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PTERIDOPHYTES OF NORTHEAST ALABAMA AND ADJACENT HIGHLANDS.

I. ANNOTATED CHECKLIST AND KEY TO FAMILIES

INTRODUCTION

This project is a guide to all the ferns and fern allies of the northeast Alabama. Plant species occurring in adjacent highland counties are also part of this flora. The study area includes 84 specific and infraspecific taxa, representing a total of 38 genera, 18 families, 6 orders, 4 classes, and 3 divisions. Pteridophytes in our area include the following divisions: Equisetophyta, Lycopodiophyta, and Polypodiophyta. The first two divisions are the so-called fern allies, which are really not allied to the true ferns (Polypodiophyta). The guide will include illustrations, maps, identification keys, habitats, distributional data, conservation status, uses, and pertinent synonymy.

The area delineated as Northeast Alabama includes Blount, Calhoun, Cherokee, Clay, Cleburne, Cullman, DeKalb, Etowah, Jackson, Jefferson, Limestone, Madison, Marshall, Morgan, Randolph, Saint Clair, Shelby, and Talladega counties. Adjacent highland counties include Bibb, Chambers, Chilton, Coosa, Lauderdale, Lawrence, Tallapoosa, Tuscaloosa, Walker, and Winston (Figure 1). The highlands of Alabama consists of the following Provinces: Interior Low Plateau (Highland Rim), Appalachian Plateau (Cumberland Plateau), Ridge and Valley, and Piedmont Plateau (Figure 2).

CHECKLIST OF PTERIDOPHYTES

Format

The checklist is based primarily upon herbarium specimens deposited at the Jacksonville State University Herbarium (JSU). The list also includes data from pertinent literature and other southeastern herbaria (Auburn University [AUA], University of Alabama [UNA], University of North Alabama [UNAF], and Vanderbilt University [VDB]). Taxa not found at JSU are noted by a herbarium acronym or literature reference. Nomenclature follows Flora of North America [FNA] (1993+) and more recent publications. Synonymy is primarily from Radford eta). (1968) and is italicized within brackets. Families, genera, specific and infraspecific taxa are arranged alphabetically within major vascular plant groups (divisions). Introduced taxon are followed by a dagger (+). Federal or state listed species are followed by a star (*) and their coded status designation (recent listing in plainface type; formerly listed preceded by an x and in italics). The coded ranks are defined as follows: S1, Critically Imperiled; S2, Imperiled; S3, Rar e; SH, of historical occurrence; LE, Endangered Species; LT; Threatened Species. Data are from the Alabama Natural Heritage tracking and inventory lists (1994, 1996, 1997, 1999).

Annotated Checklist

DIVISION EQUISETOPHYTA

EQUISETACEAE (Horsetail Family)

Equisetum arvense L., Field Horsetail.* S2

E. hyemale L. ssp. affine (Engelm.) A.A. Eaton, Scouring-rush

DIVISION LYCOPODIOPHYTA

ISOETACEAE (Quillwort Family)

Isoetes butleri Engelm., Glade Quillwort; Butler's Quillwort.* S2

I. engelmannii A. Br., Appalachian Quillwort; Engelmann's Quillwort.* xS3

I. melanopoda Gray & Durieu, Blackfoot Quillwort.* S1? {Cherokee/VDB}

I. piedmontana (Pfeiffer) Reed, Piedmont Quillwort.* S2

LYCOPODIACEAE (Clubmoss Family)

Diphasiastrum digitatum (Dill, ex A. Br.) Holub, Running Ground-pine; Ground-cedar. [Lycopodium digitatwn; L. flabelliforme]

D. trisrachyum (Pursh) Holub., Blue Ground-cedar; Deep-root Clubmoss * xSR [Lycopodium tristachywn]

Huperzia lucidula (Michx.) Trevisan, Shining Clubmoss. S2 [Lycopodiwn luciduiwn] {Jackson/Whetstone 1981}

H. porophila (Lloyd & Underw.) Holub, Rock Clubmoss.* S1 [Lycopodium porophilum] {Winston/UNA}

Lycopodiella appressa (Chapm.) Cranfill, Southern Clubmoss; Slender Clubmoss. [Lycopodiun appressum]

L. alopecuroides (L.) Cranfil, Fox-tail Clubmoss. [Lycopodium alopecuroides]

Lycopodium obscurum L., Ground-pine; Tree Clubmoss.* S1

SELAGINELLACEAE (Spikemoss Family)

Selaginella apoda (L.) Spr., Meadow Spikemoss.

S. arenicola L. spp. riddellii (Van Eselt.) Tryon, Sand Spikemoss; Riddell's Spikemoss.* S2 [S. riddellii {Shelby/VDB}

S. braunii Baker, Braun's or Treelet Spikemoss. [+]

S. rupestris (L.) Spring, Rock Spikemoss; Ledge Spikemoss.* S253

DIVISION POLYPODIOPHYTA

ASPLENIACEAE (Spleenwort Family)

Asplenium bradleyi D.C. Eat., Cliff Spleenwort; Bradley's Spleenwort.* S2

A. x ebenoides Scott, Scott's Spleenwort.* S1 [A. plazyneuron x A. rhizophyliwn] {Jefferson/Whetstone 1981}

A. x gravesii Maxon, Graves' Spleenwort. [A. bradleyi x A. pinnatifidum] {Jackson, DeKalb & Etowah/Short 1978}

A. monanthes L., Single-sorus Spleenwort.* S1 {Jackson & Talladega/VDB}

A. montanwn Wilid., Mountain Spleenwort.

A. pinnatifidum Nutt., Lobed Spleenwort.

A. platyneuron (L.) Oakes, Ebony Spleenwort.

A. resiliens Kunze, Blackstem Spleenwort.

A. rhizophyllum L., Walking Fern. [Camptosorus rhizophyllus]

A. ruta-muraria L., Wall-rue Spleenwort.* S2 {Etowah/UNA}

A. scolopendrium L. var. americanum (Fern.) Kartesz & Gandhi, Hart's-tongue Fern.* S1/LT [Phyllitis scolopendrium var. americanum]

A. trichomanes L., Maidenhair Speenwort. *S2S3 xS1

A. x trudellii Wherry, Trudell's Spleenwort. [A. montanum x A. pinnatifidum] {Jackson & Etowah/Short 1978}

AZOLLACEAE (Mosquito Fern Family)

Azolla caroliniana Willd., Mosquito Fern.

