behaved, low-maintenance choice that puts on a spectacular show of striking spring color.  Flowers are a favorite of hummingbirds, which are a principal pollinator.  Should past-bloom stems be untidy as they decline, they can be easily removed.

Eight additional species of the genus are known to occur in Arkansas of which the only one with red flowers is royal catchfly (Silene regia).  Royal catchfly is a taller clump-forming, heavily pubescent plant with more closely spaced, numerous cauline leaf pairs and un-notched petals.   Starry campion (Silene stellata), a white flowering species of the genus, has been previously addressed in this series of articles.

*The word “pink” relates to the cutting of cloth with pinking shears to prevent threads in woven cloth from unravelling.  The ends of fire pink’s petals appear to be “pinked”.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Dwarf Larkspur

Dwarf larkspur (Delphinium tricorne) of the Buttercup (Ranunculaceae) family has distinctive early spring flowers that are often dark bluish-purple (or sometimes white).  Other common names include spring larkspur and rock larkspur.  This species is typically found in rich alluvial deciduous woods and wooded rocky slopes in shady to partially sunny sites with dry to mesic soils.  It is found in the U.S. from Nebraska to Pennsylvania and from Oklahoma to Georgia.  In Arkansas, it is found primarily in the Interior Highlands of the northwestern half of the state, with scattered occurrences in better drained sites of the Mississippi Alluvial Plain.  The genus name “Delphinium” comes from a Greek word for “dolphin” based on the shape of flower spurs and buds as viewed from the side.  The specific epithet refers to the horn shape of the three follicles that constitute the fruit.

This herbaceous perennial has an ephemeral nature in that it has new basal growth in mid-winter, blooms in early to mid-spring and then goes dormant in late-spring.  Roots are elongate, shallow and branched pointy tubers with long fibrous roots.  The upper ends of the tubers have growth points from which open clusters of basal leaves and weakly erect flowering stems (peduncles) grow.  Stems, in early spring, are straight and puberulent (covered with almost imperceptible downy hairs).  Cauline leaves, on stems up to 2 feet tall, are limited to a few alternate leaves along the lower portion of stems.  The uppermost of these few cauline leaves subtend single flowers while higher up single flowers are subtended by narrow, linear, pointed bracts that decrease in size up-stem.

Basal leaves that first appear in mid-winter and those basal leaves that grow later have the same appearance, and the cauline leaves, too, are similar, although these leaves may be smaller.  Basal and cauline leaves, which remain vibrant throughout the bloom period, have an overall orbicular shape that is palmately compound (trifoliate).  The three leaflets are then deeply incised to form wedge-shaped lobes which in turn are futher deeply cut.  The central leaflet of a leaf tends to be incised into three lobes and the two lateral leaflets tend to be incised into two lobes.  The leaf blade may appear to be five-lobed.  Smooth margins of all lobes are interrupted by additional minor lobing.

Mid-winter leaves have reddish, flat-topped and pubescent petioles and leaf blades have ciliate margins and pubescent undersides.  Petioles of mature basal leaves are about ¾ inch long, slender, medium green and slightly pubescent.  Mature leaf blades are several inches across and mostly glabrous.*

Dwarf Larkspur - Delphinium tricornePhoto 1:  In mid-winter, basal leaves unfurl.  Note pubescence of petioles and leaf blades.

Dwarf Larkspur - Delphinium tricornePhoto 2:  Appearance of trifoliate basal and cauline leaves are similar.  Early pubescence is lost as the season progresses.  Photo taken at end of March.

Pedicels (stalks of individual flowers) grow upward from the peduncle in raceme fashion at about 45 degrees.  Racemes, to about 8 inches long, have up to about 25 loosely arranged flowers.  Peduncles and pedicels tend to be densely covered with short pubescence, which falls off with age.  Light green flower buds on long pedicels are round in cross-section and stubby.  Flower buds have an exterior that features flattened, indented faces with five rounded bumps and a backward projecting spur.  Flowers typically have dark bluish-purple sepals and petals, although lavender to white flowers may occur and the upper two petals of dark blue flowers may be whitish.  Sepals, the most prominent part of the flower, are broad and petal-like with a gently tapered tip.  Sepals have an indented area near their apices that corresponds to the bumps that were present on the bud’s face.  Along with the upper sepal, which bears the long upward-angled, backward-pointing tubular spur, two sepals are opposite the flower’s center and two lower sepals are down-angled to either side.  The spur contains nectar that is pilfered by bumblebees by cutting into the spur and legitimately taken by smaller bees that enter from the front and typically effect pollination.  At anthesis, the corolla faces slightly downward, and the backward projecting spur is angled upward at about 90 degrees to the pedicels.  The soft half-inch-long spurs have a diameter that gradually decreases to a blunt tip.  Flowers have small round calices with a small elongate bract at each side.

