Tennessee Naturalist Program

Forbs, Ferns, Mosses, and More Herbaceous Plants and Fungi of Tennessee

Enhanced Study Guide

1/2019 Tennessee Naturalist Program www.tnnaturalist.org

Inspiring the desire to learn and share Tennessee’s nature

These study guides are designed to reflect and reinforce the Tennessee Naturalist Program’s course curriculum outline, developed and approved by the TNP Board of Directors, for use by TNP instructors to plan and organize classroom discussion and fieldwork components and by students as a meaningful resource to review and enhance class instrucon.

This guide was compiled specifically for the Tennessee Naturalist Program and reviewed by experts in these disciplines. It contains copyrighted work from other authors and publishers, used here by permission.

No part of this document may be reproduced or shared without consent of the Tennessee Naturalist Program and appropriate copyright holders.

Unless otherwise noted, all photographs are by Margie Hunter.

2 Forbs, Ferns, Mosses, and More Herbaceous Plants and Fungi of Tennessee

Objecves -- Examine fungi (mushrooms and lichens) and plants (mostly herbaceous plants, including nonvascular and spore-producing vascular plants). Learn the structure behind taxonomic classificaon, nomenclature, and organism morphology for family, genus, and species idenficaon via dichotomous keys. Understand the mechanisms behind invasive pest species and their ecological disrupons.

Time -- Minimum 4 hours -- 2 classroom, 2 field.

Suggested Materials (* recommended but not required, ** TNP flash drive) • Fern Finder, Anne Hallowell and Barbara Hallowell * • Wildflowers of Tennessee, the Ohio Valley, and the Southern Appalachians, Dennis Horn and Tavia Cathcart * • A Field Guide for the Idenficaon of Invasive Plants in Southern Forests, James H. Miller, Erwin B. Chambliss, and Nancy J. Loewenstein (U.S. Forest Service Pub. GTR-SRS-119) ** • “Invasive Exoc Pest Plants in Tennessee – 2009” (TN-EPPC) ** • “Tennessee’s Nave Plant Alternaves to Exoc Invasives” (TN-EPPC) ** • Forbs, Ferns, Fungi, and More Enhanced Study Guide, TNP **

Expected Outcomes -- Students will gain a basic understanding of 1. fungi and lichens, general ecology and idenficaon 2. plant kingdom organizaon and evoluonary history 3. classificaon and nomenclature 4. nonvascular mosses, liverworts, and hornworts, general ecology and idenficaon 5. pteridophytes, general ecology and idenficaon 6. spermatophytes: gymnosperms, angiosperms, monocots, and dicots 7. flowering plant morphology (flower/foliage) and family characteriscs 8. dichotomous keys and their use 9. invasive pest species, their characteriscs, mechanics of invasion, disrupons

3 Plants and Fungi Curriculum Outline

I. Fungi -- Kingdom Eumycota (Mycology) A. Roles and relaonships 1. heterotrophs 2. relaonships 3. decomposers 4. mycorrhizae a. endomycorrhizae b. ectomycorrhizae c. other associaons 5. fungus anatomy B. Phyla 1. Ascomycota (sac fungi) 2. Basidiomycota (club fungi) 3. other phyla and slime molds C. Reproducon and spore dispersal D. Idenficaon 1. general fruing body forms 2. field idenficaon characteriscs

II. Lichens A. Symbioc relaonship 1. mycobionts (fungi) 2. photobionts (algae) B. Common forms 1. crustose 2. foliose 3. frucose 4. squamulose C. Ecological roles and uses 1. soil formaon 2. bioindicator 3. wildlife 4. cultural uses D. Idenficaon

4 III. Plants -- Kingdom Plantae (Botany) A. Plant funcon and phylogenec evoluon 1. photosynthesis -- autotrophs 2. species divergence – vascular ssue, seeds, flowers 3. sexual reproducon B. Classificaon 1. Kingdom, Phylum or Division, Class, Order, Family, Genus, Species 2. nomenclature 3. species, genus, and family (common traits) C. Nonvascular plants 1. ecology of bryophytes 2. hornworts (Anthocerotophyta) 3. liverworts (Marchanophyta or Hepacophyta) 4. mosses (Bryophyta) D. Vascular plants 1. Ferns (Pteridophyta) a. fern allies (clubmosses) b. true ferns c. fern life cycle d. fern idenficaon 2. Spermatophyta a. gymnosperms -- the conifers (Coniferophyta) b. angiosperms -- the flowering plants (Magnoliophyta) i. monocotyledon (Liliopsida) ii. dicotyledon (Magnoliopsida) c. reproducon d. plant morphology 3. Habitat, role in idenficaon 4. Plant ecology and uses

IV. Dichotomous Keys A. Guidelines for use

5 V. Invasive Pest Species A. Movement of species from nave range B. Mechanics of invasion 1. loss of checks and balances -- compeon, climate, predaon C. Non-nave invasive species definion D. Disrupons to nave biological systems E. Invasive plant characteriscs F. Scope and impact

VI. Management A. Natural areas B. Invasive plants C. What you can do

VII. Resources

VIII. Review Quesons

Appendix A: Common mosses, liverworts, and hornworts Appendix B: Botanical Lan pronunciaon guide

6 I. Fungi -- Kingdom Eumycota

Roles and Relaonships

The study of fungi is mycology, myco- from the Greek mukes or mykes meaning “fungus.” Fungi are heterotrophs, organisms that cannot manufacture their own food and therefore, must feed on other organisms. They receive needed nutrion in two primary ways -- as decomposers of nonliving organic maer and through symbioc relaonships with other living organisms, both as harmful parasites and as mutualisc partners. At one me, fungi were included in the plant kingdom. Beer understanding of these unique organisms demonstrated the need for a separate kingdom. Certain characteriscs of fungi appear more closely related to animals than plants. They have chin in cell walls like insects rather than the cellulose of plants, and they store food as glycogen like animals rather than starch like plants.

Earthstar (Geastrum sp.), saprotrophic

Decomposers A large number of fungi are saprotrophs, feeding on dead organic maer, and serve a crical role as decomposers in ecosystems. Fungi must digest their food before ingesng it. They excrete enzymes to break down complex organic molecules for absorpon through fungal cell walls. This funcon plays an important role in releasing and cycling nutrients through the enre system.

7 As a whole, fungi can break down virtually any organic substance. Bacteria do most of the work decomposing animals, but there are fungi capable of breaking down components such as collagen and keran. Fungi are essenal, however, in the decomposion of plant material. They are able to break down the toughest organic compounds, including lignin, a complex polymer in the cells of woody plants that enables transport of fluids and provides structural support. Fungi possessing enzymes to degrade both cellulose and lignin are termed “white-rot” fungi. The residual ssue is fibrous and pale in color. Those that only break down cellulose are referred to as “brown-rot,” leaving brown, blocky fragments that disintegrate. Certain species target foliage decomposion, their spores oen aaching to leaves before they fall. There are fungi quite specific to a parcular organic substrate, such as the husks of walnuts and hickories or the fruit cones of magnolias. Aquac fungi are important to the well being of stream fauna. Autumn leaves and spring runoff load streams with plant material, that need to be ‘condioned’ by organisms, including fungi, to make these organic materials more palatable. In addion, the fungi mycelia and spores are considered nutrious food. Animal waste (dung) is largely plant material that has not been digested. Dung-loving or coprophilous fungi play an important role here too. Spores of these fungi are released in a manner to maximize placement for consumpon by herbivores, as the spores of many species will only germinate aer passing through an animal’s digesve tract.

Yellow Patches (Amanita flavaconia), mycorrhizal

8 Symbioc Relaonships In symbioc relaonships with plants, fungi may be parasites, harming but usually not killing other organisms, or mycorrhizal partners in mutually beneficial relaonships. As parasites, fungal smuts and rusts that infect plants cause harm in various ways, including interrupon of primary processes, e.g., sexual funcon of a flower in the case of smuts. Some parasic fungi are pathogenic, aacking living ssue to play a roll, direct or indirect, in the organism’s death, such as white-rot fungus Honey Mushroom (Armillaria) in trees, Chytrid fungus in amphibians, and “insect destroyer” fungi (Entomophthora) with flies and other invertebrates. There are even fungi that parasize other fungi -- Witches’ Buer (Tremella mesenterica) and Lobster Mushroom (Hypomyces lacfluorum). On the other hand, mutualisc associaons between plant roots and fungi are extremely important to the health of a majority of plants and the ecosystems they support. Fungi forming these mutualisc associaons are called mycorrhizae (“fungus root”). The benefits to the plant are many. First and most obvious, fungi hyphae absorb water and soil nutrients for the plant -- calcium, copper, magnesium, molybdenum, zinc, potassium, and parcularly phosphorus, a liming nutrient in many soils. Hyphae are able exploit a far greater volume of soil for these essenal elements than plant roots alone. Fungi are also able to store some of these mineral nutrients (again, parcularly phosphorus) and make them available for plants during periods of acve growth or nutrient deficiency. Mycorrhizal associaons help plants in deficient soils grow beer -- soils that are less ferle, have wider pH variances, or are polluted. Plants are beer able to survive transplant shock, withstand environmental stresses like drought, and resist soil-borne pathogens. In exchange for these valuable services, plants translocate sugars from their roots to the fungus where the food is converted to fungal carbohydrates. Esmates of the amount of photosynthec producon plants share with their fungal partners reach 10% or more. There are two primary types of mycorrhizae -- endomycorrhizae and ectomycorrhizae. Endomycorrhizal fungi hyphae penetrate the root’s cells and form highly branched structures called arbuscules that oen fill the cells’ interior, yet they do not interfere with the vascular funcon of the root. Water, nutrients, and plant sugars are exchanged in these arbuscules. Some (but not all) of the fungi produce another structure called a vesicle for storage. Endomycorrhizal fungi produce large, asexual spores in the soil. Considering that approximately 80 to 85 percent of all plants (300,000, especially in grasslands and tropical ecosystems) form this type of fungal associaon, there are surprisingly few species (130) of endomycorrhizal fungi idenfied, and all of them are obligates -- they cannot grow outside the associaon with plant roots. [Kendrick, 2000] Ectomycorrhizal fungi form a dense network around plant roots called a mantle or sheath. Hyphae on the outside of the sheath extend into the soil. Hyphae on the inside enter the root and form a network around, but do not penetrate, the root’s cells called the Harg net. Most ectomycorrhizae are basidiomycetes (fungi that produce most forest mushrooms); a few are ascomycetes. Only about 10 percent of all plants, some 2,000 species, form these associaons, yet they are the important tree species of temperate and boreal forests. One tree may have several ectomycorrhizal partners at any given me, and the species of fungi may

9 change as the tree ages. Trees may be linked underground in a vast network of mycorrhizal hyphae. There are a handful of specialized mycorrhizal associaons. Some basidiomycetes form a unique blend of the two types described above in the cells of plants in the Heath family (Ericaceae). Non-photosynthesizing plants in the genus Monotropa, form associaons with ectomycorrhizal fungi, taking some of the food the fungus gets from its tree partner. Mycorrhizal associaons have been found with nonvascular liverwort and hornwort plant species. Certain fungi form relaonships with orchids, both seeds and plants. Orchid seeds have lile to no food reserves and must establish a fungal associaon to grow. Microscopic fungi called endophytes have been found living inside plant leaves and stems. While their exact funcons and relaonships are not fully understood, they do not seem to harm the plant and may confer resistance to insect predaon and deter herbivore browsing. Nearly 95 percent of all plants form some sort of mutualism with a mycorrhizal partner. Mycorrhizae are found in all terrestrial ecosystems, comprising one of the largest biomass components and serving as the largest consumer of net primary producvity. On any given plant, its mycorrhizal partner may colonize up to 95 percent of its primary root length. Mycorrhizae are just beginning to be appreciated for their roles in increased carbon fixaon and total seasonal carbon gain as evidenced in increased plant growth and larger root systems, their importance in successful reclamaon and restoraon of highly disturbed sites (mine spoils and burned areas), their stabilizaon of soil through hyphae binding parcles, and their protecon at the soil-root interface to reduce establishment of pathogenic organisms, including the producon of secondary metabolites with an-microbial properes.

Mushrooms and Toadstools For some mycologists, the term “mushroom” is generically applied to all macro-fruing fungi. Others restrict usage to the fungi that produce gilled sporocarps. Technically, the more correct term for gilled fruing bodies would be “agaric.” Historically, general cultural usage of mushroom was oen applied only to edible fungi. Another informal term is “toadstool,” which has oen been used to disnguish poisonous species and is associated with the classic cap and stalk fungal fruing form. A Brish arcle* notes “toadstool” dates to the fourteenth century. Various iteraons include toad’s cap, toad’s meat, toad’s cheese, and toad’s paddock. Somemes “frog” appeared in place of “toad.” The pairing of a toad and fungus likely resulted from common belief that all toads were poisonous. Other associaons that may have influenced the pairing include warty mushroom caps resembling toad skin, dung (toads and excrement are linked in some languages and fungi oen grow on dung), the ancient belief that both amphibians and fungi come from slime, and a tendency to associate toads and some fungi with the devil. Toadstool has no precise meaning today.