BLECHNACEAE (Chain Fern Family)

Woodwardia areolata (L.) Moore, Netted Chain fern; Net-veined Chain Fern. [Lorinseria areolata]

W. virginica (L.) Smith, Virginia Chain Fern; Southern Chain Fern.

DENNSTAEDTIACEAE (Cuplet Fern Family)

Dennstaedtia punctilobula (Michx.) T. Moore, Hay-scented Fern; Boulder Fern.* xS3

Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underw., Eastern Bracken Fern.

P. a. var. pseudocaudatum (Clute) Heller, Southern Bracken Fern; Tailed Bracken Fern.

DRYOPTERIDACEAE (Wood Fern Family)

Athyrium filix-femina (L.) Roth var. asplenioides (Michx.) Farw., Southern Lady Fern; Lowland Lady Fern. [A. asplenioides]

Cystopteris bulbifera (L.) Bern., Bulblet Bladder Fern; Berry Bladder Fern.* xS?

C. protrusa (Weath.) Blasdell, Spreading Bladder Fern; Lowland Bladder Fern. [C. fragilis var. protrusa]

C. tennesseensis Sh., Tennessee Bladder Fern.*S2

Deparia acrostichoides (Sw.) Kato, Silvery Glade Fern. [Athyrium thelypteroides]

Diplazium pycnocarpon (Spreng.) M. Broun, Glade Fern. [Athyrium pycnocarpon]

Dryopteris x australis (Wherry) Small, Southern Wood Fern.* S1

D. celsa (Palmer) Knowlton, Log Fern.* S1

D. intermedia (Muhl.) Gray, Fancy Fern; Evergreen Wood Fern.

D. marginalis (L.) Gray, Marginal Shield Fern; Leather Wood Fern.

Onoclea sensibilis L., Sensitive Fern; Bead Fern.

Polystichum acrostichoides (Michx.) Schott, Christmas Fern.

Woodsia obtusa (Spreng.) Torr., Blunt-lobed Cliff-fern; Common Woodsia.

HYMENOPHYLLACEAE (Filmy Fern Family)

Hymenophyllum tayloriae Farrar & Raine, Taylor's Filmy Fern; Gorge Filmy Fern.* S1 {Lawrence & Winston/UNAF}

Trichomanes boschianum Sturm, Bristle Fern; Filmy Fern.* xS3

T. intricatum Farrar, Weft Fern.

T. petersii Gray, Dwarf Filmy Fern; Peter's Bristle Fern.* S2

LYGODIACEAE (Climbing Fern Family)

Lygodium palmatum (Bernh.) Sw., American Climbing Fern.* S2? xS1

L. japonicum (Thunb.) Sw., Japanese Climbing Fern. +

MARSILEACEAE (Water-clover Family

Pilularia americana Braun, Pillwort.* S1 {Lauderdale/UNAF}

OPHIOGLOSSACEAE (Adder's-tongue Family)

Botrychium biternatum (Savigny) Underw., Southern Grapefern.

B. dissectum Spreng., Cut-leaf Grapefern; Common Grapefern. [B. d. var. obliquum]

B. jenmanii Underw., Alabama Grapefern.* SH {DeKalb/Dean 1968}[B. alabamense]

B. lunarioides (Michx.) Sw., Winter Grapefern.* SH {Morgan/AUA}

B. virginianum (L.) Sw., Rattlesnake Fern; Virginia Grapefern.

Ophioglossum engelmannii Prantl, Limestone Adder's-tongue.* S2S3

O. crotalophoroides Walt., Bulbous Adder's-tongue.* xS3

O. vulgatum L., Common Adder's-tongue. [O. v. var. pycnostichum]

OSMUNDACEAE (Royal Fern Family)

Osmunda cinnamomea L., Cinnamon Fern.

O. regalis L. var. spectabilis (Willd.) Gray, Royal Fern.

POLYPODIACEAE (Polypody Family)

Pleopeltis polypodioides (L.) Andr. & Wind. var. michauxiana (Weatherby) Andr. & Wind. Resurrection Fern; Gray Polypody. [Polypodium polypodioides]

Polypodium virginianum L., Rockcap Fern; Common Polypody. [Inc. P. appalachianum]

PTERIDACEAE (Maidenhair Fern Family)

Adiantum capillus-veneris L., Southern Maidenhair Fern; Venus'-hair Fern.

A. pedatum L., Common Maidenhair Fern; Northern Maidenhair Fern.

Astrolepis integerrima (Hook.) Benh. & Wind., False Cloak Fern; Star-scaled Cloak Fern. *S1 [Notholaena integerrima]{Bibb/Allison 1996}

Cheilanthes alabamensis (Buckl.) Kunze, Alabama Lip Fern. *xS3

C. lanosa (Michx.) D.C. Eat., Hairy Lip Fern.

B. tomentosa Link, Woolly Lip Fern.

Pellaea atropurpurea (L.) Link, Purple Cliff-brake.

Pteris multifida Poir. ex Lam., Spider Brake; Wall Fern. [+]

THELYPTERIDACEAE (Marsh Fern Family)

Macrothelypteris torresiana (Gaud.-Beaup.) Ching, Mariana Maiden Fern. [+] [Thelypteris torresiana]

Phegopteris hexagonoptera (Michx.) Fee, Broad Beech Fern. [Thelypteris hexagonoptera]

Thelypteris kunthii (Desv.) Morton, Widespread Maiden Fern; Southern Shield Fern.

T. noveboracensis (L.) Nieuwl., New York Fern.

T. ovata R. St. John, Ovate Maiden Fern. * S3 {Bibb/Allison 1996}

T. palustris Schott var. pubescens (Lawson) Fern., Marsh Fern.

T. pilosa (Mart. & Gale) Crawf. var. alabamensis Crawf., Alabama Streak-sorus Fern. * S1/LT [Leptogramma pilosa var. americana]{Winston}

VITTARIACEAE (Shoestring Fern Family)

Vittaria appalachiana Farrar & Mickel, Appalachian Shoestring Fern.