In addition to the prominent sepals, flowers, up to 1 inch across and 1½ inches long, have two upper petals, two lower petals, a group of stamens and three pistils.  The two upper petals, which may be whitish or the same color as the sepals, are upward flared and overlapping.  The two lower petals, same color as sepals, are larger than the upper two petals, lobed and down-flared with numerous whitish, tangled hairs on their outer surface.  Stamens, grouped behind the lower petals (when viewed from front of flower) have whitish filaments and black dangling anthers.

Dwarf Larkspur - Delphinium tricornePhoto 3:  Flower buds, as seen at left as well as in right foreground, suggest the shape of a dolphin, the source of the genus name Delphinium.  Sepals and petals of this particular plant are all the same color.  Stamens can be seen behind the two lower hairy petals.

Dwarf Larkspur - Delphinium tricornePhoto 4:  Trifoliate leaves have many elongate, broad lobes with rounded apices.  Basal leaves remain vibrant throughout the growing season.  Photo taken in early April.

With fertilization, the three ovaries produce three free-standing, bean-pod-like follicles that are joined at their bases.  The half-inch-long glabrous follicles, with long pointed beaks, arch backward away from their common center, with sutures bowed-up.  When mature and dry in mid-spring, follicles split along sutures and small, black, smooth seeds are released.  With the fruiting cycle completed about late May, plants quickly return to dormancy.

Dwarf Larkspur - Delphinium tricornePhoto 5:  In early May, follicles are beginning to dry.  Trifoliate upper leaves exhibit relatively little lobing.

*Another widespread larkspur in Arkansas is Carolina larkspur (Delphinium carolinianum), with three subspecies in the state.  It also typically has dark blue or white flowers, but flower stems are up to 4 feet tall and plants are found in sunnier, drier sites.  In mid-winter, the basal leaves of the most common and widespread variety of D. carolinianum, var. carolinianum, and D. tricorne can be easily confused, both being trifoliate and similarly lobed.  However, in the case of D. carolinianum var. carolinianum, lobes of its later leaves become increasingly narrow and numerous.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Ohio Buckeye

Ohio Buckeye (Aesculus glabra) of the Soapberry (Sapindaceae) Family, formerly of the Horsechestnut (Hippocastanaceae) family, is a medium to large deciduous tree with opposite, palmately compound leaves.  It is native from central Texas to western Pennsylvania and as far north as central Iowa to southern Michigan and south into northern Alabama and the highlands of the northwestern half of Arkansas.  Other common names include “American buckeye” and “fetid buckeye”, based on the odor of crushed leaves, bark and twigs.  The name “buckeye” relates to the dark, ovoid poisonous seeds that have a lighter colored hilum (scar).

Ohio buckeye, with long taproots bearing lateral roots, typically occurs in rich, well-drained soils in sheltered areas of valleys, ravines and slopes in partial to full sunlight.  In shady drier sites, it tends to remain more shrub-like and may reach 15 feet in height, while in sunny favorable sites in Arkansas, it may be 40 feet*.  Trunk diameter remains relatively small at up to about 2 feet.  Large trees in sunny sites have numerous lateral branches that produce a broad rounded crown.  Lower lateral branches are lost in shady sites.  Plants as young as three years old can produce flowers.

Ohio Buckeye - Aesculus glabraPhoto 1:  This 4-inch seedling, in late January, would already have a long taproot.  Seeds in inset were similarly planted outdoors, but had not yet sprouted.