* “The Word ‘Toadstool’ in Britain,” by Tony Baker, Mycologist 4:25-29, 1990. (www.fungi4schools.org)

10 Anatomy of Fungi: The Parts of a Mushroom

“A mushroom is the fruing body (sporocarp) that arises from the larger fungal organism, the vegetave body or mycelium (a mass of branching threadlike filaments called hyphae), which is typically concealed beneath the soil or within decaying wood or other substrate. “A mushroom begins as an immature form called a buon. Depending on the species, the buon may inially be enrely surrounded by a membranous structure known as the universal veil. As the mushroom within expands, it stretches and tears the universal veil, oen leaving remnants on the cap. These remnants are referred to as patches or warts. There may also be a cup-like remnant of the universal veil called a volva around the base of the mushroom stalk. Most mushrooms lack a universal veil and have neither patches nor warts nor a volva. “The typical mature mushroom has a cap and a stalk. The stalk may be aached to the cap’s center, off its center, or at its side. On the cap’s underside, there may be gills, which are radiang, blade-like structures upon which spores are produced. In place of gills, there may be teeth or spines or tubes (the open end of each tube is called a pore). The tubes are packed closely together; their collecve pores are known as the pore surface. Both teeth or spines and tubes serve the same basic reproducve funcon as gills. “In some species, the underside of the immature cap is covered by a piece of ssue stretching from the cap’s edge or margin, to the stalk. This ssue, the paral veil (not shown in illustraon), covers and protects the developing gills or tubes. As the mushroom cap expands, the paral veil tears, oen leaving remnants on the cap margin or adhering to the stalk, where it forms a skirt or ring.”

Alan E. Bessee, William C. Roody, Arleen R. Bessee, Dail L. Dunaway, Mushrooms of the Southeastern United States (Syracuse, NY: Syracuse University Press, 2007), pages 2-3. Text and drawing used by permission.

11 Phyla in the Kingdom Fungi

Primary Phyla Associated with Macrofungi in Terrestrial Habitats Ascomycota (Sac Fungi) -- Represenng the largest phylum with over 30,000 described species, ascomycetes are very diverse. However, all are characterized reproducvely by the typical formaon of eight spores in a microscopic saclike structure called an ascus. Asci form small embedded groups or large layers on ferle surfaces. Sac fungi include valuable species like unicellular yeasts that vegetavely reproduce through budding, bothersome species such as sooty molds that grow on sugary wastes of aphids in the garden, deadly pathogens responsible for Dutch elm disease, chestnut blight, and beech bark disease, and economically damaging apple scab and gray mold on so fruits. The macrofungi ascomycetes are an interesng mix. Cordyceps spp. send up a fruing body from the parasized larvae of insects, buried pupae of moths, and false truffles. Blue-green wood stain (Chlorociboria aeruginascens and C. aeruginosa) discolors infected dead wood. Forest species include Jelly Clubs (Leoa), Earth Tongues (Microglossum, Trichoglossom, and Geoglossum), cup fungi (Common Brown Cup, Peziza badioconfusa, Scarlet Cups, Sarcoscypha, Eyelash Cup, Scutellinia scutellata), and Dead Man’s Fingers (Xylaria polymorpha). Dead Man’s Fingers start out white, coated with asexual spores called conidia. When those fall off, the fungus is black and releases sexual spores. Related Carbon Balls (Daldinia concentrica) are round, brown, hard structures on dead wood and look like charcoal inside when broken. The black interior is zoned in concentric bands. The fruing bodies of some fungi grow underground, hypogeous. Truffles are the best known examples. Both the Ascomycota and Basidiomycota have subterranean species. One of the ascomycete species of False Truffle (Elaphomyces) is mycorrhizal with Red Spruce and a food source for Northern Flying Squirrels. Hypogeous sporocarps emit an odor when mature encing animals to dig them up. Spores are spread aer passing through their digesve tracts. The most sought ascomycetes (aer truffles) are morels. Unlike most mushrooms, morels fruit in spring, producing sporocarps with disncve ridges and pits, Yellow or Common Morel (Morchella esculentoides) and Black Morel (M. angusceps). Though delicious, raw morels contain toxins and are considered poisonous. They should only be eaten aer thorough cooking and should not be consumed with alcohol. Some species of False Morel, also spring fruing, are highly toxic and deadly. The resemblance is superficial as False Morels are usually larger and brown or reddish-brown with convoluted caps as opposed to the yellow or black conical caps with ridges and pits found on true morels. Members of this phylum comprise the vast majority of those fungi that form lichen associaons with algae. Lichens will be covered in the next secon.

Basidiomycota (Club Fungi) -- Basidiomycetes form a club-shaped structure called a basidium typically with four small extensions, each containing a spore. The club fungi have three main subphyla. Two of them, comprising less than a third of the 29,000 described species,

12 contain plant parasites and pathogens, the rusts and smuts. The third subphyla consists of the typical common “mushrooms” of which there are two primary groups. In one group, the ferle layer is disnct and exposed allowing spores to be forcibly discharged. It includes the agarics, so-bodied sporocarps with gills (the mushrooms) or a layer of tubes (the boletes) and the aphyllophorales, sporocarps without these two features (chanterelles, tooth fungi, and coral fungi) or that look substanally different (hard bracket or shelf polypores). The other group has the ferle mass of spores enclosed within a sterile layer of hyphae -- pualls, earthstars, bird’s nests, and snkhorns. There are a handful of isolated taxa that are hard to place in any group, including Witches’ Buer (Tremella mesenterica), Wood Ears (Auricularia auricula), and Jelly Tongue (Pseudohydnum gelanosum). Basidiomycetes (and ascomycetes) are good at digesng resistant materials like cellulose, collagen, and parcularly lignin. A handful of basidiomycetes make up the remaining lichenized fungi. While many taxa in this phylum are saprotrophic, it also features those fungi that form ectomycorrhizal associaons with forest trees in the temperate and boreal zones.

Other Phyla True and related fungi taxa have undergone dramac reorganizaon in recent years from DNA analysis. The number of phyla within the fungi kingdom is sll in flux and some taxa may change phyla. • Chytridiomycota - Chytrids, mostly unicellular and microscopic, have mole spores with single flagellum. There are saprotrophs, a few parasites, and one deadly pathogen that causes the infecous disease chytridiomycosis in amphibians. It affects their skin, impeding the osmoc funcon, and may release toxins. • Blastocladiomycota - Primarily saprotrophic fungi feeding on dead organic maer in soil, mud, and water, these fungi inhabit mostly aquac habitats and have mole spores. • Glomeromycota - This recent phylum contains the arbuscular mycorrhizal fungi (endomycorrhizae), which are usually obligate symbionts (cannot grow without the host plant) and are associated with approximately 80% of all plants. This associaon dates to 400 million years ago. Many species in this phylum are not known to reproduce sexually. • Zygomycota - A highly diverse group ecologically, including saprotrophs and pathogens disnguished by unique reproducon through zygospores, this phylum is undergoing much change, perhaps being dismantled, and many of its species are now considered part of the Glomeromycota, including Bread Mold.

Slime Molds (Moulds) At one me slime molds represented four separate phyla in the Kingdom Eumycota. Since these organisms are unrelated to true fungi, they are now in the Kingdom Prosta though they are typically sll studied by mycologists. The largest and best known group of slime molds (formerly Myxomycota or myxomycetes) is today known as the Class Myxogastria. The other classes of slime molds are largely microscopic.

13 Myxogastria are plasmodial slime molds that feed on bacteria and other microorganisms living in dead plant material and are common on the forest floor, logs, and soil. They have two trophic (feeding) stages and a reproducve stage. The first trophic stage involves germinaon of amoeboid cells (haploid) from spores. Each cell has a single nucleus and feeds and divides to build up large populaons. The molds are nearly impossible to detect in this stage. With the second feeding stage, two haploid cells join as a diploid zygote that undergoes repeated nuclear division forming a mass of protoplasm with no cells walls and thousands of nuclei called a plasmodium. It resembles a glistening, veinlike blob and can creep, up to an inch per hour, in search of food. In experiments, plasmodia have successfully negoated mazes to find food. Eventually, it develops one or more fruing bodies with spores, the reproducve stage. The plasmodium (most species are small, but some are quite large) and the sporocarps are the visible stages in the life cycle of a slime mold. Species idenficaon is based on morphological features of the sporocarp and microscopic examinaon of the spores. Two regularly seen slime molds are Dog Vomit Slime Mold (Fuligo sepca) oen found on garden wood mulch in the plasmodial stage and Wolf’s Milk (Lycogala epidendrum) found on forest logs as sporocarps.

Wolf’s Milk or Toothpaste Slime Mold (Lycogala epidendrum)

14 Reproducon in Fungi

Fungi are very diverse organisms. Different groups employ different processes for reproducon. There are a few species in which sexual reproducon has never been observed. Others engage in both sexual and asexual reproducon, usually at different mes. These two reproducve phases of the same fungus have somemes been confused as belonging to two different species. For the most part, fungi reproduce asexually, somemes through fragmentaon of the mycelium, and largely through the mitoc division of haploid spores (conidia). When environmental condions are good, this is the preferred method, allowing each fungus to more rapidly colonize a favorable locaon. Adverse environmental condions may prompt sexual reproducon. In the more complex fungi, like the mushroom-producing ascomycetes and basidiomycetes, hyphae seek contact with another compable mycelium of the same species. When they meet, the two hyphae fuse, a process call plasmogamy that produces a “secondary mycelium.” The two different nuclei in the mycelial cells of the fused hyphal strands remain separate for a while. In basidiomycetes, this separate state (dikaryoc) may last many years. Eventually, the two nuclei in the cells fuse (karyogamy) to create diploid zygotes, which divide by meiosis into haploid cells to make spores with new genec combinaons. The secondary mycelium produces the complex fruing body, or sporocarp, differenang into separate parts of the mushroom -- stalk, gills, etc.

Spore Release and Dispersal Fungal spores are released and dispersed in various, oen ingenious ways. The ejecon of spores occurs by acve or passive means. Acve mechanisms involve internal forces creang tension or pressure to propel spores far enough to escape the fruing body. In some species, the discharge is explosive, shoong the spores up to eight feet. In gilled and pore mushrooms, the distance may be no more than half a millimeter, just enough to clear the gill or pore surface without hing the opposite side and fall from the cap, a mere flick. Ascomycetes absorb water to build internal pressure for an explosive release. Changes in humidity can create a mass release in cup fungi, somemes making an audible hissing sound. The air turbulence of hundreds of thousands of spores released at once helps elevate the spores, increasing their distance and the opportunity to catch a breeze. Passive mechanisms rely on external forces, such as raindrops, wind, or animals to eject or spread the spores. Pualls (Calvaa, Lycoperdon, Scleroderma, Calostoma, etc.) and Earthstars (Geastrum, Astraeus) rely on the force of falling raindrops to discharge a puff of spores and wind to carry them away. Snkhorns (Munus, Phallus) mire their spores in smelly slime that aracts flies to carry the spores away on their feet. Bird’s Nest Fungi (Cyathus, Crucibulum) have a uniquely designed splash cup to use the force of a raindrop to propel the spore packets (peridioles) up and out of the cup some distance. The peridiole is weakly aached to the cup by a long thread that easily snaps loose. The thread’s end is scky to grab nearby foliage and wrap around it, allowing the spore packet to release

15 spores from an elevated posion. There are species that grow on dung, and the spore release mechanism posions the peridioles for consumpon and later eliminaon by browsers.

Fluted Bird’s Nest Fungus (Cyanthus striatus)

Idenficaon General Groups of Fungi Based on Fruing Body Form (As found in Mushrooms of West Virginia and the Central Appalachians by William Roody)

Gilled Mushrooms have radiang vercal plates (gills) on the underside of the cap tapered to a ‘sharp’ edge. Spores are produced on the surfaces of the gills. Gilled mushrooms are so- fleshed and may be umbrella shaped with a stalk and cap or shelf-like without a stalk. They may be mycorrhizal or saprotrophic, and a few are parasic. Substrates include wood, humus, or soil. Large genera, such as Amanita, Russula, Lactarius, and Cornarius, each represent several hundred species. The deadly nave Destroying Angel (Amanita bisporigera), and its close relave the Death Cap (A. phalloides, introduced from Europe) are responsible for most mushroom fatalies in North America.

Chanterelles are usually grouped with the gilled mushrooms, however, the folds under their caps are not true gills. Unlike ‘sharp-edged’ true gills the folds of chanterelles are blunt ridges with many forks and cross veins. The folds are typically decurrent, running down the stalk. Chanterelles are funnel or trumpet shaped, slightly to deeply depressed in the center of the cap, and oen brightly colored red or egg-yolk yellow. A few species have smoother ferle surfaces that appear more wrinkled than ridged, such as Black Trumpets (Craterellus). The so-called ‘Scaled or Vase-shaped Chanterelles’ (Gomphus), vaguely resemble chanterelles and were historically treated within that group. Recent genec studies show the genus having closer es to the club fungi and earthstars listed below than true chanterelles.

16 Boletes are stalked mushrooms, so and fleshy with a thick layer of tubes under the caps forming a surface of small pores through which spores are released. This tube layer separates easily from the rest of the cap. Some boletes have a ring-producing paral veil. Most boletes form mycorrhizal associaons and are found on soil associated with both hardwoods and conifers. They can be very colorful, and species are eaten by a variety of wildlife.

Stalked Polypores have a shallow tube layer of surface pores that cannot be separated from the cap with ease. Most polypores are annuals, yet persist due to their woody or leathery texture, and either feed on dead organic maer or are parasites. They predominantly grow on woody substrates. If growing laterally on wood, some may appear stalkless, more closely resembling bracket or shelf fungi. Polypores can generate three different types of hyphae -- generave, skeletal and binding. Skeletal hyphae is thick-walled and contributes to the tough nature of many polypores. Ganoderma lucidum, known as Reishi or the “Mushroom of Immortality,” is widely coveted for alleged medicinal benefits.