KEY TO PTERIDOPHYTE FAMILIES

1. Plant floating on water or stranded on moist substrate (not rooted in soil) Azollaceae

1. Plant rooting in soil or growing on trees or rocks.

2. Plant with gametophyte generation only, lacking sporangia; plant filamentous or resembling a liverwort; growing in non-calcareous rock crevices.

3. Plant filamentous or ribbon-like Hymenophyilaceae (in part)

3. Plant leaf-like, not filamentous or ribbon-like Vittariaceae

2. Plant with sporophyte generation; plant not filamentous or not resembling a liverwort; habitat various.

4. Stems hollow, jointed, and ridged; leaves inconspicuous and sheathed around stem; sporangia in cone-like strobili terminating stem Equisetaceae

4. Stems not hollow, non-jointed, and lacking ridges; leaves conspicuous and not forming sheaths around stem; sporangia variously borne.

5. Plant grass-like; sporangia borne at leaf base.

6. Plant tufted; sporangia enclosed in a cavity at leaf base Isoetaceae

6. Plant on short creeping stems, not tufted; sporangia numerous in a sporocarp attached by short stalk (1-3 mm) at base of leaf Marsileaceae (Pilularia)

5. Plant not grass-like; sporangia variously borne on leaf surfaces.

7. Plant moss-like; leaves simple and greatly reduced; blades bearing a single midvein or midveins wanting; sporangia in axils of leaves or in cone-like strobili.

8. Leaves tightly appressed (imbricate) and scale-like.

9. Plant more than 5 cm tall; strobili cylindric and on branching slender stalks or sessile Lycopodiaceae (Diphasiastrum)

9. Plant less than 5 cm tall; strobili quadrangular (4 sided) and sessile Selaginellaceae (in part)

8. Leaves spreading or loosely appressed, not scale-like.

10. Leaves dimorphic, of two different shapes and sizes (lateral and median leaves); strobili quadrangular Selaginellaceae (in part)

10. Leaves not dimorphic, all of similar sizes; strobili cylindric or sporangia solitary in axils of leaves Lycopodiaceae (in part)

7. Plant not moss-like; leaves often large and elaborate, simple or compound; blades with numerous lateral veins; sporangia usually borne on leaf surfaces, not in leaf axils or in cone-like strobili.

11. Plant vine-like, twining, and usually climbing on vegetation Lygodiaceae

11. Plant not vine-like and not climbing on vegetation.

12. Leaves not deeply lobed or dissected.

13. Sporangia borne on a separate fertile, spike-like sporophore, usually arising from a single leaf (occasionally 2 or more leaves in some species) Ophioglossaceae (in part)

13. Sporangia borne on the underside (abaxial surface) or margin of leaf, not on a sporophore.

14. Leaves less than 3 cm long, very thin and translucent (1 cell thick) Hymenophyllaceae (in part)

14. Leaves more than 3 cm long, thicker and not transparent (2 or more cells thick) Aspleniaceae (in part)

12. Leaves deeply lobed to dissected.

15. Blades on sterile (non-sporulating) leaves, 1-pinnatifid (sometimes pinnate at base).

16. Leaves monomorphic, sterile and fertile (sporulating) leaves similar in appearance and both pinnatifid (deeply lobed); leaves evergreen.

17. Sori round and without indusia (a protective covering); leaf stalk (petiole) green Polypodiaceae

17. Sori elongate with indusia; leaf stalk brown, at least near base Aspleniaceae (in part)

16. Leaves dimorphic, fertile (sporulating) leaves greatly dissimilar to sterile (non-sporulating) leaves, only sterile ones pinnatifid; leaves not evergreen.

18. Sterile leaves with mostly with opposite pinnae, margins undulating to lobed; fertile leaves forming bead-like clusters Dryopteridaceae (Onoclea)

18. Sterile leaves mostly with alternately arranged pinnae, margins with small teeth (serrulate); fertile leaves pinnate, not forming bead-like clusters Blechnaceae (in part)

15. Blades on non-sporulating leaves 2-pinnatifid, pinnate or more than 1- pinnate.

19. Sporangia borne on branched fertile segments ("spikes") arising from a single leaf Ophioglossaceae (in part)

19. Sporangia borne on underside or margin of leaf, not on branched fertile segments.

20. Sterile and fertile leaves or leaflets strongly dimorphic; fertile portions lacking leafy tissue; flat stipules present at base of leaves; sporangia without well defined annulus (row of thick-walled cells) Osmundaceae

20. Sterile and fertile leaves similar or only slightly dimorphic; fertile portions with leafy tissue; stipules lacking at leaf bases; sporangia with well developed annulus.

21. Leaves thin and translucent (1 cell thick between veins); leaf blades usually less than 20 cm long; sporangia borne along margins of leaflets in a tubular cup-like structure (involucre) with an exserted bristle; plant growing on acidic rock (e.g., sandstone)Hymenophyllaceae (in part)

21. Leaves much thicker; sporangia borne on the underside (abaxial surface) or margins of leaflets, but not in an involucre with an exserted bristle; plants growing in soil or on various types of rock (basic or acidic).

22. Sori located along margins of leaflets, either covered by rolled under (revolute) leaflet edge or in cup-like indusia.

23. Sori in cup-like indusia and not covered by edge of leaflets; leaf blades and rachises bearing gland-tipped whitish hairs (with a hay-like fragrance); rhizome hairy Dennstaedtiaceae (Dennstaedtia)

23. Sori covered by rolled under leaflet edge; leaf blades glabrous or pubesecnt, but lacking white gland-tipped hairs; rhizome scaly or hairy.

24. Leaf stalk 3-branched; leaves (3 main divisions) broadly triangular; rhizome hairy Dennstaedtiaceae (Pteridium)

24. Leaf stalk not 3-branched; leaves not broadly triangular; rhizome scaly Pteridaceae

22. Sori not located along the edge of leaflets, or sori marginal with kidney-shaped indusium and not covered by rolled under leaflet edge.