In winter, Ohio buckeye has round tapered lateral or axillary buds with sharp points that grow from just above previous year’s leaf scars. Terminal buds also form on nonflowering shoots.  Buds are covered with six or so layers of tightly imbricated scales with pointed tips.  Scales may have free-standing tips and are slightly keeled at their center.  Most new growth is from the uppermost lateral buds–the terminal inflorescence is shed in the fall or winter, leaving no true terminal bud on the original stem–as well as from terminal buds on sterile stems.   Many lateral buds do not develop. Because the terminal inflorescence is shed after seed dispersal, the dominance of the uppermost lateral buds gives the buckeye crown a very jagged architecture, which is best seen in winter.

In mid to late winter, the most-exterior scales just drop off while inner scales become light green and grow into long, leaf-like appendages before they, too, drop off.  By the time flowers reach anthesis, bud scales are gone and stem growth for the year has mostly stopped.

New stem growth is green and finely pubescent and becomes light gray and glabrous later in the year.  Previous years’ stems bear large, shield-shaped scars where leaves from previous years have dropped off.  Older branches are smooth and gray while larger trees have bark that has small scaly to knotty, unfurrowed plates that uniformly cover the trunk.

When new leaves first unfurl, the opposite leaves are a golden green which evolves to a pale green at time of bloom and a shiny, dark green upper surface in summer while the lower surface is a lighter green.  Leaves, glabrous on upper and lower surfaces, typically have five spreading leaflets, all attached at the tip of the petiole or leaf stalk (palmate), but three leaves and seven leaves may occur.  Leaves have slender, glabrous and light green petioles up to 6 inches long, with a broad clasping base and leaflets that are 3 to 6 inches long.  The central leaflet is the largest and, along with the next two lower leaflets, is at its broadest 1- to 2-inch-width in its upper portion with a gentle taper to its nearly stalkless base and a sharp taper to its apex (somewhat obovate).  The lower two broadly lanceolate leaflets are angled downward within the plane of the leaf.  While the lower portions of leaflets have entire margins, the upper portions have crenulated margins with tiny teeth.  In fall, leaves may become a showy reddish-orange.  Venation of the leaflets is pinnate with straight, equally spaced opposite or offset veins.

New terminal growth of more vigorous stems continues unabated to produce an upright panicle of flowers 6 to 8 inches long.  The peduncle of the inflorescence begins immediately above the last set of new leaves, initially with the same appearance as the stem below.  Above the peduncle, groups of flowers are spread around the rachis with spaces between groups.  Stalks of the floral groups are initially coiled against the rachis, but as growth continues, flower buds enlarge and open and the coils gradually straighten.  Flowers all along the rachis open in unison from the rachis to the ends of the straightened coils. Flowering continues for several weeks.  At the time of bloom, the color of the stem, peduncle, rachis and pedicel are the same light green with very short pubescence.  New stems and rachises become tan with rounded white lenticels (pores) by mid-summer.  Peduncles fall off after seeds mature and the capsules split open.

Ohio Buckeye - Aesculus glabraPhoto 2:  Terminal and lateral buds produce rapid spring growth.  These two stems grew from separate lateral buds at the top of previous year’s growth.

Ohio Buckeye - Aesculus glabraPhoto 3:  This plant, only several years old, produced its first inflorescence.  Note enlarged reddish bud scales about to fall off and (on the trunk) leaf scars, undeveloped buds and lenticels.

Ohio Buckeye - Aesculus glabraPhoto 4:  Opposite leaves typically have five leaflets with pinnate venation and crenulated margins.  Note axillary leaf buds for next years’ growth.

The inch-long greenish yellow flowers, on short pedicels, tend to be mostly male (staminate) with perfect (bisexual) flowers near the base of the cluster.  Flowers have greenish yellow, round and slightly elongated calyxes with five rounded lobes enclosing four greenish yellow petals.  The length of petals protruding from the calyx is greater than the length of the calyx itself.  Two long petals, that are flared upward, may bear orange, feather-like markings extending from flower’s throat to near the petals’ ends.  Two shorter broader petals flare outward in near-horizontal fashion.  Ends of the four petals are roundly flexed toward the flower center.  Flowers have seven stamens with white filaments and brown anthers that are exserted beyond the corolla, and a white pointed pistil (in perfect flowers only) equally exserted beyond the corolla.  The pistil of a flower becomes fully exserted (with open stigma) before the corolla has opened and before stamens are exserted.  Pistils are covered with a fine pubescence while filaments and the exterior of petals have longer hairs.