Bracket or Shelf Polypores, grow on wood and are mostly fibrous or woody and leathery in texture. Most are annuals but oen persist due to the tough nature of the fruing bodies. There are some perennial species which add a new ferle layer of tubes each year. Most aach laterally to the wood substrate and are stalkless. Many of them can break down lignin and are important decomposers of wood. Chicken of the Woods (Laeporus spp.) are tasty when cooked, having a meaty texture like chicken. Turkey Tails (Trametes versicolor) appear in overlapping clusters.

Tooth Fungi have a ferle surface of hanging or projecng spines (“teeth”) on which the spores are produced. They have a variety of shapes (standard stalk and cap to bracket forms) and grow on a variety of substrates. Some species are saprotrophic, some mycorrhizal, and a few are considered parasic, found growing from wounds on living hardwood trees. Hericium spp. grow on wood and form a beard-like, globose mass or have a branched, frozen waterfall appearance.

Club Fungi are narrowly club shaped or finger-like in appearance with simple stalks, typically not branched or lobed, growing singularly or in clusters on various substrates. The ferle poron is usually clearly differenated from the stalk. Most are saprotrophic. One genus (Cordyceps) is parasic on the pupae of underground insects and other fungi like False Truffles. Jelly Clubs (Leoa), Earth Tongues (Microglossum, Geoglossum, and Trichoglossom), Snkhorns (Munus) and Dead Man’s Fingers (Xylaria polymorpha) are club shaped. [Note: This grouping is based on the physical appearance of the mushroom and should not be confused with the basidiomycetes, the so-called “club fungi,” a term which refers to the microscopic reproducve structures of that phylum. To emphasize this point, the species noted above are all ascomycetes, or “sac fungi.”]

17 Coral Fungi resemble marine corals. They may grow in clusters of simple tubular clubs (called Worm Corals) or may be branched and shrub-like, usually on the ground though somemes on decaying wood, and may be brightly colored. Many are saprotrophic; some are mycorrhizal. This category includes the bright Yellow and Orange Spindle Corals (Clavulinopsis spp.) Though unrelated to true coral fungi, the large leafy masses of Cauliflower Mushroom (Sparassis) have a marine-like quality too.

Pualls and Earthstars produce spores within a globular structure (peridium) or spore case that typically develops a central opening. They do not discharge spores through internal mechanisms, relying instead on external forces like raindrops to prompt spore release. Most are saprotrophic; a few are mycorrhizal. Earthstars (Geastrum, Astraeus) feature the starlike collar of their split outer covering. Stalked Pualls (Calostoma) may have a similar collar (C. lutescens) or a pile of sloughed, gelanous shavings (C. cinnabarina).

Bird’s Nest Fungi have spores contained in small packets called peridioles resembling eggs in a small splash cup that aids dispersal of the ‘eggs’ during rainfall. [See Spore Release and Dispersal] Similar in appearance to cup fungi, they are related to pualls and are saprotrophic.

Jelly Fungi have a rubbery, gelanous texture, especially when moist, usually appearing as a crimped or convoluted mass on wood. They may shrivel in dry condions and revive in wet. Some are saprotrophic; some, such as Witches’ Buer (Tremella mesenterica), are parasites on other fungi (mycoparasic). Jelly Tooth or Jelly Tongue (Pseudohydnum gelanosum) has a gray translucent tongue-shaped fruing body with spines like tooth fungi.

Cup Fungi usually appear in groups on the ground or on wood as open, shallow cups. The ferle inner surface is lined with spore sacs (asci) that forcibly rupture shoong spores into the air. There are species with easy-to-spot coloraon, like Orange Peel (Aleuria aurana) and Scarlet Cups (Sarcoscypha spp.), and less noceable ones, Hairy Rubber Cup (Galiella rufa) and Common Brown Cup (Peziza). Blue-green Wood Stain (Chlorociboria) rarely produces its small clusters of bright turquoise fruit yet may easily be found in decaying wood stained the characterisc color. Many species are saprotrophic. Some fruit in spring.

Morels, False Morels, Saddle Mushrooms typically fruit in spring. All contain potenally dangerous toxins, but cooking alters this property in true morels (Morchella), which are tasty edibles. False morels (Gyromitra) and saddle mushrooms appear convoluted, folded, and brain- like. True morels are columnar to conical and arranged with obvious pits divided by ridges. They are saprotrophic and can also form mycorrhizal associaons.

18

Turkey Tails (Trametes versicolor) Shelf or Bracket Fungus

Witches’ Buer (Tremella mesenterica) Jelly Fungus

19 Field Idenficaon Characteriscs Various physical (morphological) characteriscs of the fruing bodies (sporocarps) can be assessed to determine a tentave idenficaon of fungi in the field. Spore color may be determined with a spore print (set cap on paper for a few hours). The most accurate idenficaon requires examinaon of various microscopic features. Size, shape, and surface features of spores are among the characteriscs oen crucial for idenfying many fungi at the species level. Macroscopic field characteriscs include:

• Gills, pores, or teeth under the cap -- This is the first and most important determinaon for idenficaon. • Gill characteriscs -- Gills aached to the stalk or free from the stalk (cung a mushroom in half lengthwise through the center will show gill aachment), gill color, spacing between gills, branching gills, thickness of the gills, and the presence of veins running between the gills can be important disncons. • Cap and stalk -- Color, paerns, presence of warts or patches, surface texture, stalk ring or skirt, stalk diameter and uniformity, stalk base, overall size, and height should be noted. Many of these qualies may change as the mushroom ages. With mulple fruing bodies, note if the stalks are separate or fused at the base. • Bruising -- Exterior flesh may change color when damaged. Exposure to air can cause interior flesh to change color in some species. • Exudates -- Exudes milky latex (Lactarius) when cut or other liquid, somemes naturally occurring on the fruing body. Older specimens may be dry or slow to produce exudate. • Pore characteriscs -- The outer edge of pores may be smooth or jagged. • Odor -- A few species are known for having a disnct odor. • Substrate -- Fungi may be found growing on soil, organic ground lier, downed branches, logs, stumps, or live trees. • Quanty of fruing bodies present -- Some species appear singly, others are typically found in groups of several to many individuals (gregarious). • Season -- Most fungi fruit in summer and fall, but some species are found in spring and even winter. Spring fruing fungi include true and False Morels (Morchella and Gyromitra), Oyster Mushroom (Pleurotus), several Inky Caps (Coprinus), and some of the cup fungi such as Common Brown Cup (Peziza), Scarlet Cup (Sarcoscypha), and Devil’s Urn (Urnula) Others, like Witches’ Buer (Tremella) and Wood Ears (Auricularia), can be found in winter as well. • Neighboring plants -- Mycorrhizal fungi are oen closely associated with certain tree species.

Primary fungi sources: (full bibliographical informaon in secon VI. Resources) Mushrooms of West Virginia and the Central Appalachians, William C. Roody The Kingdom Fungi: The Biology of Mushrooms, Molds, and Lichens, Steven L. Stephenson The Fih Kingdom (Third Edion), Bryce Kendrick Fleshy Fungi of the Highlands Plateau course, Andrew S. Methven, Highlands Biological Staon

20 II. Lichens

Symbioc Relaonship The accepted definion of a lichen by the Internaonal Associaon for Lichenology is “an associaon of a fungus and a photosynthec symbiont resulng in a stable vegetave body having a specific structure.” More humorous takes include “a fungus and an alga that take a ‘lichen’ to one another” and “fungi that have discovered agriculture.” Lichens do not have one common ancestor, deriving from mulple independent origins over me. The fungal component, or mycobiont, is most likely from the phylum Ascomycota. More than 40% of nearly 30,000 species of ascomycetes form lichens, plus a small number of species within the phylum Basidiomycota (most mushroom producing fungi and relaves). Less than 15% (14,000 spp.) of all currently known fungi form lichen associaons. Lichens are classified taxonomically under the name of their fungal component. The algal component, or photobiont, is mostly green algae. Some lichens contain cyanobacteria, also known as blue-green algae, and a few enterprising lichens use both. A smaering of other algal types have been found. According to Brodo, et.al., “only a dozen genera [of algae], however, represent the photobionts in the vast majority of lichens.” Any parcular lichen fungus will usually associate with the same photobiont(s). An alga, however, may associate with many different fungi. Species in the green alga genus Trebouxia are rarely found in a free-living state and comprise the most common algae in lichens. Green algae contain chloroplasts (specialized bodies of chlorophyll) in their cells and produce a true green color in lichens. Cyanobacteria have chlorophyll throughout the cell fluid and produce a dark blue-green or blue-gray color. They are free-living in nature, and Nostoc is a common genus found in jelly lichens and many species of Dog or Pelt Lichens (Pelgera spp.) Each component can be grown separately in culture, and each looks different as a disnct organism. In parcular, the fungus forms a shapeless mass of hyphae. When the two are mixed together in appropriate condions, the characterisc form of the lichen is achieved, a process called morphogenesis. It is believed that the fungus contains all the informaon to form the lichen, but the presence of the alga is essenal to spark the transformaon process. However, the natural lichen structure was never fully replicated. Recent research revealed a third organism. Basidiomycota yeasts have been found in a majority of the macrolichens studied on six connents. The nature of this complex relaonship and role the second fungus plays are not yet understood. Is the symbioc relaonship parasism, commensalism, or mutualism? A case can be made for each. The nature of the relaonship varies among species and can be rather complex. In most instances, the fungus invades the alga’s cells, eventually killing them, but the cells reproduce faster than they are destroyed. Branches of fungal hyphae surround and press against the alga’s cell wall. Somemes ny structures called haustoria penetrate the cell wall. The presence of the fungus causes algal cell walls to become more permeable, allowing some of

21 the carbohydrates produced in photosynthesis to leak through and be absorbed by the fungus. The rest is used by the algae. The relaonship is far from one-sided. The fungus contributes several benefits, most notably the lichen’s structure, protecng the photobiont. The ssue provides a more consistently moist environment, as the outer cortex protects against moisture loss and acts as a sun screen against excess light. At the same me, the broad leafy form exposes more surface to the light. These qualies allow the algae to survive in habitats they could not colonize on their own, such as rocks and dry tree bark. The partnership between fungus and alga requires a delicate balance. If anything affects one partner or the other, the relaonship could break down quickly leading to the lichen’s death.

Common Lichen Forms The vegetave body of a lichen is called the thallus (thalli, pl). Thalli have four main growth forms by which lichens are characterized: crustose, foliose, squamulose, and frucose.

Crustose lichens adhere directly to the substrate and form no lower surface or cortex. They can only be removed by taking part of the substrate. The rest of a crustose thallus is more or less strafied with upper cortex, algal, and hyphal layers. Crustose lichens have several typical surface textures and can be brightly colored. They grow outward covering more substrate surface, and the leading edge of this growth (called the prothallus) contains no algae, only fungal ssue, and is usually dark in color. Gold Dust Lichen (Chrysothrix candelaris) looks like bright yellow spray paint on rocks.

22 Foliose lichens have flaened, leaflike thalli that are lobed and typically sub-erect. Some may be large and more three-dimensional, such as Lung Lichen (Lobaria pulmonaria), others are closely aached to the substrate (Flavoparmelia). Foliose thalli are usually layered with an upper cortex that may contain a variety of pigments, the algal layer, a layer of fungal hyphae (the medulla), and a lower cortex. Some have no lower cortex. A few are not strafied, and the algae and hyphae mix uniformly, as with jelly lichens. Foliose lichens aach to the substrate through hyphae of the lower cortex or medulla or small root-like structures called rhizines. Hairlike appendages on the edge of the thallus are called cilia.

Squamulose lichens are an intermediate form between foliose and crustose. They are scalelike with each ny “scale” aached separately in small to large patches. The structure of each scale thallus is similar to foliose lichens, but their overall size places them with crustose lichens as microlichens. Brish Soldiers and Pixie Cups (Cladonia spp.) are squamulose lichens.

23 Frucose lichens are three-dimensional with either an erect or pendulous growth habit and no obvious upper and lower layers. They may be unbranched filaments or highly branched and shrubby in appearance. The cortex surrounds the stalked thalli and the algal and hyphal layers are arranged inside. Tough carlaginous ssue offers support. These lichens are usually aached to the substrate at one or just a few points. Beard Lichens (Usnea) are common examples.

Lichen Funcon A colorless upper cortex generally yields a gray or greenish gray lichen. When wet, the cortex becomes more transparent and the brighter green color of the algae is more apparent. Several bright pigments may be present in the cortex of various species. These colors are most oen found in areas of high solar radiaon and exposure, where these pigments offer more protecon to the algae. Surface textures, such as hairs or scaly cells of dead ssue, break up the light. Color can also absorb heat energy or reflect it in habitats with harsh temperatures. Lichens cover 8% of the earth’s surface. They are small, slow growing, and long lived, with the ability to absorb moisture rapidly through all surfaces of the thallus from humidity, fog, and dew as well as rain. Their physical design helps them trap moisture and slow evaporaon. Because they do not have a protecve waxy coang like plant leaves, lichens dry out quickly and survive droughts by entering a dormant state. As they do not have true roots, lichens absorb minerals needed for growth from the atmosphere or substrate through moisture. This trait can be harmful when more toxic compounds are absorbed, both for the lichen and any organism that feeds on the lichen. In nitrogen poor areas, lichens oen employ cyanobacteria, either exclusively or in conjuncon with green algae, for their ability to fix atmospheric nitrogen. Lichens without cyanobacteria in

24 otherwise nitrogen poor environments can be found near bird or animal populaons to derive this essenal element from their waste products.