25. Veins areolate along mid-veins of leaflets, forming a series of chain-like loops; son elongate and located along each side of mid-vein in single chain-like rows Blechnaceae (in part)

25. Veins of leaflet not areolate; sori round or elongate and located along lateral veins and mid-veins of leaflets, but not in single end to end rows.

26. Leaves small to medium-sized (less than 12 cm wide) and evergreen; leaf stalks (petioles) wiry; sori elongate bordering veins only along one side (except in Thelypteris pilosa).

27. Leaves with transparent, needle-like (pointed-tipped) hairs; sori lacking indusia; plant known only from Winston County growing on sandstone cliffs Thelypteridaceae (T. pilosa)

27. Leaves lacking transparent, needlelike hairs (hairs have blunt tips, if present); sori with indusia; plants widespread in various habitats Aspleniaceae (in part)

26. Leaves large (more than 12 cm wide end deciduous; leaf stalks stout, sori round or elongate and usually partially covering veins.

28. Leaves with transparent, needlelike (pointed-tipped) hairs that are simple to branched; sori with or without indusium; rhizome creeping (mostly subterranean) and slender, less than 1 cm in diameter Thelypteridaceae (in part)

28. Leaves with blunt-tipped, simple hairs (not transparent or needlelike) hairs (or leaves glabrous); sori with indusium; rhizome ascending and thicker, more than 1 cm in diameter Dryopteridaceae (in part)

ACKNOWLEDGMENTS

We are especially grateful to the late Warren Herb Wagner, Jr. who thoroughly reviewed this flora. He was a magnificent botanist who inspired us with his zeal. Special thanks goes out to Robert Kral for his many comments and suggestions. He is the authority on Alabama plants and his review made this flora complete. We also thank Jack Short for reading through the manuscript and providing valuable insight. The grammatical review of Verna Gates of Birmingham greatly improved this work. We thank Marion Montgomery, of the Anniston Museum of Natural History, who painstakingly drew the original illustrations. Finally, we appreciate the contributions of Tern Ballard (Equisetaceae, Ophioglossaceae, Lygodiaceae), Tim Hofmann (Dennstaedtiaceae), and Steve Threlkeld (Adiantum).

REFERENCES CITED

Alabama Natural Heritage Program [ANHP]. 1994. Vascular Plant Inventory Tracking List, April edition. Montgomery, Alabama.

Alabama Natural Heritage Program. 1996. Species Inventory List. Montgomery, Alabama.

Alabama Natural Heritage Program. 1997. Inventory List of Rare, Threatened, and Endangered Plants, Animals, and Natural Communities of Alabama. Montgomery, Alabama.

Alabama Natural Heritage Program. 1999. Inventory List of Rare, Threatened, and Endangered Plants, Animals, and Natural Communities of Alabama. Montgomery, Alabama.

Allison, J. R. 1996. A "lost world" in Bibb County, Alabama. Georgia Department of Natural Resources, Georgia Natural Heritage Program.

Dean, B. E. 1968. Ferns of Alabama. Southern University Press, Birmingham.

Flora of North America [FNA] Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 4+ vols. New York and Oxford.

Radford, A. E., H. E. Ahles and C. R. Bell. 1968. Manual of the Vascular Flora of the Carolinas. The University of North Carolina Press, Chapel Hill.

Short, J. W. 1978. Distribution of Alabama Pteridophytes. M.S. Thesis, Auburn University, Auburn.

Whetstone, R. D. 1981. Vascular flora and Vegetation of the Cumberland Plateau of Alabama... Ph.D. Dissertation, Department of Botany, University of North Carolina at Chapel Hill.

II. EQUISETOPHYTA AND LYCOPODIOPHYTA

INTRODUCTION

The divisions covered in this work are Equisetophyta and Lycopodiophyta, which include horsetails, scouring-rushes, quillworts, club-mosses, and spike-mosses. These "fern allies," produce spores like the true ferns (division Polypodiophyta), but look more like mosses, grasses, or rushes. The leaves of fern allies also lack a leafstalk (petiole) and the leaves are small (except in quillworts) and veinless or with only a single unbranched vein.

Information on specific and infraspecific taxa is set up in the following format: Number. Name author(s) [derivation of specific and infraspecific epithets]. VERNACULAR NAME. Habit; nativity (if exotic). Sporulating dates. Habitat data; highland provinces; relative abundance; [occurrence on Coastal Plain]. Conservation status. Wetland indicator status. Comments. Synonyms.

Introduced taxa are followed by a dagger (+). Species of conservation concern are followed by a star (*). The coded state ranks (ANHP 1994, 1996, 1997, 1999) are defined in Table 1. Wetland indicator status codes (Reed 1988) are defined in Table 2. Relative abundance is for occurrence in the study area and not for the whole state. Frequency of occurrence is defined as followed, ranging in descending order: common (occurring in abundance throughout), frequent (occurring throughout but not abundant), occassional (known in more than 50% of the region but in scattered localities), infrequent (known in less than 50% of the region in scattered localities), rare (known from only a few counties and restricted to a specific localities), and very rare (known from only a single or few populations; mostly narrow endemics, disjuncts, and peripheral taxa). Synonyms are from Mohr (1901)-- M; Small (1938)-- S; Radford et al. (1968)-- R; and Lellinger (1985)-- L. Suggested pronunciation, author(s), date of citation, common n ame, and derivations are provided after each genus.

Distribution maps are typically for 18 counties in the northeast region of Alabama. The maps are expanded to adjacent highland counties for taxa that are rare or peripheral. Key to symbols are as follows: Filled circle (*) = documented at Jacksonville State University herbarium; filled square (*) = documented at another herbarium; open circle (*) = reported in literature.

Positive or negative signs indicate a frequency toward higher (+) or lower (-) frequency of occurrence within a category.

Division I. EQUISETOPHYTA

Class 1. EQUISETOPSIDA

Order 1. EQUISETALES

1. EQUISETACEAE (Horsetail Family) [*]

(*.) Contributed in part by Terri L. Ballard

1. EQUISETUM {eh-quih-SEE-tum} Linnaeus 1753 * Horsetails * [Latin equus, horse, and seta, bristle; the bushy, branching pattern in some species somewhat resemble a horse's tail.]