Ohio Buckeye - Aesculus glabraPhoto 5:  Flowers that are positioned along the upper sides of the clusters.   Pistil of a flower is exserted before flower’s stamens appear, as seen at lower left.

Ovaries of fertilized perfect flowers form green, round, spiky fruits.  Typically, only a few fruits at the base of the panicle will reach maturity.  Mature fruit is an ovoid, thick-skinned and light brown leathery capsule.  The capsule, up 1-½ inches in diameter, contains one to three seeds in separate chambers.  By the time fruit has matured, with multiple seeds, the seed exterior becomes flattened due to seed-to-seed contact.  At dehiscence (splitting of fruit) in late summer, capsules with sharp prickles split, so that the smooth, shiny mahogany-colored seeds with a light tan hilum, drop out.  Seeds, 1 to 1-½ inch wide and long, may germinate where dropped or wherever buried by squirrels or deposited by flowing water.  Seeds can germinate in mid-winter.  With germination, taproot and leaves quickly become fully functional so that cotyledons are not needed for photosynthesis.

Ohio Buckeye - Aesculus glabraPhoto 6:  Fertilized ovaries develop into fruit with long, weak spines.  Typically, only a few fruits per panicle will reach maturity.  Photo in mid-April.

Ohio Buckeye - Aesculus glabraPhoto 7:  Upper portion of panicle beyond developing fruit has dropped off in this mid-July photo.  Note two opposite brown buds at base of panicle’s peduncle (for next year’s growth).

Ohio buckeye can be a desirable garden or wildland plant where people, pets or livestock would not consume the seriously poisonous (contains aesculin) seed or other plant parts (squirrels apparently are not affected).   The plant is quite ornamental with its large attractive leaves, showy flowers and interesting fruit.  It has full-size flower panicles and fruit within several years of being planted by seed.  It also has nice fall color.  It does best in a mesic (moderately and evenly moist) soil where is protected from strong sun and wind.  Leaves may drop early in response to dry, hot conditions.

Red buckeye (Aesculus pavia) is the only other species of the genus native to Arkansas.  Red buckeye, occurring across most of the state, is more shrub-like, with red flowers and smooth seed capsules.  Native to the Appalachians and sometimes cultivated, yellow or big buckeye (Aesculus flava) is dissimilar to Ohio buckeye in various ways, including 1) its lower leaflets extending horizontally, 2) pistil and stamens that are not exserted, 3) fruit capsules that are smooth, and 4) typical height at maturity of 70-90 feet.  Bottlebrush buckeye (Aesculus parviflora) is a species native to the Southeast U.S., primarily Alabama, that is also frequently cultivated.  It differs from Ohio buckeye in having a definite shrubby habit, with multiple stems and suckers from the base, and longer panicles of white flowers with long-exserted filaments.

*  National champion Ohio buckeye in Casey County, KY, is 148 feet tall.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Arkansas Yucca

Arkansas yucca (Yucca arkansana) of the Agave (Agavaceae) family, formerly of the Lily (Liliaceae) family, an evergreen shrub, occurs in Texas, Oklahoma, southeastern Kansas, southern Missouri and Arkansas.  In Arkansas, it occurs throughout much of the Interior Highlands.  The genus name originates from a misapplication by Carl Linnaeus in 1753 of a common name for cassava (Manihot esculenta), “yuca”, to the plants for which he named and described the genus Yucca.  Arkansas yucca, also called soft-leaf yucca and soapweed, is found in well-drained soils in sunny prairies, glades and rocky hillsides as well as along partially sunny edges of woodlands and thickets.

A mature Arkansas yucca has a several-feet-long taproot-like rhizome (round in cross-section) along with near-surface, short lateral side-rhizomes with scattered fibrous roots.  Rhizomes, that may be one or more inches in diameter, have a brown epidermis with a white potato-like interior.  Ends of lateral rhizomes turn toward the surface, where apical buds each produce a single rosette of leaves.  A mature plant may have a half-dozen scattered leaf rosettes so that a single plant appears to be a colony.  Single, stout, woody flower stalks grow from the apical bud within leaf rosettes.  After flowering, those rosettes and their rhizome segment die and decay, leaving a circular scar or nub on the surviving portion of the root.