Sexual Reproducon in Lichens Lichens reproduce sexually forming ascospores of the associated fungus. The lichen can produce two different structures, (1) apothecia, disk or cup shaped fruing bodies aached to the thallus or raised on stalks (podea), whose inner surfaces contain the spore sacs (asci) or (2) perithecia, flask-shaped bodies parally embedded in the thallus with a narrow opening for spore release. To be successful, the spores must find a suitable algal partner and a conducive environment for growth. As a backup, lichens also reproduce asexually in several ways. Soredia are fine, powdery bits of hyphae wrapped around a few algal cells that develop in the algal layer and are released through cracks in the cortex. Isidia are smooth, peg-like outgrowths of the thallus cortex containing hyphal and algal cells. They are fragile and easily broken for distribuon. Bits of the thallus itself can break away or form minute lobes that drop off. Lichens can also produce asexual spores called conidia in ny structures (pycnidia) that resemble perithecia.

Lichen Ecological Roles Soil formaon -- Thanks to their self sufficiency and ability to withstand harsh condions, lichens can colonize bare, or newly exposed surfaces. They are oen the dominant life forms in extreme environments. The fungal hyphae of lichens can penetrate several millimeters into rock surfaces, and chemicals in many species can soen rock to speed weathering. Lichens catch and hold parcles of soil and contribute to soil formaon when they die. On bare soil, they stabilize erosion, hold moisture, provide shade, and contribute organic maer and nitrogen. In these ways, lichens iniate biological succession. Wildlife -- Various browsers eat different species of lichens from forest to tundra. Small mammals, such as flying squirrels and voles, eat lichens. Many ny arthropods eat lichens or live within lichens (and mosses), including waterbears (or moss piglets, Tardigrades). Many bird species and flying squirrels use lichens as nesng material. Certain insects, such as some moths, and the gray and bird-voiced treefrogs are paerned to blend into lichen-covered tree bark and escape detecon through camouflage. Green salamanders resemble the lichen-covered rocks of their habitats. Environmental indicators -- Since lichens absorb moisture and minerals from the air, they can be excellent bioindicators of air quality. Foliose (such as Lobaria) and frucose (such as Usnea) species are more sensive. The numbers and species of lichens change dramacally from city to forest. Not only can their presence and health be monitored, they can be analyzed to assess specific contaminants. Lichens can also indicate ecological connuity and age. Certain species are only found in old growth communies and play an important role in the complex funconing of previously unknown ecosystems, such as those found in the tree canopies of ancient forests.

25 Human uses -- Lichens have been used for natural dyes, perfumes, medicines, poisons, food, clothing, and decoraons. By calculang their growth rate, lichen colonizaon of a surface can be dated through lichenometry.

Lichen Idenficaon Careful examinaon of the morphological characteriscs (physical features of form and structure) of lichens will idenfy many species in the field. Make note of the primary growth form, shape, size, color (which may be different when wet), presence and appearance of specialized structures such as cilia (hairlike appendages along thallus edge), rhizines (rootlike structures on thallus undersurface), podea (short stalks under apothecia), or reproducve structures (apothecia, perithecia, isidia, soredia), the substrate, and habitat. Lichens contain various compounds that are known to react chemically to parcular agents. Use of chemical reagents for spot tests will produce predictable results to confirm identy based on color change. These compounds may not be found in all parts of the lichen, therefore, the locaon of color change is oen diagnosc as well. Among the most commonly used chemicals are potassium hydroxide (KOH) and sodium hypochlorite (household bleach). Certain species of lichens will fluoresce under ultraviolet light. In some instances, microscopic examinaon of the spores is necessary to determine the species.

Primary lichens sources: (full bibliographical informaon in secon VI. Resources) Lichens of North America, Irwin M. Brodo, Sylvia Duran Sharnoff, Stephen Sharnoff Lichens, William Purvis

26 III. Plants -- Kingdom Plantae

Plant Funcon and Evoluon

Botany is the scienfic study of plants and plant life.

Plants and Photosynthesis Organisms that can make their own food are called autotrophs or primary producers. All other organisms must consume these producers directly or indirectly to live and grow. The vast majority of plants are autotrophs. All green plants (and some algae) have chloroplasts in certain leaf and stem cells. These chloroplasts contain chlorophyll which enables them to harness the energy of the sun and manufacture food through photosynthesis. The photosynthec process occurs in two stages. First, chlorophyll absorbs energy from red and blue light (reflecng green, thus the color we see). This energy absorpon, called the light reacon, is stored as a chemical compound. The second stage, called the dark reacon, uses this stored energy in the presence of water to convert carbon dioxide into sugar and release excess oxygen.

6CO2 + 6H2O + Sunlight Energy --> C6H12O6 + 6O2 Plants use these sugars and starches during respiraon for growth and reproducon and oen store excess food, water, or nutrients in roots, stems, or leaves for future growth. Food stored in fruits and seeds ences animals for dispersal and supports germinang seedlings unl they unfurl seed leaves and begin independent photosynthesis.

Plant Evoluon During their evoluon over 500 million years, species have diverged from common ancestors to populate the globe with well over 300,000 different plants. A simple chart called a cladogram demonstrates four main developmental milestones: waxy cucle and stomata, vascular ssue, seeds, and flowers.

1. Waxy cucle and stomata -- For green algae to begin the transion from their aquac environment to life on land, they first had to devise a way to retain internal moisture. A waxy cucle on the surface composed of lipids protects foliage against desiccaon but creates another problem with gas exchange. Development of small pores, called stomata, allow leaves to take in CO2 and conduct photosynthesis.

2. Vascular ssue -- Mosses and other bryophytes (dominant plants in the Ordovician Period, 450 mya) lack true vascular ssue having no roots and no lignin for the development of water conducng structures. Lignin, a complex polymer, is present in all vascular plants. These plants must pull water and nutrients up the xylem (composed of dead cells) for photosynthesis and distribute sugars by way of the phloem (composed of living cells) throughout the plant’s ssues

27 for respiraon. Leaf stomata release water vapor (transpiraon) in the process of drawing water from roots to leaves. Vascular ssue also provides structure necessary for plants to grow in height. Ferns in the late Devonian Period (360 mya) are characterisc of early vascular plants.

3. Seeds -- Mosses and ferns reproduce by spores, and both are dependent on water for ferlizaon, as male gametes must swim to reach female gametes, and germinaon. Spores contain no nutrive ssue and must land in a hospitable locaon under proper condions to germinate and connue the life cycle. The development of vascular ssue allowed plants to grow taller making water-dependent ferlizaon even more difficult. Plants evolved the ability to retain the gametes on the sporophyte unl ferlizaon occurred and beyond, offering both protecon and nutrion. Ferlized seeds are shielded by a seed coat unl germinaon and a layer of nutrional ssue or endosperm to help them survive a wider range of condions following germinaon. Fernlike plants called “seed ferns” were the first to have aached seeds. Gymnosperms developed in the late Paleozoic Era (300 mya) and dominated during most of the Mesozoic Era (250 to 65 mya).

4. Flowers -- Gymnosperms are largely wind pollinated. Their seeds develop on the surface of the reproducve structures, such as the scales of a cone, and are considered “naked” (gymno) since there is no enclosing structure. Angiosperms (angio means “vessel”) or flowering plants evolved an enclosing structure called a carpel (or pisl), comprised of the sgma, style, and ovary for their seeds. Rather than rely on wind, the development of colorful petals and nectar in associaon with the pisl (containing female gametes) and stamens (containing male gametes) serves to aract insects and animals as pollinators. Aer ferlizaon, the ovary develops into fruit with a hard or so outer coang containing the seed(s) which aracts insects or animals for dispersal. Angiosperms rose to dominance by the end of the Cretaceous Period (140 to 65 mya) and have connued their diversificaon throughout the Cenozoic Era (65 mya to today).

28

Sexual Reproducon in Plants Sexual reproducon in all plants (and fungi) is a two stage process called the alternaon of generaons. This process for seed plants and spore-producing fungi, bryophytes, and ferns alternates between the gametophyte (haploid stage) and sporophyte (diploid stage). The nuclei of the gametophyte’s haploid cells have one set of chromosomes; the sporophyte’s diploid nuclei have two sets. For all vascular plants -- ferns, gymnosperms, and angiosperms -- the sporophyte generaon is the dominant generaon, the ferns, trees, shrubs, and wildflowers. The sporophyte is much larger, longer lived, and able to support itself. The gametophyte generaon is very reduced in size, occurs over a brief me span, and except for ferns, is dependent on the sporophyte for nutrion. Fern gametophytes are independent organisms. With nonvascular plants, the gametophyte generaon is dominant. The green moss, liverwort, and hornwort plants are the gametophytes. They are photosynthesizing, self supporng, long lived, and larger than the sporophyte generaon, which are ephemeral structures growing from and dependent on the gametophyte for nutrion. The basic processes of meiosis (cell division reducing the chromosomes by half), mitosis (cell division retaining the same number of chromosomes), and ferlizaon (fusion of the male gamete [haploid] and female gamete [haploid] to form a zygote [diploid] and develop an embryo) occur with all plant reproducon. The processes for fungi are somewhat similar but more complicated with unique differences. Addional details are provided for each group where they are discussed in the guide. However, simply reading about the processes behind plant and fungi sexual reproducon can be confusing. The selecon of online videos below feature graphics that may prove helpful.

-- Alternaon of generaons: hp://www.youtube.com/watch?v=SCTNKTfa-s0 -- Angiosperms: hp://www.youtube.com/watch?v=yjpbKpKgUSs -- Angiosperm carpel and seed development: hp://www.youtube.com/watch?v=UBfY_9jeq6s -- Gymnosperms: hp://www.youtube.com/watch?v=Fa7mjzEgTAQ hp://www.youtube.com/watch?v=2gWEgrMwMe0 -- Mosses: hp://www.youtube.com/watch?v=ZLrcfaeiFHM -- Ferns: hp://www.youtube.com/watch?v=Fhk-Y0duNjg hp://www.youtube.com/watch?v=WR5PIqOcfKw

29 Scienfic Classificaon Two terms are used in associaon with the classificaon of any living (or exnct) organism -- systemacs and taxonomy. These terms are oen used interchangeably. Systemacs has been described as having six components: (1) provide scienfic names for organisms (2) describe organisms (3) preserve collecons of organisms (4) provide classificaon, idenficaon keys, and distribuon data on organisms (5) invesgate organisms’ evoluonary histories (6) consider organisms’ environmental adaptaons Components one through four relate to classificaon and are addressed by taxonomy. Taxonomy literally means “order method.” Each organism species and each grouping of organisms (genus, family, order, etc.) is considered to be a taxon, e.g., Quercus alba (White Oak) is a taxon, the genus Quercus (all oaks) is a taxon, the family Fagaceae (beech family) is a taxon, etc. Collecvely, they are taxa (plural).

Main Biological Kingdom Ranks Biological kingdoms demonstrate a hierarchical relaonship, a family tree of sorts, and have seven primary ranks (categories) as follows: Kingdom, Phylum (Division), Class, Order, Family, Genus, Species In the plant kingdom, “division” has typically been used in place of “phylum,” though this convenon is becoming less common. There can be subcategories under each rank. Each organism is a disnct species. Closely related species are grouped into a single genus. Related genera (plural) are grouped into a single family, and so on. Many mnemonics have been created to keep the rank order straight. • Kings Play Chess On Fine Green Silk (or Fat Green Stools) • Keeping Precious Creatures Organized For Grumpy Sciensts • Kids Prefer Cheese Over Fried Green Spinach • King Phillip Chews On Funny Ginger Snaps

Nomenclature: The Binomial In the eighteenth century, Swedish botanist Karl Linne (Carl von Linne), whose name was Lanized to Carolus Linnaeus, proposed the modern system of biological nomenclature that simplified all names to a binomial. Before this, organisms had mulword names, up to 15 descriptors, that varied country to country. Linnaeus established a two-name system for each organism consisng of its genus and species. He is considered the father of modern taxonomy. In Linnaeus’ day, organism relaonships were judged on the similarity of physical traits. In the case of plants, the sexual characteriscs of the flower provided the primary basis for this judgement. DNA studies offer a new way to gauge relaonships, resulng in the reorganizaon of families and the reclassificaon of many species under new names. Any previous name remains aached to the species as a synonym.

30 The naming of plant species is strictly governed by the Botanical Congress and its Internaonal Code of Botanical Nomenclature, which sets specific protocols for naming new species or changing an established name. Scienfic evidence must be presented in a professional arcle and published in a peer-reviewed journal before any proposed classificaon change is accepted. Unfortunately, individuals and organizaons may or may not adopt these accepted changes. Various floras and plant checklists oen represent a hodgepodge of past and current nomenclature. Therefore, it is advisable to choose one published lisng to follow. In Tennessee, the new state flora, Guide to the Vascular Plants of Tennessee, 2015, incorporates the most recent taxonomic changes. The use of Lan in organisms’ names, chosen for its universal scholarship in Linnaeus’ day, oen serves to deter and inmidate people today. Many taxonomic groups with fewer species, such as reples, mammals, or birds, have widely established, uniform common names. Common names of plants, however, are many and variable, with ample opportunity for confusion. Plants may have several (five or more) common names by which they are known. Different plants share the same common name. Accepted common names may vary from one region to another. In contrast, scienfic names (and synonyms) are consistent worldwide and provide the surest method of accurate identy. Lan names are not chosen randomly. There is oen a direct connecon to some aspect of the organism -- an outstanding feature or idenfying characterisc, the geographic locaon in which it was first found and described, etc. Books that translate the Lan meanings oen provide welcome insight and bring to life these names mired in a dead language through interesng stories in the hands of a good naturalist. See Resources: Stearn’s Diconary of Plant Names for Gardeners. Pronunciaon of botanical Lan follows the English use of vowel sounds and is not as difficult as it appears. Pracce helps. Since Lan is a dead language and has several iteraons (academic, liturgical, biological), don’t sweat it. Pronunciaons vary region to region and professor to professor. As one Texas botanist said, “You can put whatever ‘twang’ you want to on it.” There are a few general guidelines in Appendix B of this study guide.