Selected references: Hauke, R. L. 1961. A resume of the taxonomic reorganization of Equisetum subgenus Hippochaete, I. Amer. Fern J. 52: 29-123. Hauke, R. L. 1962. A resume of the taxonomic reorganization of Equisetum, subgenus Hippochaete, III. Amer. Fern J. 52: 57-63. Hauke, R. L. 1993. Equisetaceae. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 3+ vols. New York and Oxford. Vol. 2, pp. 76-84.

1. Stems dimorphic; sterile (non-sporulating) stems green (chlorophyllus) with whorls of branches at node; fertile (sporulating) stems brownish (achlorophyllus) unbranched; dark bands not present above or below nodal sheaths; strobili ("cones") 1-3 cm long, stalks 2-5 cm long, apex rounded, maturing early spring before sterile stems; aerial stems annual, flexuous E. arvense

1. Stems monomorphic: both fertile and sterile stems green and unbranched; dark bands present above and/or below nodal sheaths; strobili 0.5-1.5 cm long, stalks sessile to subsessile, apex mucronate, maturing late spring through summer; aerial stems evergreen, rigid E. hyemale

1. Equisetum arvense * Linnaeus [of cultivated fields]. FIELD HORSETAIL. Figure 1. Deciduous perennial. Sporulates March -- April. Low open woods and seepy areas; Cumberland Plateau, Ridge and Valley; rare; [Coastal Plain]. State Rank, S2. Wetland Indicator Status, FAC. It is only known from two places in northern Alabama; near Willett Spring in Calhoun County and Hughes Spring in Morgan County. Further south, it has been found growing on the banks of the Black Warrior River in Greene and Marengo counties. This species also occurs in Europe where the juice of the plant mixed with vinegar has been used as a remedy for ulcers and dropsy (Abbe 1981).

2. Equisetum hyemale Linnaeus [of winter] subsp. affine (Engelmann) Calder & Taylor [allied]. COMMON SCOURING-RUSH; TALL SCOURING-RUSH. Figure 2. Evergreen perennial. Sporulates May -- September. Low woods, streambanks, seepages, and pond margins; Interior Low Plateau, Cumberland Plateau, Ridge & Valley; occasional; [rarely Coastal Plain]. Wetland Indicator Status, FAC +. Equisetum hyemale occurs in Europe and Asia to northwestern China. The specific epithet is referring to the fact that this species persists throughout winter (Thieret 1980). Equisetum is considered poisonous to horses and other livestock when consumed with hay (Pohl 1955). Because of the hard sand-like substance (silica) found in the plant, the rush-like stems were used to polish pewter, scrub floors, and scour pots, pans and other utensils (Abbe 1981), hence the common names. The high silica content makes it a folk remedy for arthritis; the theory is silica smooths joints. Synonyms: Equisetum praealtum Rafinesque-S; Equisetum hyemale Linnae us var. affine (Engelmann) A. Eaton-- L.

DIVISION II. LYCOPODIOPHYTA

Class 1. ISOETOPSIDA

Order 1. ISOETALES

I. ISOETACEAE (Quillwort Family)

1. ISOETES {eye-so-EE-teez} Linnaeus 1753 * Quillworts * [Greek isos, equal, and etos, year; in reference to evergreen habit of some members of this genus.]

Selected references: Boom, B. M. 1982. Synopsis of Isoetes in the southeastern United States. Castanea 47: 38-59. Brunton, D. F., D. M. Britton, and T. F. Wieboldt. 1996. Taxonomy, identity, and status of Isoetes virginica (Isoetaceae). Castanea 61: 145-160. Taylor, W. C., N. T. Luebke, D. M. Britton, R. J. Hickey, and D. F. Brunton. 1993. Isoetes. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 3+ vols. New York and Oxford. Vol. 2, pp. 64-75. Taylor, W. C., T. H. Mohlenbrock, and J. A. Murphy. 1975. The spores and taxonomy of Isoetes butleri and I. melanopoda. Amer. Fern J. 65: 33-38.

1. Sporangium wall whitish (unpigmented); megaspores with honeycomb-like ridges (reticulate); plant aquatic (submerged or emergent), usually not associated with rock outcrops I. engelmannii

1. Sporangium wall pigmented with brown; megaspores with short ridges, wart-like projections (tuberculate to papillate) or almost smooth; plant terrestrial or amphibious in wet clay soils or on rock outcrops.

2. Plant of granite outcrops (rarely sandstone); sporangium brown and less than 6 mm long I. piedmontana

2. Plant of limestone outcrops or prairie-like habitats; sporangium mottled with brown and usually more than 6 mm long.

3. Leaf bases pale to brown; leaves usually less than 1 mm wide at middle; megaspores papillate, more than 450 um in diameter (0.50-0.65 mm average); plant of limestone outcrops I. butleri

3. Leaf bases usually shiny black; leaves usually more than 1 mm wide at middle; megaspores almost smooth to slightly tuberculate, less than 450 um in diameter (0.30-0.40 mm average); plant not associated with limestone outcrops I. melanopoda

1. Isoetes butleri * Engelmann [G. D. Butler, 1850-1910]. BUTLER's QUILLWORT; GLADE QUILLWORT. Figure 3. Deciduous perennial. Sporulates April -- October. Limestone outcrops; Interior Low Plateau, Cumberland Plateau; rare. State Rank, S2. Wetland Indicator Status, OBL. Named in 1878 by George Engelmann in honor of its discoverer, George Dexter Butler.

2. Isoetes engelmannii * A. Braun [G. Engelmann, 1809-1884]. ENGELMANN'S QUILLWORT. Figure 4. Evergreen perennial. Sporulates April -- October. Small streams, ponds, pools, and ditches; all highland provinces; infrequent; [Coastal Plain]. State Rank, previously S3 (ANHP 1994). Wetland Indicator Status, OBL. The most widely distributed quillwort in North America. Species was discovered in 1842 near St. Louis by George Engelmann, it was named in his honor by Alexander Braun. A recently described species, Isoetes appalachiana Brunton & Britton (Appalachian Quillwort) is very similar to I. engelmanni and occurs in Georgia and other states. It has a distinct cytology and spore morphology: I. appalachiana is tetraploid and its megaspores have a broken-reticulate pattern with ragged-crests; I. engelmanni is hexaploid and its megaspores have an unbroken reticulate pattern with generally smooth crests (Brunton and Britton 1997).