Arkansas Yucca - Yucca arkansanaPhoto 1:  Rhizome segment at coin is decaying after producing a flower stalk marked with a red asterisk.   Lateral roots grew from either side of dead segment.  Taproot is below coin.

Leathery, flexible, radiating leaves, up to about 2 feet long, are widest at mid-leaf with a width of about ¾ inch.  Reduction of leaf width towards its pointed, pliable tip is very gradual.  White-edged margins are straight and entire with scattered, long, loosely curly white filaments that exfoliate from margins.  Leaves, a dull medium green above and below, become narrow near their bases before finally widening within the rosette base.  Leaves are convex below and concave above, with a thickened stout midrib.  Toward mid-leaf, the blade gradually widens and the midrib becomes less prominent and less stout where leaf may bend down and become twisted.  Parallel venation is not prominent on the upper or lower leaf surfaces.  Bases of previous years’ leaves gradually weaken so that old leaves gradually lower to lie on the ground where they remain for several years, gradually decaying.  Plants are usually stemless (acaulescent), although a short leaning (decumbent) stem, mostly hidden by decaying leaves, may develop.   Leaf rosettes may be 2 feet tall and wide.

Arkansas Yucca - Yucca arkansanaPhoto 2:  Long, radiating leaves are thin and pliable with a flexible point.  Photo taken in late February.

Arkansas Yucca - Yucca arkansanaPhoto 3:  Curly marginal hairs are not twisted as on some other species.  Appreciable stems do not develop as leaf rosettes add new central leaves from year to year.

The inflorescence in early spring consists of a raceme on a tall, stiffly erect, somewhat woody and glabrous stalk (rachis) that grows from the apex of a leaf rosette.  Stalks, usually 2 to 3 feet tall (but may reach 6 feet), have narrow, triangular and purplish bracts that are erect, both below and within the raceme.  As erect flower buds on the upper portion of the stalk enlarge behind bracts, those bracts wither and the pedicels bend down at the base of flowers so that, at anthesis, flowers droop and face the ground.  Individual flowers are on short pedicels (attached directly to the stalk) or in a group of several flowers (small panicles) on short pedicels growing from a short peduncle (which, in turn, is attached to the stalk).   Individual plants typically do not bloom every year.

Arkansas Yucca - Yucca arkansanaPhoto 4:  In this early May photo, purplish bracts on elongating flower stalk subtend and cover developing flower buds.  Bracts are relatively short, as compared to other species.

Arkansas Yucca - Yucca arkansanaPhoto 5:  Flowers, on short pedicels and peduncles, occur in a raceme-type inflorescence.  Bracts on the lower portion of the stalk do not dry while flowers are in bloom.  Site shown is mostly shady with loam soil.

The bell-shaped (campanulate) flowers, up to 2½ inch long and 2 inches wide, have six greenish-white, thickened tepals (three sepals and three petals) that have narrow bases, gentle tips and broadened mid-sections.  The whitish green superior ovary is topped by a pale green lobed style that terminates in a three-lobed stigma with a recessed central orifice.  Six round, white, post-like filaments, attached to flower receptacle below the ovary, bear pollinia (packets of pollen) at their rounded apices.  Yucca moths are the only insects that can successfully pollinate yucca flowers and developing yucca fruit is the only larval food source for yucca moths.*

Arkansas Yucca - Yucca arkansanaPhoto 6:  Heavy flowers cause pedicels to bend down.  As seed capsules develop, strengthened pedicels bend upward.

Arkansas Yucca - Yucca arkansanaPhoto 7:  The lobed green style sits above a large ovary that is surrounded by post-like filaments and thick tepals.  Honey ants can be seen feeding on flower bud resin.

As flowers pass anthesis, pedicels twist upward so that developing green, tough, seed capsules are upright.  Capsules, about 2½ inches long and 1 inch wide on woody pedicels and stalks, are composed of three elongate carpels that are divided into two locules, each with a stack of seeds.  A capsule, with an exterior that is rounded along each elongate carpel, contains 100+ seeds.  When capsules dry in late fall, they become papery and split from the top between carpels.  As capsules open, hundreds of flattened, rounded papery black seeds are shaken out on windy days and become scattered.   Dead stalks and racemes are somewhat woody and may persist for several years.