31 Key Ranks: Species, Genus, and Family

Species -- The definion of a species is a group of individuals who interbreed and produce viable offspring only with each other and are idenfied by a combinaon of unique characteriscs. A few genera of species do hybridize naturally. This tendency is parcularly noteworthy among some ferns (spleenworts and wood ferns), oaks (Quercus), hawthorns (Crataegus), and coneflowers (Echinacea). As a word, “species” is used for both singular and plural applicaons; abbreviaons are sp. (singular) and spp. (plural). The species name is the specific epithet. It is an adjecve, wrien in lower case and italicized (or underlined), describing a plant characterisc Mitchella repens (creeping), nong geographic locaon Echinacea tennesseensis, or acknowledging a person Lilium michauxii (André Michaux). There are ranks below species level. A subspecies (ssp.) or variety (var.) is a naturally occurring form or structural variaon(s) having a geographic, ecological, or phylogenec basis. The two terms are largely equivalent in pracce. Once a parcular term is assigned, however, the other term may not be used interchangeably. This name appears aer the species name: Acer saccharum ssp. nigrum (Black Maple) or Chrysogonum virginianum var. brevistolon (Green- and-gold).

Genus -- The genus (genera, plural) is a noun. It is capitalized and italicized. There may be one or several species within a parcular genus based on a set of similar characteriscs. There is usually gender agreement between the genus and specific epithet with feminine (-a) or masculine (-us) endings; (-um) is neutral. Trees are an excepon to this rule. In Greek and Roman mythology, wood nymphs lived in trees, which were given a feminine gender in Lan. Botanical Lan honors that tradion, giving the majority of tree specific epithets a feminine ending even when the genus is masculine -- Quercus rubra (Red Oak), Cornus florida (Flowering Dogwood), Juniperus virginiana (Eastern Red Cedar)

Family -- A grouping of related genera forms a family. The family name derives from one genus that is recognized as the “type” characterizing the group. The ending -aceae is added to the generic name, and it is capitalized but not italicized. Example: For the mustard family, the type genus is Brassica, and the family name is Brassicaceae. The ending is pronounced ‘ay-see-ee; in the example, bras-sih-‘cay-see-ee. The set of similar traits found within family groupings is a great asset in field idenficaon. Determining which family or families demonstrate certain physical characteriscs narrows the opons significantly, improving idenficaon speed and accuracy. The book Botany in a Day (Elpel) presents the paerns of family traits for vascular plants. For instance, the majority of species in the mint family, Lamiaceae, will have opposite leaves, bilaterally symmetrical flowers, and square stalks. All will have four nutlets in the seed capsules. Many will be aromac. Not all families are this disncve, but each has its own set of idenfying characteriscs.

32

Buercup family (Ranunculaceae) has variable numbers of petals and numerous stamens.

Rose family (Rosaceae) has five petals and numerous stamens.

33

Mint family (Lamiaceae) has bilaterally symmetrical flowers, opposite leaves, and square stems.

Mustard family (Brassicaceae) has four flower petals and radially arranged seedpods, oen as upright narrow structures called siliques.

34 !!!!!!!!!! Kingdom Plantae

!! ! ! ! ! !!!Green Algae !! ! ! ! ! ! ! ! ! !!!Land Plants !!

!!! ! Nonvascular!! !!!!(Subkingdom)!! ! ! Vascular (Trachiobionta) !!Mosses (Bryophyta)!! ! !!Liverworts (Marchantiophyta)!! !!Hornworts (Anthocerotophyta)! !

! Seedless Plants! !!!(Superphyla)!! ! Seed Plants (Spermatophyta)

Lycophyta!!!!Monilophyta!! ! (Phyla)!! ! Gymnosperms (Coniferophyta) ! Club mosses!! ! ! Horsetails!! ! ! ! ! ! ! Spike mosses!! ! Ferns!! ! ! ! ! ! ! ! Angiosperms (Magnoliophyta) ! Quillworts!! ! ! ! ! ! ! ! ! ! ! ! !

!!!!!!!!!!!! !!!!!!!!!!!!! (Class)! !!!Monocots (Liliopsida) !!!!!!!!!!!!!!!!!!!Dicots (Magnoliopsida)

The major divisions (phyla) in the plant kingdom • nonvascular plant divisions for mosses, liverworts, and hornworts • ferns and their relaves • seed plants, gymnosperms and angiosperms

35 Nonvascular Plants: Mosses, Liverworts, and Hornworts

Mosses, liverworts, and hornworts are nonvascular plants, among the earliest land plants, and are found throughout the world in a variety of habitats from Antarcca to the tropics. Collecvely referred to as bryophytes, each group is its own division. Bryophyta -- mosses Marchanophyta (Hepacophyta) -- liverworts Anthocerotophyta -- hornworts Nonvascular species differ from vascular plants in several ways. Lacking lignin to form water- conducng structures, their physical design and habitats allow them to absorb needed water and nutrients from the atmosphere, rain, or surface streams through their leaves and stems. Without the structural support of vascular ssue, they remain short, typically a few cenmeters in height. Having no need for roots, small rhizoids help them aach to various rock, soil, or wood substrates. They reproduce by spores, and unlike vascular plants, their dominant life stage is the gametophyte. [A list of common moss, liverwort, and hornwort species is in appendix A.]

Bryophyte Ecology Because of their life cycle requirements, most bryophytes live in moist places, such as waterfall spray cliffs, rock seeps, bogs, or areas of high rainfall. Liverworts and hornworts, in parcular, need a damp to wet environment. In more moderate environments, bryophytes typically grow close to the ground or in sheltered microhabitats. Some moss species become dormant in dry spells and revive with precipitaon. Many bryophytes have adapted to grow well in moderate to deep shade. Typical substrates include rock, soil, rong wood, and tree bark. Some species may be restricted to just one type of substrate. The pH level (acidity/alkalinity) of soil or rock may encourage or deter certain species. Most bryophytes are terrestrial, but a few are aquac. Bryophytes, parcularly mosses, capture and hold large amounts of water and water-borne nutrients, protect soil from erosion, serve as nursery beds for germinang seed plants, and provide habitat for numerous insects and microorganisms.

Atrichum altecristatum, Wavy Starburst Moss

36 Mosses Esmates on the number of moss species worldwide range from 10,000 to 15,000. Mosses vary significantly in form, texture, habitat, etc. Yet most mosses possess certain defining characteriscs. The leaves on a moss stem are spirally arranged imparng a round appearance. Each leaf forms a single point at the p and typically has a central midrib. Moss leaves are generally one cell thick, except for the midrib, and absorb water through all their surfaces. The green moss plants are the gametophytes (haploid stage, one set of chromosomes). At the ps of leafy branches, male moss gametophytes produce antheridia that generate the male sperm, each with two flagella, and female gametophytes produce archegonia that contain the eggs. In wet condions, sperm ‘swim’ from the male plant to the female plant ferlizing the eggs (forming zygotes) which mulply through mitosis in the now developing sporophytes, the diploid stage (two sets of chromosomes). The sporophytes grow into small capsules on long stems aached to the moss plant. As the sporophytes mature, the diploid cells split in half by meiosis to produce haploid spores that will be released to germinate, grow into green moss plants, and begin the cycle anew. Moss sporophytes are oen persistent aer spores are released. The shape and structural details of the sporophytes help idenfy species. These structures include the seta (stem), capsule, calyptra (hood covering the capsule), operculum (cap on the capsule), and peristome (ring of tooth-like projecons around the capsule opening regulang the release of spores). A few species reproduce asexually by several means, including the producon of gemmae, ny plant buds of about a dozen cells each, in lile splash cups. Raindrops propel the ny buds out of the cup to develop a new, genecally idencal plant.

Moss sporophytes

37 Liverworts There are two types of liverworts, leafy (the more common form) and thalloid. Leafy liverworts superficially resemble mosses, but for most species, there are simple disnguishing characteriscs. The leaves of a leafy liverwort appear in two rows on the stem, giving them a flat appearance. The leaves normally overlap one another like shingles, are either rounded or lobed at the p, and do not have a midrib. Mosses differ in each of these traits. Thalloid liverworts have a sheet of green ssue that grows in lobes but is not differenated into leaves and stems. They are larger and easier to spot. Decent-sized colonies can be found coang soil or rocks near or in streams, seeps, wet cliffs, etc. The process of reproducon in liverworts is similar to mosses. However, unlike mosses, liverwort sporophytes typically develop undetected close to the plant, quickly shoot up on a fragile stalk to release spores, and wither in a few days. Some liverworts can also reproduce asexually by gemmae (ny plant buds) like mosses.

Dumorera hirsuta, thalloid liverwort

Hornworts The hornwort gametophyte looks a lot like a thalloid liverwort (flat green ssue), but the sporophyte is a thin, green, hornlike projecon that connues to grow from the base. The p of the sporophyte splits down the center to connually release mature spores as the sporophyte grows from the base producing new spores. This adaptaon allows it to produce and discharge spores over a prolonged period. They also appear to incorporate colonies of Nostoc (cyanobacteria) in their gametophyte ssue, taking advantage of its ability to fix nitrogen. Hornworts are sll something of a puzzle, and only about 100 species are known worldwide.

Primary bryophytes sources: (full bibliographical informaon in secon VI. Resources) A Trailside Guide to Mosses and Liverworts of the Cherokee Naonal Forest, Paul G. Davison Outstanding Mosses and Liverworts of Pennsylvania and Nearby States, Susan Munch Gathering Moss: A Natural and Cultural History of Mosses, Robin Wall Kimmerer

38 Vascular Plants

Ferns -- Pteridophyta

From the Carboniferous period (Mississippian and Pennsylvanian Periods) to today, ferns have survived and adapted first as large land plants. In the Cretaceous Period as angiosperms rose in size and dominance, ferns became smaller and evolved the ability to capitalize on light wavelengths in a dim understory. Ferns are found all over globe, and some have broad ranges as spores can travel great distances.

Fern Cladogram

Ferns and Fern Relatives (Allies)

Lycophyta Monilophyta Spermatophyta (seed plants)

Clubmosses Eusporangiate Ferns Huperzia Ophioglossum Diphasiastrum Botrychium, etc. Lycopodium, etc.

Horsetails Leptosporangiate Ferns Equisetum Dryopteris Quillworts Spikemosses Asplenium, etc. Isoëtes Selaginella

Adpated from a diagram in Ferns of Northeastern and Central North America, 2nd Ed., Boughton Cobb, Elizabeth Farnsworth, and Cheryl Lowe

A fern is a seedless vascular plant that has large complex leaves with branching vein paerns and reproduces by spores. Plants in the Lycophyta -- clubmosses, quillworts, and spikemosses -- also reproduce by spores but diverged earlier in evoluonary me from the “true ferns” and seed plants. While they are sll studied alongside ferns, they are no longer believed to be as closely related to true ferns as once thought. Several species of clubmosses are found in Tennessee forests, including most commonly Shining Clubmoss (Huperzia lucidula, East Tennessee), Southern Ground-Cedar (Diphasiastrum digitatum, statewide), and Flat-branch Tree Clubmoss (Dendrolycopodium dendroideum, East

39 Tennessee). Quillworts and spikemosses are not as readily encountered. The life cycles for all lycophytes and horsetails are similar yet differ in some details to that of true ferns.

Southern Ground Cedar (Diphasiastrum digitatum)

Monilophyta are true ferns (leptosporangiate) and their closest relaves (horsetails and eusporangiate ferns). The differences between eusporangiate and leptosporangiate ferns center on characteriscs of the sporangia (small structures containing spores) and the form of emerging fronds (leaves) in spring. Eusporangiate ferns produce vast numbers of spores in large sporangia with thick walls that develop from a group of cells. The sporangia simply spill spores when mature. In spring, their leaves emerge from the ground erect or bent double. In contrast, leptosporangiate fern leaves emerge coiled in the familiar fiddlehead associated with most common ferns. Their sporangia are small, developing from a single cell, with thin walls that feature a ring of thickened cells (annulus) designed to break open and forcibly eject the spores. Each sporangium contains 64 spores. Our eusporangiate ferns include adder’s-tongues (Ophioglossum spp.), Ralesnake Fern (Botrypus virginianus), grape ferns (Sceptridium spp.), and moonworts (Botrychium spp.) The leptosporangiate ferns include those genera whose species are common plants in the forest -- wood ferns (Dryopteris spp.), spleenworts (Asplenium spp.), Christmas Fern (Polyschum acroschoides), New York Fern (Thelypteris noveboracensis), Cinnamon Fern (Osmundastrum cinnamomea), bladder ferns (Cystopteris spp.), and many others. Horsetails (Equisetaceae), one of the oldest fern families, have hollow, grooved stems containing silica that were used to scour pans. Leaves are reduced and fused into a small sheath at each node on the stem. Spore-producing cones develop at the top of ferle stems. Tennessee has two species of horsetails, including Common Scouring Rush (Equisetum hyemale ssp. affine).