3. Isoetes melanopoda * Gray & Durieu [black-footed]. BLACK-FOOTED QUILLWORT; MIDLAND QUILLWORT. Figure 5. Deciduous perennial. Sporulates April -- October. Wetland Indicator Status, OBL. Wet fields and prairies; Interior Low Plateau, Ridge and Valley; rare. State Rank, S1. The specific epithet means black-foot" referring to the leaf bases, though they are not always black.

4. Isoetes piedmontana * (N. E. Pfeiffer) C. F. Reed [of the Piedmont]. PIEDMONT QUILLWORT; BLACK-BASED QUILLWORT. Figure 6. Deciduous perennial. Sporulates April -- October. Granite outcrops on the Piedmont Plateau; rare; very rarely sandstone outcrops on the Cumberland Plateau. State Rank, S2. Wetland Indicator Status, OBL. Found only in Alabama, Georgia, and South Carolina. Another granite outcrop species, the Black-spored Quillwort (I. melanospora Engelmann) was erroneously reported for Alabama. It can be distinguished from I. piedmontana by its blackish spores, leaves that are usually less than 8 cm long, and unpigmented sporangium wall.

Order 2. SELAGINELLALES

1. SELAGINELLACEAE (Spike-moss Family)

1. SELAGINELLA {sell-lah-jih-NELL-uh} Palisot de Beauvois 1805 * Spike-mosses * [Selago, an ancient name for Lycopodium, a genus resembling Selaginella, and Latin -ella, diminutive suffix.]

Selected references: Valdespino, I. O. 1993. Selaginella. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 3+ vols. New York and Oxford. Vol 2, pp. 38-63. Van Eseltine, G. P. 1918. The allies of Selaginella rupestris in the southeastern United States. Contr. Nat. Herb. 20: 159-172.

1. Sterile leaves dimorphic; arranged in 4 ranks on stem (2 medial, 2 lateral); lateral leaves appressed to stem.

2. Plant erect; large aerial branches above main stem; plant of moist areas (usually in close association with original ornamental planting) S. braunii

2. Plant prostrate; branching along length of main stem; plant of wet rocks, creek banks, and low wet areas where species occurs naturally S. apoda

1. Sterile leaves monomorphic; no distinct arrangement on stem.

3. Stems erect or ascending (seldom decumbent or creeping); roots usually well developed; megaspores with honey-combed surfaces S. arenicola subsp. riddellii

3. Stems decumbent or creeping (never erect); roots few to non-existent; megaspores nearly smooth S. rupestris,

1. Selaginella apoda (Linnaeus) Spring [footless]. MEADOW SPIKE-Moss. Figure 7. Deciduous perennial. Sporulates May -- September. Creek banks, sheltered wet areas, moist sandstone bluffs, and low wet areas; all highland provinces; frequent; [Coastal Plain]. Wetland Indicator Status, FACW+. The specific epithet refers to the lack of a well developed creeping rootsystem; thus the plant is "footless" (Snyder and Bruce 1986). Synonyms: Selaginella apus (Linnaeus) Spring-- M; Diplostachyum apodum (Linnaeus) Beauvois--S.

2. Selaginella arenicola * L. Underwood [of the sand] subsp. riddellii (Van Eseltine) R. M. Tryon [J. L. Riddell, 1807-1865]. SAND SPIKE-MOSS; RIDDELL'S SPIKE-MOSS. Figure 8. Persistent perennial. Sporulates May -- September. Dry sandy areas, sandstone and granite outcrops; Cumberland Plateau, Ridge and Valley, Piedmont Plateau; rare; [Coastal Plain]. State Rank, S2. Wetland Indicator Status, UPL. The range of S. arenicola subsp. arenicola includes Florida and southern Georgia. Van Eseltine named this species in honor of John Leonard Riddell, inventor of the binocular microscope and author of Catalogus Florae Ludovicianae published in 1852.

3. Selaginella braunii + Baker [E. M. Braun, 1889-1971]. BRAUN'S SPIKE-Moss; TREELET SPIKE-Moss. Figure 9. Persistent perennial; native to China. Sporulates May -- September. Moist forested and open areas (escapes from ornamental plantings); Cumberland Plateau; very rare. Wetland Indicator Status, NI. This species is an escape from cultivation and is doubtfully well-established in our range. Named in honor of Emma Lucy Braun, an American ecologist.

4. Selaginella rupestris * (Linnaeus) Spring [of rocks]. ROCK SPIKE-Moss; DWARF SPIKEMOSS. Figure 10. Persistent perennial. Sporulates May -- September. Flat sandstone and granite outcrops, sandy areas; Cumberland Plateau, Ridge and Valley, Piedmont Plateau; rare. State Rank, S2S3. Wetland Indicator Status, UPL.

Class 2. LYCOPODIOPSIDA

Order 1. LYCOPODIALES

1. LYCOPODIACEAE (Clubmoss Family)

Selected reference: Wagner, W. H. and J.M. Beitel. 1993. Lycopodiaceae. In: Flora of North America Editorial Committee, eds, 1993+. Flora of North America. 3+ vols. New York and Oxford. Vol. 2, pp. 18-37.

1. Strobili occurring along the length of the stem; sporophylls (fertile leaves) similar to sterile leaves Huperzia

1. Strobili occurring terminally along upper one-third of stem; sporophylls distinctly different from sterile leaves.

2. Main erect stem unbranched; plants pale green, usually dying back during winter; plants primarily of wetland communities Lycopodiella

2. Main erect stem branching into several finger- or fan-like branchlets; plants shiny green, evergreen; plants primarily of upland communities.

3. Strobili sessile; leafy branches usually 5 to 8 mm wide Lycopodium

3. Strobili with distinct peduncles; leafy branches usually 3 mm or less wide Diphasiastrum

1. DIPHASIASTRUM (dye-phase-ee-ASS-strum} Holub 1975 * Ground-cedars; Ground-pines * [Incomplete likeness to the genus Diphasium.]