For a partially shady to sunny garden or natural area, Arkansas yucca can provide a dominant accent.  It has year-round presence, but its occasional flowering would be the highlight of a year.  Its large flowers dramatize its flower parts and its mutulistic relationship with yucca moths can be easily viewed.  In addition to the yucca moth, yucca plants are host plants for caterpillars of giant-skipper butterflies, including Megathymus yuccae, which is rare in Arkansas.

Identification of yucca species and their varieties has found disagreement among various authorities.  In Arkansas, four additional species are recognized: Yucca filamentosa (common names: Adam’s-needle, Spanish-bayonet or yucca), a Southeastern U.S. native introduced to Arkansas;  Yucca flaccida (same common names: Adam’s-needle, Spanish-bayonet or yucca), also a Southeastern U.S. native introduced to Arkansas; Yucca louisianensis (common names: Louisianna yucca, Gulf Coast yucca or sandhill yucca), a native to south Arkansas; and Yucca freemanii (common name: Freeman’s yucca), a native species discovered new to Arkansas in Miller County in 2014.  Characteristics that can help distinguish Yucca arkansana from these other species:  1) leaf rosettes of a plant are separated by a short distance, instead of bunched-up, 2) there are relatively few leaves per rosette and these are thin and pliable with weak pointed tips, 3) curly marginal leaf hairs are not twisted, 4) flowering stalks are generally less than 4 feet, 5) inflorescence is a raceme, instead of a panicle, and 6) color of style is a contrasting green within a whitish green flower.

*   Yuccas and yucca moths, totally dependent on each other for natural reproduction, represent an example of mutualism.  The moth species that pollinates Arkansas yucca (as well as some other yuccas) is Tegeticula yuccasella.   Female moths pack pollinia into a ball that they carry in their coiled palps to another plant.  Moths use their egg-laying abdominal appendage (ovipositor) to insert an egg into a chamber (locule) of the ovary.  The transported pollinium ball is then forced into the recessed orifice of the stigma to effect pollination.  The moth caterpillar, as it grows, ties a dozen or so ovules together as they are eaten, leaving hundreds of viable seed.  Mature caterpillars drop to the ground to pupate in the soil until the next year’s flowers are available.

Article and photographs by ANPS member Sid Vogelpohl

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Know Your Natives – Marginal Wood Fern

Marginal wood fern (Dryopteris marginalis) of the Wood Fern (Dryopteridaceae) family, an evergreen, twice-cut fern, occurs throughout much of the eastern U.S. from Texas and Minnesota to the Atlantic Coast, but is mostly absent from the Gulf Coastal Plain and unknown from Louisiana and Florida.  In Arkansas, it has been documented to occur in the northwestern half of the state in the Interior Highlands.  The genus name is based on the Greek words for “oak” and “fern” which relates to occurrence of wood ferns among oak forests.  The specific epithet relates to the location of sori (spore clusters) near the margins of the leaf divisions.  Other common names include marginal shield fern, evergreen wood fern and leather-leaf wood fern.  This fern favors areas with damp to dry soils found in partially to fully shaded, rocky ravines, stream banks and north-facing slopes.

This medium-large fern has an elongate, reclined and compact rootstock, with dense, shallow and branching fibrous roots that grow directly from the rootstock.  New fronds grow from the center of the rootstock in late winter as tightly curled fiddleheads (or croziers).  Fiddleheads are densely covered with light-brown, soft scales and filaments, especially near their bases.  Scales and filaments, mostly absent along the leaf rachis (the midrib of the compound leaf), remain on the stipe (leaf stalk) of mature leaves.  The stipe and rachis, yellowish-green overall, with the stipe having a reddish base, have a central upper groove along the flattened upper side and a convex lower side.  The plant has a vase-like overall shape with an airy appearance.  Old fronds (an alternative term for fern leaves) wither around the rootstock, with the old stipes persisting for several years.

photo-1-january-11Photo 1:  Rootstock, as seen in this mid-January photo, bears green fronds which will decline when new fronds appear in late winter .  Previous year’s stipes remain.  Fibrous roots have been removed.

photo-2-april-6Photo 2:  In early April, fiddleheads grow from center of rootstock.  Previous year’s and older stipes can be seen.