40

Ralesnake Fern (Botrypus virginianus) and Mountain Wood Fern (Dryopteris campyloptera).

Fern Life Cycle Ferns produce spores on the undersides of frond leaflets or in separate modified leaflike structures. In leptosporangiate ferns, the ny sporangia, each bearing 64 spores, are grouped in small clusters. Each cluster of sporangia is called a sorus (sori, plural). The sori are oen arranged in paerns on the back side of frond leaflets. In certain fern species, each sorus has a special covering called the indusium that peels back or splits open when the sporangia mature and are ready to shed their spores. The shape of the sorus, the presence of an indusium, the paern and locaon of the sori on the frond are all diagnosc characteriscs for idenficaon. Some ferns have enre fronds or porons of a frond that never develop sporangia and are considered sterile. Fronds that do produce sporangia are ferle and may have a different shape or appearance. Ferns with differing ferle and sterile fronds are considered dimorphic (two forms). In spring, sori are oen very underdeveloped and may be difficult to detect, hindering idenficaon. On young plants or in unfavorable condions, a fern may produce few or no sori. In some leptosporangiate ferns (Cinnamon, Sensive, Royal, Interrupted, Ostrich), the ferle frond (or poron) is greatly modified and oen substanally reduced, amounng to lile more than ghtly grouped clusters of sporangia in place of pinnae (leaflets). Eusporangiate ferns produce one sterile frond and a sporophore which is their equivalent of a ferle frond. Fern spores need a moist to wet environment to complete the plant’s life cycle. Upon germinaon, a single cell structure develops into a mul-celled gametophyte, called a prothallus. At one end of this 1/4-inch, heart-shaped prothallus, male organs called antheridia develop and produce flagellated haploid sperm. At the other end, female organs called archegonia develop, each containing a single haploid egg. In the presence of water, the sperm

41 swims to an archegonium to ferlize the egg. Within three to four months, a new growing fern (the diploid sporophyte) displaces the gametophyte. Archegonia can exude a chemical to aract the sperm. To avoid self ferlizaon, the antheridia and archegonia mature at different mes in some species. The flask shaped archegonia arch away from their own antheridia and somemes secrete chemicals to suppress development of archegonia in nearby gametophytes, reducing compeon and increasing the chance of cross ferlizaon. Eupsporangiate ferns are an excepon. Their gametophytes develop underground usually resulng in self ferlizaon.

Sori of Silvery Glade Fern (Deparia acroschoides)

Fern Idenficaon Along with noted sporangia characteriscs, other morphological traits are important in fern idenficaon. There are two primary growth forms or habits. Those with disnct individual plants are clump-forming and oen vase-shaped, as with the wood ferns (Dryopteris spp.) or Cinnamon Fern (Osmundastrum cinnamomea). Ferns that spread by rhizomes (horizontal, rootlike stems) form colonies, somemes quite large. New York Fern (Thelypteris noveboracensis) and Hayscented Fern (Dennstaeda punclobula) are colonizing ferns. The shape of the frond (primary leaf or blade) may range from narrowly linear (Ebony Spleenwort, Asplenium platyneuron) to large and triangular (Bracken Fern, Pteridium aquilinum). The blade may taper at the p only or at both ends, appearing widest in the middle. Frond branching paerns (number of divisions) vary among species and include an enre or undivided blade (Walking Fern, Asplenium rhizophyllum, and Southern Adder’s-Tongue, Ophioglossum pycnoschum), once-divided or pinnate blade (Christmas Fern, Polyschum acroschoides), twice divided or bipinnate blade (Lady Fern, Athyrium filix-femina ssp. asplenioides), and thrice-divided or tripinnate blade (Bracken Fern, Pteridium aquilinum). Blades that are once-divided or pinnate have mulple small leaflets called pinnae (plural, singular pinna) branching featherlike to either side of the rachis (stem poron within the blade). In twice-divided or bipinnate ferns, each of these pinna is subdivided into smaller leaflets called pinnules. Fern blades or pinnae that are not divided all the way to the blade rachis or pinna midvein are called pinnafid (Broad Beech Fern, Phegopteris hexagonoptera). The resulng lobes are somemes called segments.

42 Leaflet margins for pinnae and pinnules may be smooth, toothed, or lobed. The spe (peole stem below the blade) and rachis (stem secon within the blade) may have hairs or flat scales. Hairs and scales may drop off as the fronds age. As with any organism, habitat oen provides a clue to identy, narrowing the range of possibilies based on species’ preferences regarding soil moisture, light exposure, soil chemical composion, and elevaon.

Gardening with the Nave Plants of Tennessee: The Spirit of Place Margie Hunter, Univ. of Tennessee Press, 2002. Used by permission.

Primary ferns source: (full bibliographical informaon in secon VI. Resources) Ferns of Northeastern and Central North America, 2nd Ed., Boughton Cobb, Elizabeth Farnsworth, and Cheryl Lowe

43 Spermatophytes -- Gymnosperms and Angiosperms Seed-bearing plants evolved from spore-producing plants. The first non-fern seed plants were gymnosperms. The term means “naked seed” and refers to seed development on the exposed surface of reproducve structures such as the scales of a pine cone. In contrast, the seeds of angiosperms, which evolved later, are fully enclosed in the ovary of a flower to form fruit. The success of this enclosed seed development is evidenced in angiosperms’ rapid diversificaon and rise to dominance.

Gymnosperms in Tennessee Of the four plant kingdom divisions represenng gymnosperms, only one -- Coniferophyta, the Conifers (somemes seen as Pinophyta) -- has species nave to Tennessee. In Tennessee, gymnosperms are represented in the following families: • Cypress (Cupressaceae) -- Eastern Red Cedar (Juniperus virginiana), Bald Cypress (Taxodium dischum), Arbor Vitae or White Cedar (Thuja occidentalis) • Pine (Pinaceae) -- Fraser Fir (Abies fraseri), Red Spruce (Picea rubens), two species of hemlock (Tsuga spp.), six species of pine (Pinus spp.) • Yew (Taxaceae) -- Canada Yew (Taxus canadensis) Three of these species -- Fraser Fir, Carolina Hemlock (Tsuga caroliniana), and Canada Yew -- are listed as threatened or endangered species. One pine species, Loblolly Pine (Pinus taeda), has been widely planted outside its normal range for soil stabilizaon, game management, and other commercial interests. Its historical range encompasses only a few counes along the state’s southern border. Except for Canada Yew, a shrub, the remainder are trees. Conifer foliage is narrowly linear, needlelike, or scalelike and remains on the plant longer than one year (evergreen), being shed and replaced gradually throughout the year rather than all at once. Bald Cypress is the lone excepon with annually deciduous foliage. Most Tennessee conifers are monoecious, each tree producing separate male and female cones. Pollen bearing male cones are small and so, dropping aer pollen release. Larger female cones, oen woody and persistent, contain the seeds. Eastern Red Cedar and Canada Yew are dioecious, producing male and female cones on separate plants. Red Cedar and Bald Cypress have berry-like female cones; yews produce individual seeds, each surrounded by an open-ended, fleshy red aril. Gymnosperms are considered ‘sowoods,’ a term that has nothing to do with the actual hardness of the wood, referencing only its ‘naked seed’ development. Woody angiosperms are termed ‘hardwoods.’

Eastern Hemlock (Tsuga canadensis)

44 Angiosperms -- The Flowering Plants There is only one division of angiosperms, the Magnoliophyta (somemes seen as Anthophyta). The vast majority of plants in Tennessee are angiosperms. Besides the unique reproducve structures and processes associated with angiosperms, there are other defining characteriscs that separate them from gymnosperms. There are annual (one-year life span) and biennial (two-year life span) species, however, most are perennial (living a few to many years). They may be herbaceous or woody. Above ground growth on most perennial herbaceous plants typically dies back in winter, but roots are sll alive and send up new growth the following year. Woody species in all spermatophytes retain stems and branches above ground and exhibit secondary growth, the addion of a new outer layer of conducve vascular ssue each year increasing stem diameter. Primary growth is the increase in height, extending the plant’s main axis. Leaves of angiosperms are typically wide and flat (broadleaf). Most woody species are deciduous, dropping leaves in autumn in temperate latudes. As with conifers, evergreen woody angiosperms retain green leaves beyond one year replacing them individually. Broadleaf evergreens are most common in tropical climates. Herbaceous species that retain foliage through winter, such as Liverleaf (Hepaca spp.), Allegheny Spurge (Pachysandra procumbens), and some sedges (Carex spp.), generate fresh foliage in spring.

Celandine Poppy (Stylophorum diphyllum)

45 Monocots and Dicots Magnoliophyta is divided into two classes, represenng the main types of flowering plants: Liliopsida, the monocots, and Magnoliopsida, the dicots. The primary differences are outlined.

Characterisc Dicot Monocot

embryo two cotyledons one cotyledon

seed leaves two seed leaves one seed leaf

leaf venaon veins reculated or branched veins parallel (start and end (need) at a common point)

flowers parts in mulples of four or five parts in mulples of three

secondary growth oen present absent

herbaceous/woody herbaceous and woody herbaceous only

stem vascular system vascular bundles arranged in a ring vascular bundles scaered throughout stem

pollen grain surface three furrows or pores one furrow or pore

roots taproot fibrous roots

Among the monocot families are grass, orchid, lily, onion, spiderwort, sedge, iris, caail, arum, rush, and bunchflower, which includes trilliums. There are many excepons to the typical disncons between monocots and dicots. For example, veins in the leaf-like bracts of trilliums are branched rather than parallel. Dicot families include buercup, rose, heath, aster, mustard, mint, pea, and many others. Just as with monocots, there are a few dicot species with certain traits more characterisc of the other class. Recent DNA study has shown that these two groupings do not represent plants’ actual evoluonary development. It’s a bit more complex, parcularly with dicots. There are now four groups -- Basal Angiosperms, Magnoliids (a type of dicot), Monocots, and Eudicots (‘true’ dicots).

46 Crested Iris (Iris cristata), monocot

Sourwood (Oxydendrum arboreum), dicot

Reproducon in Spermatophytes Unlike mosses, liverworts, and ferns with their flagellated male sperm cells, spermatophytes do not need water as a medium to facilitate ferlizaon. Gymnosperms rely on wind to carry pollen grains, each containing the male sperm cells, to the female eggs within the scales of cones. Specialized staminate strobili, referred to simply as male cones, produce pollen which is released in vast quanes for the hit or miss possibility of finding a female cone of that species. Angiosperms somemes use wind, but most have adapted a more targeted approach, aracng a variety of other organisms with real (or imagined) benefits to pick up and move

47 pollen from one individual to the female reproducve organs of another. This efficient ferlizaon mechanism requires less pollen, improves success rate, and assists genec diversity. Spermatophytes produce two types of spores -- microspores (male) and megaspores (female). [Ferns and mosses just have one spore type.] Microspores develop into microgametophytes or pollen. In angiosperms, this occurs within the anthers of a flower. Megaspores develop into megagametophytes within an ovule. The simplified essence of the reproducve process in both gymnosperms and angiosperms leads a sperm cell from the pollen grain to ferlize the megagametophyte’s egg, which develops into an embryo, the next sporophyte generaon for seed plants. Each ovule in gymnosperms sits exposed on the surface of a scale in the cone. The ovule in angiosperms is enclosed within a special structure called a carpel or pisl. It consists of a sgma (scky top to catch pollen grains), style (a connecng neck-like tube), and an ovary or chamber within which ferlizaon and development of the seed takes place producing fruit. Any seed, whether from a gymnosperm or angiosperm, will have an embryo, endosperm, and seed coat. The seed coat protects the young embryo unl condions are right for germinaon. Seeds may have to experience warm or cold periods (straficaon) or be exposed to some chemical or mechanical process to break down the seed coat (scarificaon) to spur germinaon. Using the endosperm as a food and energy source, the embryo begins growth by sending a root (radical) into the soil. Seed leaves (cotyledons) of the green shoot emerge and begin photosynthesis. The seedling transions from endosperm nutrion to manufacturing its own food and produces the first true leaves.

Plant Morphology Morphology entails the physical form and structure of an organism. Technical terms are usually given to an organism’s physical parts and/or characterisc traits. Botany is blessed (or burdened) with a wealth of specialized terminology. Efforts to idenfy and/or research plants will likely expose naturalists to many of these terms. There are dozens of surface characteriscs, innumerable specialized structures, and technical jargon from acnomorphic (radially symmetrical or regular flower) to zygomorphic (bilaterally symmetrical flower). A good glossary is indispensable. Most floras and field guides contain a glossary of terms used. Naturalists and amateur botanists may find the following book helpful -- Plant Idenficaon Terminology: An Illustrated Glossary, 2nd Ed., James G. Harris and Melinda Woolf Harris. Angiosperms display many different growth forms -- woody trees, shrubs, and vines and herbaceous vines, grasses, and other flowering plants. Trees typically develop a single trunk and grow over 12 feet tall. Shrubs are mul-stemmed and remain under 15 feet. Woody vines (lianas) and non-woody vines develop long stems that creep or climb over other plants for support in their quest to reach sunlight using twining stems or leaf peoles, clinging rootlike structures, tendrils, holdfast pads, or thorns. Herbaceous plants include grasses and grass-like sedges and rushes with long, narrow leaves and flower-producing herbs in a variety of sizes and shapes. These non-grass herbaceous species are called forbs.