Selected references: Holub, J. 1975. Diphasiastrum, a new genus in Lycopodiaceae. Preslia 14: 97-100. Wagner, W. H. and J.M. Beitel. 1993. Diphasiastrum. In: Flora of North America Editorial Committee, eds, 1993+. Flora of North America. 3+ vols. New York and Oxford. Vol. 2, 28-32.

1. Rhizomes close to ground surface (usually within 1 cm of surface or within leaf litter); underside 2 rows of sterile leaves shorter than 2 upperside rows; sterile leaves green D. digitatum

1. Rhizomes well below ground surface (usually 1-6 cm below surface); underside 2 rows of sterile leaves similar in size to 2 upperside rows; sterile leaves bluish (glaucous) D. tristachyum

1. Diphasiastrum digitatum * (Dillenius ex A. Braun) Holub [finger-like]. RUNNING GROUND-PINE; SOUTHERN RUNNING-PINE; CROWFOOT CLUB-MOSS. Figure 11. Evergreen perennial. Sporulates April - August. Mixed upland woods, roadsides, powerline rights-of-way, and other open areas; all highland provinces; occasional. Species of Special Concern (Freeman et al. 1979). Wetland Indicator Status, FACU. The specific epithet of this species refers to the finger-like appearance of the branchlets. Used as a decoration in wreaths. Often confused with the similar Northern Running-pine, Diphasiastrum complanatum (Linnaeus) Holub. Synonyms: Lycopodium flabelliforme (Fernald) Blanchard--S. R; Lycopodium digitatum Dillenius ex A. Braun--L.

2. Diphasiastrum tristachyum * (Pursh) Holub [three-spiked] GROUND-CEDAR; SLENDER GROUND-PINE; DEEP-ROOT CLUB-MOSS. Figure 12. Evergreen perennial. Sporulates April - August. Mixed upland woods, roadsides, powerline rights-of-way, and other open areas; Cumberland Plateau; very rare. State Rank, SR. Wetland Indicator Status, UPL. The state rank was based on an old record by E. W. Graves who collected it in 1917 on top of Sand Mountain near Higdon in Jackson County. It has recently been collected from Jackson County. Diphasiatrum digitatum and D. tristachyum are known to interbreed to form the infertile hybrid Diphasiastrum X habereri (House) Holub. Synonym: Lycopodium tristachyum Pursh--S, R, L.

2. HUPERZIA {hew-PURR-zee-uh} Bernhardi 1801 * Fir-mosses; Hanging Club-mosses * [For Johann Peter Huperz, a German fern horticulturist.]

Selected references: Wagner, W. H. and J.M. Beitel. 1993. Huperzia. In: Flora of North America Editorial Committee, eds, 1993+. Flora of North America. 3+ vols. New York and Oxford. Vol. 2, pp. 20-24. Waterway, M. J. 1986. A reevaluation of Lycopodium porophilum and its relationship to L. lucidulum (Lycopodiaceae). Sys. Bot. 11: 263-276.

1. Sterile leaves widest above middle, leaves of varying lengths resulting in a tufted (shaggy) appearance, leaves with toothed upper margins; main stem frequently branching so that plant often forms dense clumps; plant mainly of soil in rich rocky woods... H. lucidula

1. Sterile leaves widest below middle, leaves of essentially same length, leaves typically with entire margins (sometimes toothed); main stem usually branching only 2 to 3 times; plant growing directly on acidic rocks H. porophila

1. Huperzia lucidula * (Michaux) Trevisan [somewhat shining]. SHINING CLUB-Moss; SHINING FIR-Moss. Figure 13. Evergreen perennial. Sporulates June -- September. Rich rocky woods, plant mainly grows directly from soil; Cumberland Plateau; rare. State Rank, S2. Wetland Indicator Status, FACW. The common names are derived from the shiny appearance of the leaves. This species and Huperzia porophila form a sterile hybrid known as Huperzia X bartleyi (Cusick) Kartesz & Gandhi. Synonym: Lycopodium lucidulum Michaux-- S, R, L.

2. Huperzia porophila * (F. Lloyd & L. Underwood) Holub [lover of stone]. ROCK CLUB-MOSS; CLIFF CLUB-MOSS. Figure 14. Evergreen perennial. Sporulates June -- September. Rich rocky acidic woods, plant usually grows directly on rock outcroppings; Cumberland Plateau; rare. State Rank, S1. Wetland Indicator Status, UPL. This species is known to occur in Winston County, Alabama. Synonyms: Lycopodium porophilum F. Lloyd & L. Underwood-- M, S, R, L.

3. LYCOPODIELLA {lye-koh-POH-dee-ell-uh} Holub 1964 * Bog Club-mosses * [A diminutive form of Lycopodium.]

SELECTED REFERENCE: Wagner, W. H. and J.M. Beitel. 1993. Lycopodiella. In: Flora of North America Editorial Committee, eds, 1993+. Flora of North America. 3+ vols. New York and Oxford. Vol. 2, pp. 34-37.

1. Horizontal stems creeping flat on ground (rooting throughout on ventral surface), leaves entire to slightly toothed; strobili (cones) only slightly wider than adjacent stem (1-2 mm wider); leaves of upright stem and sporophylls appressed L. appressa.

1. Horizontal stems arching (only rooting where touching ground), leaves strongly toothed; strobili distinctly wider than adjacent stem (3-6 mm wider); leaves of upright stem and sporophylls spreading to ascending L. alopecuroides.

Note: Lycopodiella prostrata (Harper) Cranfill [Lycopodium prostratum Harper], Feather-stem Club-moss, Prostrate Club-moss, occurs in Coastal Plain portions of counties adjacent to our study area. It is very similar to L. appressa, the horizontal stems creep flat on the ground, but the leaves are strongly toothed and the leaves of upright stem and sporophylls are not appressed.

1. Lycopodiella alopecuroides (Linnaeus) Cranfill [resembling Alopecurus, foxtail grass]. FOXTAIL CLUB-MOSS; FOXTAIL BOG CLUB-MOSS. Figure 15. Deciduous perennial. Sporulates July - September. Wet areas including ditches, emergent wetlands, pond margins, and forested wetlands; Ridge and Valley; rare; [chiefly Coastal Plain]. Wetland Indicator Status, OBL. This species is believed to be one of the first American members of this group examined by Linnaeus (Thieret 1980). Synonym: Lycopodium alopecuroides Linnaeus--M, S, R, L.