Stipes are one-fourth to one-third as long as the fronds: fronds are up to 18 inches long and 6 inches wide, with stipes being about 8 inches long.  Plants, with fronds ascending and arching, may have a height and width each of about 2 feet.  Fronds are a bluish-green above and a lighter yellowish-green below.  Fronds have linear-lanceolate, quickly tapering pinnae (leaflets) that are mostly opposite lower on the rachis, changing to alternate higher toward the apex.  The leathery, glabrous pinnae, with tips arching toward frond apex, are longest mid-frond and slightly shorter toward the frond base (a semi-tapered fern).  Separation of pinnae, greatest at frond base, decreases toward frond tip as pinna length quickly shortens to form an acuminate frond tip.  Pinnae approaching frond tip are lobe-like.  Lower pinnae have very short petiolules (stalks) that are perpendicular to the rachis, with upper, short pinnae becoming sessile without overlap.  Pinnae are divided (a twice-divided fern) into oblong pinnules (sub-leaflets or secondary leaflets) which have rounded tips and margins that vary from slightly crenulate to well lobed near the base of the rachis.  Fronds may have 15 or so easily distinguishable pairs of pinnae before the pinnae become indistinct at the frond apex.  Similarly, pinnae may have 15 or so easily distinguishable pairs of pinnules before they become indistinct at the pinna tip. Main and secondary veins of pinnules, purplish on the underside, are of about equal size, nearly straight and forked.   Main veins of pinnules extend onto pinna midribs.

photo-3-january-11Photo 3:  In early January, fronds remain green.  All fronds of a plant grow from center of rootstock without any off-sets.

Fertile and sterile fronds are both photosynthetic and have the same appearance when viewed from above.  However, fertile fronds bear fruit-dots or sori* (singular, sorus) on the undersides of their upper (or distal) pinnae. Each sorus is a cluster of spore-producing sporangia.  From one to five sori are located along the margins of pinnules, except on those pinnules near the frond apex that are more lobe-like. Single sori are located at sinuses between the lobes.  Sori, which develop in late spring, are at first green, but then become dark brown.  They are covered by umbrella-like shields called indusia (singular, indusium).  Indusia are attached to the pinnule at the sori’s depressed center, loosely extending over many sporangia.  Each sporangium or spore case produces 50+ spores.  The depressed center of a sorus along with a depressed side cause the round sorus to appear reniform (kidney-shaped).  As the spore cases grow, they push the indusium aside.  When air moisture is low, spore cases pop open and dust-like spores are dispersed into the air.**

photo-4-marginal-fmid-right-1Photo 4:  Upper surfaces of an infertile frond (left) and a fertile frond (right) appear the same.  Lower side of a fertile frond, which has already dispersed its spores, is shown in the middle.  As shown, pinnae may be opposite or alternate.  Photo taken in mid-July.

photo-5-marginal-yPhoto 5:  Sori are located at margins of pinnules or singly at sinuses between lobe-like pinnules, as shown.  Note venation and slightly crenulate margins.  Photo taken in mid-July.

In a garden or natural areas, this evergreen, medium-large fern with its nice form and good texture can serve as an accent plant that provides interest throughout the year.  The plant does not colonize or spread by rootstock.  Dead fronds, spread around the rootstock, quickly disintegrate and tougher stipes, which disintegrate in two years, are not especially noticeable.  Marginal wood fern does well in partial to full shade.  It is not a preferred food choice of deer or rabbits.  Once established, it can survive dry periods.

photo-6-may-15Photo 6:  Marginal wood fern is an excellent accent plant for a shady garden.

In Arkansas, 12 taxa in the wood fern family (Dryopteridaceae) have been documented outside of cultivation, of which nine (including several of hybrid origin) are native.  Of the natives, only two species are widespread evergreen ferns, namely the subject species marginal wood fern (Dryopteris marginalis) and Christmas fern (Polystichum acrostichoides).  Christmas fern is a distinctive, once-cut fern with darker green and more leathery pinnae.

*   Sori – Greek for “heaps”.
**  With dispersal of spores, the above-ground “diploid sporophyte phase” of a fern’s life cycle (referred to as “alternation of generations”) concludes.  In the soil, spores germinate to produce a prothallus as the “haploid gametophyte phase”.  The prothallus produces mobile sperm gametes and attached egg gametes.  With fertilization of the egg, a diploid zygote may develop into a new diploid sporophyte plant.

Article and photographs by ANPS member Sid Vogelpohl

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