48 Gardening with the Nave Plants of Tennessee: The Spirit of Place Margie Hunter, Univ. of Tennessee Press, 2002. Used by permission

The illustraon above presents a very simplified view of a typical angiosperm’s flower. There are many variaons on this arrangement and more detailed structures among the diversity of flowering plants. ‘Flowers’ on certain species, such as grasses or sedges, differ significantly. In a complete flower, four whorls or series -- sepals, petals, stamens, and pisl(s) -- are present; incomplete flowers lack one or more series. Sepals (calyx) and petals (corolla) comprise the perianth and are sterile. Stamens (anther, filament) produce pollen, the male gametophyte. The pisl or carpel (sgma, style, ovary) is associated with producon of the female gametophyte. Flowers containing both male and female reproducve structures are said to be perfect. Unisexual flowers (imperfect) have either male structures (staminate flowers) or female structures (pisllate flowers). If both staminate and pisllate flowers are present on the same plant, the species is monoecious (‘one house’). If the differently sexed flowers are only found on separate plants, as with hollies (Ilex spp.), the species is dioecious (‘two houses’). Before DNA studies, plant classificaon was based primarily on physical characteriscs of the flower. These characteriscs are sll among the easiest to use in idenfying species in the field and can point toward a parcular family, facilitang idenficaon. A good understanding of flower morphology is an important skill in field work. Not every individual of a species will look idencal to the next. Besides genec diversity from sexual reproducon, an organism’s environment can influence its appearance. This quality has a 50-cent term -- phenotypic plascity -- the ability of any given genotype (set of genes) to produce different phenotypes (outward physical appearance) in response to varied environmental condions, e.g., too much or too lile sunlight, not enough water, poor soil, etc. This is not an important concept to know. Just one to keep in mind when a plant doesn’t seem to ‘fit’ all aspects of its official descripon.

49 Gardening with the Nave Plants of Tennessee: The Spirit of Place Margie Hunter, Univ. of Tennessee Press, 2002. Used by permission.

The posion and arrangement of flowers on the plant are important features to note. Flowers may be solitary or occur in a cluster (inflorescence). Clusters can be arranged in several different forms, broadly rounded to flat umbels, corymbs, and cymes; tall, narrow spikes and racemes; and open to dense panicles among other types. Flowers may appear at the p of the stem (terminal) or in the juncon of stem and leaves (axillary). The shape, margin, p, base, venaon, division, aachment, and arrangement of leaves have many more opons than presented in the graphic here, each with its own special term, as well as leaf or stem surface textures and characteriscs. There are several types of underground stem structures -- bulbs, corms, tubers, rhizomes, stolons -- that form roots and store food. Unique twig characteriscs permit idenficaon of woody deciduous shrubs and trees in winter based on shape, color, buds, leaf scars, and other traits. [See TNP forests study guide.]

50 Gardening with the Nave Plants of Tennessee: The Spirit of Place Margie Hunter, Univ. of Tennessee Press, 2002. Used by permission.

51 The ripened ovary containing mature seeds constutes a plant’s fruit. Fruits may be simple (single flower with one pisl, holly, tomato, peach), aggregate (single flower with many pisls, raspberry, magnolia), or mulple (many flowers fuse in a ght cluster, mulberry, sweetgum, Osage orange). There are a variety of fruit types, oen with characteriscs disncve enough to idenfy plants to species. Fleshy fruits include berries, drupes, and pomes. Fruits derived from the ripened ovary and other floral ssue, such as strawberry or apple, are accessory or ‘false fruits.’ The strawberry has dry fruits (achenes) embedded in the expanded, ripened flesh of the receptacle. Dry fruits that open at maturity (dehiscent) to expose the seeds include the follicle, capsule, legume, loment, silicle, and silique. Other dry fruits do not open (indehiscent), such as achene, cypsela, caryopsis, nutlet, nut, schizocarp, and utricle. Some botanical differences are very slight, requiring close observaon of detail. It is oen these small details, the presence of minute hairs for example, that separate one species from another. Because differences can be subtle, it is important to look at more than one leaf, flower, fruit, or twig to be certain observed traits are representave. A malformed leaf or flower that has dropped a petal could affect idenficaon. A quality 10X hand lens is indispensable.

Plant Ecology

Plants make life possible on this earth. Their ability to harness energy, manufacture food, release oxygen, and moderate the environment are essenal contribuons to the success of other life forms. As primary producers, plants are the first link in the grazing food chain and a significant contributor to the detrital food chain. They play key roles in biogeochemical cycles (oxygen, carbon, water, nutrients), influence soil formaon, moderate soil and air temperature, slow the flow of surface water, and filter pollutants from the environment. Plants evolved symbioc relaonships with many other organisms, typically exchanging their food producon capabilies for important services, such as improved water and nutrient uptake (mycorrhizal fungi), cross ferlizaon (pollinators), and seed distribuon (insects, birds, and mammals). Wildlife depend on plants for food, shelter, predator protecon, and nesng sites. Not all plants manufacture their own food. Plant parasites tap into the roots or stems of other plants for water and nutrion. They have special organs called haustoria, which are modified roots, to draw nourishment from hosts. There are two types. Holoparasic plants have lile to no chlorophyll and infiltrate both the phloem and xylem of the host to derive all organic and inorganic nutrients, Dodder (Cuscata spp.) is an example. Hemiparasic plants only infiltrate the host’s xylem, accessing water and mineral nutrients but lile carbon. They have green leaves to photosynthesize their own food but a reduced root system; examples are Buffalo Nut (Pyrularia pubera) and Mistletoe (Phoradendron seronum) among others in the sandalwood family (Santalaceae). Orobanchaceae, the broomrape family, contains both types including holoparasites Squawroot (Conopholis americana on oak roots), Beechdrops (Epifagus virginiana on beech roots), and One-flowered Cancer Root (Orobanche uniflora on asters, stonecrops, and

52 saxifrage), and hemiparasites False Foxglove (Agalinis spp. and Aureolaria spp.), Indian Paintbrush (Caslleja coccinea), and Wood Betony (Pedicularis canadensis) on grasses. Some plants are mycotrophs, living in part or enrely as parasites on mycorrhizal fungi plus a few saprophyc fungi species as well. All orchids need these fungi to survive as seedlings and in periods of dormancy, and some species such as Coralroot orchids (Corallorhiza spp.) exist enrely on the fungi. Members of the Indian pipe family (Monotropaceae) and a few species in the heath (Ericaceae), diapensia (Diapensiaceae), and genan (Genanaceae) families obtain part or all of their nutrional needs from the food these fungi receive in their mutualisc partnerships with forest trees.

Habitat and Idenficaon A plant’s habitat oen provides clues to assist idenficaon. Conversely, idenfying plants can provide clues to the habitat.

Wetlands -- The federal definion of wetlands is “those areas that are inundated or saturated by surface or groundwater at a frequency and duraon sufficient to support, and that under normal circumstances do support, a prevalence of vegetaon typically adapted for life in saturated soil condions. Wetlands generally include swamps, marshes, bogs and similar areas." The Naonal Wetland Plant List compiled by the U.S. Army Corps of Engineers, Fish and Wildlife Service, Environmental Protecon Agency, and Natural Resources Conservaon Service is used to make wetland determinaons as part of the Clean Water Act and Naonal Wetland Inventory. Plants are categorized based on the likelihood their presence indicates the area is a wetland. These codes appear in floras and other plant lisngs such as USDA PLANTS Database Web site. The United States is divided into seven wetlands regions. West Tennessee is Atlanc and Gulf Coastal Plain Region; Middle and East Tennessee are Eastern Mountains and Piedmont Region. Aquac species (hydrophytes) grow submerged or at the water’s surface and may be rooted (Nelumbo lutea, American Lotus) or free floang (Lemna spp., Duckweed).

Indicator Code Indicator Status Meaning

OBL Obligate Wetland Almost always occurs in wetlands

FACW Facultave Wetland Usually occurs in wetlands 67 to 99 percent of the me; may occur in non-wetlands

FAC Facultave Equally likely to occurs in wetlands and non- wetlands

FACU Facultave Upland Usually occurs in non-wetlands 67 to 99 percent of the me; may occur in wetlands

UPL Obligate Upland Almost never occurs in wetlands

53 Topography and Elevaon -- Elevaons above 4,000 feet are usually accompanied by changes in climate sufficient to alter the plant community composion. Northern hardwood species (e.g., Yellow Birch [Betula alleghaniensis], Mountain Maple [Acer spicatum], Witch Hobble [Viburnum lantanoides]) begin to appear, and above 5,500 feet boreal remnants such as Fraser Fir (Abies fraseri) and Red Spruce (Picea rubens) are found. Herbaceous plants preferring higher elevaons include Mountain Wood Fern (Dryopteris campyloptera), Rugel’s Ragwort (Rugelia nudicaulis) and Roan Mountain Bluet (Houstonia purpurea var. montana). At lower elevaons, topographical features influence community composion too. Steeper slopes, south facing slopes, and ridge lines support more drought tolerant species and those preferring a more acidic soil, such as oaks, pines, hickories, and Mountain Laurel (Kalmia lafolia). North facing slopes are cooler, moister and support a greater diversity of plants.

Soil Chemistry -- Soils reflect the chemical composion of associated bedrock, plant communies, and the accumulaon or leaching of soil alkalis due to rainfall amounts and drainage. Plants that prefer less acidic to neutral soil or can tolerate alkalinity are called calciphiles. Their presence indicates a higher pH soil and underlying rock formaons, such as limestone, needed to produce that soil condion. Near Cades Cove in the Great Smoky Mountains Naonal Park, the presence of limestone is indicated from the appearance of plants not oen noted in the park -- Redbud trees (Cercis canadensis) and Purple Clirake fern (Pellaea atropurpurea) on Rich Mountain, Wild Blue Phlox (Phlox divaricata) and Shoong Star (Dodecatheon meadia) in White Oak Sink, Narrowleaf Glade Fern (Diplazium pycnocarpon) along Schoolhouse Gap Trail. In contrast, calcifuge plants do not tolerate a high pH, suffering from the lack of soluble iron in more basic soils. They prefer acidic soils and include many members of the heath family -- blueberries (Vaccinium spp.), Mountain Laurel (Kalmia lafolia), Trailing Arbutus (Epigaea repens), and Pipsissewa (Chimaphila maculata), plus others like Galax (Galax urceolata), Cinnamon Fern (Osmundastrum cinnamomea), and Painted Trillium (Trillium undulatum). These plants are most oen found on soils derived from more acidic rocks of the Cumberland Plateau or Blue Ridge Mountains or upland ridges of the Highland Rim, where rainfall leaches alkalis from the soil resulng in lower pH. Plants affect soil and its chemistry through nutrient cycling, enhancing ferlity, adding organic content, and modifying temperature and moisture. Not the passive inhabitants they appear, some plants produce chemicals and release them into the soil to inhibit growth of nearby species including their own offspring. Termed allelopathy, this ability reduces compeon for resources. Black Walnuts (Juglans nigra) are well known for this trait, producing the chemical juglone. Plant species sensive to juglone will grow poorly if at all. Seed germinaon and seedling growth are reduced. Not all plants are suscepble. Grasses, for example, grow well around Black Walnuts. Some invasive pest plants are allelopathic, using this chemical advantage to displace nave species and overrun habitats.

54 Cultural Uses Plants’ praccal uses range from elemental, e.g., food, shelter, and clothing, to medicines, dyes, and ornaments. Ethnobotany is the scienfic study of tradional plant usage by various cultures throughout human history. From ancient mes to modern medicine, herbalists have looked for plants to treat different maladies. In the sixteenth and seventeenth centuries, a philosophy known as “The Doctrine of Signatures” became popular. If some physical component of a plant resembled parts of the human anatomy, symptoms of a disease, or the cause of a medical problem, then this plant was considered a good treatment for any associated illness. Plants with liver-shaped leaves could treat liver ailments. Plants with long straight flower stalks that resembled snakes or had raling seedpods would treat snakebite. Red juice in the roots of Bloodroot (Sanguinaria canadensis) indicated its use for blood disorders. The basic concepts driving the doctrine were ‘like cures like’ and the belief that God provided the means to cure any disease and marked it with a ‘sign.’ Many plant common names derive from this idea, oen pairing the relevant part of the body with the old English word for plant, “wort” -- liverwort, lungwort, spleenwort, toothwort. Most have proven to be of lile or no value. Plants of the Cherokee by William H. Banks (Great Smoky Mountains Associaon, 2004) details the tribe’s medicinal usage for dozens of plant species. The recommended field guide for this class, Wildflowers of Tennessee, the Ohio Valley, and Southern Appalachians by Horn and Cathcart, oen cites tradional uses at the end of plant descripons.

Sharp-lobed Liverleaf (Hepaca aculoba)

55 IV. Dichotomous Keys

Species idenficaon does not have to be a guessing game, flipping through pages of a field guide in hopes of finding a suitable match. A well-developed dichotomous key achieves faster, more accurate results. Keys are reference tools, highly organized wrien devices designed to work progressively through specified physical traits and arrive at the correct idenficaon. Dichotomous keys are arranged in a numbered series of paired statements or couplets. Each couplet describes the same plant feature(s), and each statement in that couplet stands in direct contrast to the other, presenng mutually exclusive, observable traits. Selecon of the most applicable statement directs the reader to another numbered couplet. This process is repeated unl an idenficaon is reached. The process systemacally eliminates what the organism is not, narrowing the possible alternaves unl the correct species is idenfied. Keys can be organized on different scales, from a single family to enre vascular plant floras. They may be restricted to a parcular geographic region, such as a physiographic province, or focus on a parcular set of seasonal or character traits, such as winter twigs or fruit. Plant keys have one drawback: characteriscs used to separate species, for example flowers or fruits, may not be present at a given me thus impeding idenficaon.