2. Lycopodiella appressa (Chapman) Cranfill [appressed]. SOUTHERN CLUB-MOSS; APPRESSED BOG CLUB-MOSS; TIGHT-LEAF CLUB-MOSS. Figure 16. Deciduous perennial. Sporulates July - September. Wet areas including ditches, emergent wetlands, pond margins, and forested wetlands; Cumberland Plateau; rare; [chiefly Coastal Plain]. Wetland Indicator Status, OBL. The specific epithet refers to the appressed sporophylls. This species is known to hybridize with Lycopodiella alopecuroides and other members of this genus. No effort is made to separate the hybrids within this treatment. Synonyms: Lycopodium adpressum (Chapman) Lloyd & Underwood--M; Lycopodium appressum (Chapman) F. Lloyd & L. Underwood--S, R, L.

4. LYCOPODIUM {lye-koh-POH-dee-um} Linnaeus 1753 * Club-mosses * [Greek lykos, wolf, and pous, podes, foot; in reference to the resemblance of a wolf's paw].

Selected reference: Wagner, W. H. and J.M. Beitel. 1993. Lycopodium. In: Flora of North America Editorial Committee, eds, 1993+. Flora of North America. 3+ vols. New York and Oxford. Vol. 2, pp. 25-28.

1. Lycopodium obscurum * Linnaeus [obscure]. PRINCESS-PINE; TREE CLUB-MOSS. Figure 17. Evergreen perennial. Sporulates August - February. Rich wooded slopes and floodplains; Cumberland Plateau; rare. State Rank, S1. Wetland Indicator Status, FACU-. The specific epithet obscurum refers to the deep, "hidden" rhizome (Snyder and Bruce 1986). Foliage from this species has been used as a decoration during the Christmas season (Dean 1969), but should not be collected due to its rarity. The common name is derived from its tree-like resemblance.

REFERENCES CITED

Abbe E. 1981. The Fern Herbal. Comstock Publishing Associates. Ithaca and London.

Alabama Natural Heritage Program [ANHP]. 1994. Vascular Plant Inventory Tracking List, April edition. Montgomery, Alabama.

Alabama Natural Heritage Program. 1996. Species Inventory List. Montgomery, Alabama.

Alabama Natural Heritage Program. 1997. Inventory List of Rare, Threatened, and Endangered Plants, Animals, and Natural Communities of Alabama. Montgomery, Alabama.

Alabama Natural Heritage Program. 1999. Inventory List of Rare, Threatened, and Endangered Plants, Animals, and Natural Communities of Alabama. Montgomery, Alabama.

Brunton, D. F. and D. M. Britton. 1997. Appalachian quillwort (Isoetes appalachiana, sp. nov. ; Isoetaceae), a new pteridophyte from the eastern United States. Rhodora 99: 188-133.

Dean, B. E. 1969. Ferns of Alabama. Southern University Press, Birmingham.

Freeman, J. D., A. S. Causey, J. W. Short, and R. R. Haynes. 1979. Endangered, threatened, and special concern plants of Alabama. Department of Microbiology, Agricultural Experiment Statioin. Departmental Series no. 3, Auburn University.

Lellinger, D. B. 1985. A Field Manual of the Ferns and Fern-allies of the United States and Canada. Washington, D. C.

Mohr, C. 1901. Plant Life of Alabama. Cont. U. S. Nat. Herb. 6.

Pohl, R. W. 1955. Toxicity of ferns and Equisetum. Amer. Fern J. 45: 95-97.

Radford, A. E., H. E. Ahles, and C. R. Bell. 1968. Manual of the Vascular Flora of the Carolinas. The University of North Carolina Press.

Reed, P.B. 1988. National List of Plant Species that Occur in Wetlands: Southeast (Region 2). U.S. Fish and Wildlife Service, Biological Rep. 88 (24), 244 p.

Small, J. K. 1938. Ferns of Southeastern States. Published by author. New York.

Snyder, L. H. and J. G. Bruce. 1986. Field Guide to the Ferns and Other Pteridophytes of Georgia. The University of Georgia Press.

Theiret, J. W. 1980. Louisiana Ferns and Fern Allies. Lafayette Natural History Museum in conjunction with the University of Southwest Louisiana. Lafayette, Louisiana.
 Definition of state ranks.
Code Definition
S1 Critically imperiled in Alabama because
 of extreme rarity or because of some
 factor(s)
S2 Imperiled in Alabama because of rarity
 or because of some factor(s) making it
 very vulnerable to extirpation from the
 state.
S3 Rare or uncommon in Alabama.
S4 Apparently secure in Alabama, with many
 occurrences.
S5 Demonstrably secure in Alabama and
 essentially "ineradicable" under present
 conditions.
SH Of historical occurrence, perhaps not
 verified in the past 20 years, and
 suspected to be still extant.
SR Reported, but without persuasive
 documentation which would provide a
 basis for either accepting or rejecting
 the report.
SU Possibly in peril in Alabama, but status
 uncertain.
S? Not ranked to date.
 Definition of wetland indicator codes.
Code Status
OBL Obligate Wetland Species
FACW Facultative Wetland Species
FAC Facultative Species
FACU Facultative Upland Species
UPL Obligate Upland Species
NI No Indicator Status
Code Probability of Occurrence
OBL Occurs with estimated 99% probability in
 wetlands.
FACW Estimated 67%-99% probability of
 occurrence in wetlands, 1%- 33%
 probability in nonwetlands.
FAC Equally likely to occur in wetlands and
 nonwetlands (34%-66% probability).
FACU Estimated 67%-99% probability of
 occurrence in nonwetlands, 1%-33%
 probability in wetlands.
UPL Occurs with estimated 99% probability in
 uplands.
NI Insufficient information available to
 determine an indicator status.
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Author:Spaulding, Daniel D.; Whetstone, R. David; Ballard, J. Mark
Publication:Journal of the Alabama Academy of Science
Geographic Code:1USA
Date:Oct 1, 2000
Words:6224
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