Guidelines for Using Dichotomous Keys Select an appropriate key for the area -- Make sure the key being used covers the organism or its geographic area. Some keys are limited in their coverage. Vascular floras will not include bryophytes (nonvascular plants). Kentucky’s vascular flora may not help in Tennessee’s Blue Ridge Province, as this physiographic area is outside that state’s borders. Understand terminology -- Some keys may be highly technical, and success depends on accurate interpretaon of couplet statements. Do not guess. Most keys have a glossary. Use it. Read both statements of a couplet before making a choice -- Sounds obvious, but it is oen tempng to go with the first statement if it seems a good match. The second statement, however, may be a beer match. Examine more than one specimen -- Each leaf, flower, or plant may not be idencal to others of its kind. Insect damage, malformaon, or age can alter the appearance of a specimen. Before working the key, make sure the specimen is truly representave of the species. Try both choices if the answer is not clear -- Depending on how well the key is wrien, the season to which it is geared, the decisiveness of the character trait exclusion, or the quality of the specimen, it is possible to arrive at a couplet where either choice could work. Follow each lead. At some point, an inconsistent trait will likely exclude the incorrect path. Verify the identy -- Once an idenficaon is made, it may be desirable to confirm it by checking other sources. Field guides, online herbaria, full species descripons, drawings or photographs, etc., offer visual and verbal comparisons of a full range of characteriscs to help verify the organism’s identy. Keep at it -- Keys can be tricky and discouraging at first. Pracce does help. Try keying known species to get a feel for the process.

56 V. Invasive Pest Species

Review of Nave Ecosystems A review of nave ecosystems’ evoluon and funcon will provide a framework to beer understand how certain non-nave species become invasive. Nave organisms evolved with the land, climate, and each other developing interwoven relaonships through geologic me. Each species’ role has been refined within the landscape’s biological communies, and a system of checks and balances helps establish a relavely stable equilibrium. Many species develop unique adaptaons to local condions, and all play an integral role in the locality’s funcon and identy.

Species’ Movement All organisms are nave somewhere. They developed in their communies and landscapes through the processes described above. The natural movement of most organisms is typically slow and deliberate. This pace allows the organism to adapt to changing environmental condions and allows the environment to adjust to the presence of the organism. In Tennessee, nearly 2,900 species of vascular plants have been documented. Approximately 2,400 are nave, and the remainder are introduced from other areas around the world. As humans have moved around the world, they have deliberately or inadvertently brought species from other connents. Insects arrived as hitchhikers on plants or in other organic materials. Some have been brought in for specific research or commercial applicaons and escaped. Animal species are oen imported as exoc pets. Non-nave plants have served a wide range of uses from ornamental horculture to animal forage and soil stabilizaon. These introducons preclude evoluonary adjustments, leaving three possible outcomes. The organism struggles and dies in its new environment, survives under acve management, or adapts to live independently. Those that live independent of management become naturalized, successfully growing and reproducing in the wild. The laer outcome can result in a non-nave species becoming too successful and spreading to the detriment of nave species.

Mechanics of Invasion Non-nave species have the potenal to become invasive because their co-evolved biological checks and balances are missing in the new environment. Compeon from other species nave to their home range, climate limitaons in that range, and the predators, pests, and diseases that kept their populaons in check are no longer factors. The checks and balances for nave species, on the other hand, are sll in place. To make maers worse, nave species have no evolved defenses to effecvely compete with some of these newcomers. These advantages favor the non-nave species. If these organisms are equipped to reproduce or spread aggressively, they can quickly outcompete nave species for territory or resources.

57 Characteriscs of Invasive Plants Non-nave plants capable of surviving on their own in a new environment are at greater risk of becoming invasive if they exhibit one or more of the following traits. • Rapid growth -- quickly grows in size • Early flowering maturity -- matures to produce flowers (and fruit) at an early age • Copious seed producon -- produces viable seeds up to thousands or millions per plant • Effecve seed dispersal -- efficient dispersal through wind, water, or animal vectors • Rampant vegetave spread -- quickly colonizes and overspreads its surroundings

Non-nave Invasive Species In 1999, Execuve Order 13112 signed by President Clinton, established the Naonal Invasive Species Council, proscribed its dues, and set a legal definion for non-nave, invasive species: “Any species, including its seeds, eggs, spores, or other biological material capable of propagang that species, that is not nave to that ecosystem; and whose introducon does or is likely to cause economic or environmental harm or harm to human health.” Federal, state, and local government agencies, private landowners, and conservaon organizaons oen work cooperavely to idenfy pest species and take measures to eradicate or control populaons that threaten farm and rangelands, public lands, wildlife, or waterways.

Disrupt Nave Biological Systems Non-nave invasive species disrupt and harm nave biological systems in a variety of ways. Each disrupon ripples through the ecosystem causing further harm like falling dominoes. Non- nave invasive species are not simply out-of-balance, they are also nonfunconing, contribung lile to important ecosystem services. Overrun a variety of habitats -- Invasive species are not restricted to disturbed sites like roadways and abandoned lots. Through seeds spread by water, wind, and animals, they can invade undisturbed, prisne habitats. The fragmented nature of wild or undeveloped land increases this opportunity. Some habitats evolved in conjuncon with periodic disturbance to rejuvenate or maintain a certain community. These systems are especially vulnerable. Displace or destroy nave populaons -- Once invasive species get the upper hand, they can quickly outcompete nave organisms, occupying space, claiming sunlight, nutrients, and water, eang food sources, or preying on nave species. Enre groups of nave species may be displaced. Deprive dependent nave organisms of needed food sources, etc. -- Without their evoluonary partners, many organisms are deprived of proper nutrion, adequate shelter, or nurturing environments. Many insects do not recognize non-nave plants as edible or lack evolved digesve enzymes to derive nutrion. Without insects, parent birds have nothing to feed their young. Fruits of non-nave plant species may be eaten, but they oen lack the necessary nutrion to maintain proper wildlife health. Disrupt plant/animal associaons -- Communies carefully constructed through the paent wheel of evoluon are quickly torn to shreds. As nave plants are overrun by non-nave plant

58 species, wildlife dependent on nave species must go elsewhere as well. The enre dynamic of funconal roles, predator and prey, and energy transfer through the system weakens. Significantly reduce plant and wildlife diversity -- When a non-nave plant species invades a system, it can indiscriminately overrun and replace scores of nave species, somemes resulng in a near monoculture. As plant diversity drops, so does wildlife diversity. Significantly reduced diversity hobbles community funcon, resulng in a biological wasteland. Stress rare and endangered plants and animals -- Various threats such as polluon, development, and fragmentaon undermine populaons, placing organisms at risk of exncon. The impact of invasive species is now regarded as second only to habitat destrucon as a threat to endangered species. According to NatureServe, of the 40 North American freshwater fishes that have become exnct during the past century, invasive species were a contribung factor in more than two-thirds of those exncons. Hybridize with nave species to alter genecs -- Some nave plants are related to species from other connents. This relaonship could allow the species to cross ferlize. Genec contaminaon of nave stands and inherited adaptaons for non-naves are two negave consequences. Support non-nave pathogens and pests -- Non-nave plants have also introduced some of their insects and diseases, which can have dire consequences on the health of nave species not adapted to such organisms, as have Hemlock Woolly Adelgid and Chestnut Blight. Alter ecosystem processes -- (1) Fire - Non-nave species may burn hoer and more destrucvely to damage even fire-adapted communies. (2) Water - Non-nave species may overrun seasonal wetlands changing the area’s hydrology which could affect seasonal breeding habitat for amphibians. (3) Soil - Non-nave species may alter soil chemistry inhibing germinaon or growth of nave species. (4) Predaon - Non-nave species may decimate nave populaons as prey, altering community makeup and funcon (food web, pollinaon, seed dispersal).

Japanese Sltgrass (Microstegium vimineum) Chris Evans, River to River CWMA, www.invasive.org

59 The Scope and Impact Not all non-nave species are invasive or present major ecological problems. The effect of most is benign. For those that do pose an invasive threat, the degree of invasiveness varies. In Tennessee, plants are ranked as Severe, Significant, or Lesser Threats, plus a watch list, by the Tennessee Exoc Pest Plant Council. A plant’s invasive potenal changes with environmental factors such as moisture, climate, soil, etc. Species may pose a problem in one area of the country or state but not another. Naonal costs associated with invasive plant species (control, crop and land losses) were esmated at $35 billion annually (Pimentel et al. 2004). Areas must typically be treated more than once, require regular monitoring for new and recurring populaons, and oen need restoraon work to return nave species. Any disturbance opens the door for major infestaons. In the Great Smoky Mountains Naonal Park, an intense 160-acre fire burned much of the forest leaving an open, mineral soil in August 2010. Well-established populaons of Princess Tree (Paulownia tomentosa), a TN-EPPC ranked Severe Threat invasive species, on adjacent public and private land provided ample seed sources. Between 2011 and 2014, the Naonal Park Service dedicated 943 work hours to the removal of 96,137 non-nave Paulownia seedlings and saplings at a minimum labor cost of $14,000. All levels of aquac and terrestrial systems are threatened in some way by non-nave invasive species. Tennessee may not have Burmese Pythons like Florida, but European Wild Boars are mean, destrucve animals in state forests. They represent just one species in a long list of invaders that include Zebra Mussels (Asia), Purple Loosestrife (Europe), Balsam Woolly Adelgid (Europe), European Starlings, Fire Ants (South America), chestnut blight (Asia), Emerald Ash Borers (Asia), beech bark disease (Europe), dogwood anthracnose (origin unknown), Asian Carp, Kudzu (Asia), and many, many more.

Mimosa (Albizia julibrissin) James H. Miller, USDA Forest Service, www.invasive.org

Pimentel, D., R. Zuniga, and D. Morrison. Update on the Environmental and Economic Costs Associated with Alien- Invasive Species in the United States. Ecological Economics 52 (2004): 273-288.

60 VI. Management

Plant management must be approached on a community basis, taking into account the geological, geographical, and ecological characteriscs.

Natural Areas

In 1971, Tennessee passed the Natural Areas Preservaon Act establishing the state’s Natural Areas Program. The General Assembly has since designated 83 State Natural Areas represenng intact ecosystems that demonstrate the funcon of natural ecological processes. The most unmodified and unique areas are managed for research. The rest include varying degrees of public recreaon. Oen other local, state, and federal agencies and non-governmental organizaons are involved in site management. Addionally, there are 30 Registered State Natural Areas established through non-binding, voluntary agreements with private and public landowners protecng ecologically important sites. Tennessee also has thirteen Naonal Natural Landmarks overseen by the Naonal Park Service. Six are Designated State Natural Areas, and one is a Registered State Natural Area. According to the Tennessee Department of Environment and Conservaon, Division of Natural Areas webpage, “The Natural Areas Program seeks to include adequate representaon of all natural communies that make up Tennessee's natural landscape, and provide long term protecon for Tennessee's rare, threatened and endangered plant and animal life.” Rare plants in protected natural areas are monitored and managed according to their life histories -- habitat needs, growth, and reproducon. Some of these communies historically evolved in conjuncon with various natural disturbance processes. These large and small-scale disturbances, such as fire, flooding, or river scour, maintain or periodically reintroduce a set of community condions supporve to certain species by removing compeon, opening the canopy, promong seed germinaon, or enriching soils. Management goals in natural areas seek to protect or restore the occurrence of such processes and, where appropriate, provide substutes. For communies that have a known dependence on or adaptaon to fire, periodic prescribed burns may be used to iniate renewal and regeneraon. Over me, the lack of fire in such communies allows successional species to dominate and change the character of the community. Fire suppression in the Smokies has led to the populaon decline of Table Mountain Pine (Pinus pungens), whose cones require the heat of a forest fire to release seeds for germinaon. Any control or management measures employed for plant succession, habitat, or natural populaons must be based on scienfic evidence from observaon and study according to each natural area’s master plan. The establishment of buffer zones surrounding each natural area allow for firebreaks and service areas (parking, facilies) and minimize external influences from nearby development, etc.

61 Invasive Plants

Effecve management of invasive plant species must be preceded by research and observaon to understand each species’ life history, including its habitat preferences and limitaons, growth paern and reproducve cycle, seed life in the soil (seed bank), and pest and disease vulnerabilies. Invasive plant control may take several forms. • Biological control -- living organisms as means of reducing invasive species populaons through predaon. Animals (mammals and invertebrates) to browse or defoliate and pathogens (viruses, bacteria, fungi) through parasism or disease introduce checks and balances to minimize disrupons to nave ecosystems. Though research is done before the introducon of a non-nave control, unforeseen consequences to non-target species remain a possibility. • Mechanical control -- physical removal of a plant through hand pulling, digging, cung, girdling, mowing, mulching, bulldozing, etc. This method works best with light infestaons. It is a more targeted approach, but soil disrupon and high labor costs can be disadvantages. • Chemical control -- herbicides kill plants through foliar sprays, cut stump treatments, stem injecons, or basal bark applicaons. Chemicals are toxic, material costs including equipment and labor are high, and training is required. Chemical control works for large infestaons, but depending on treatment method, there is the potenal to damage or kill non-target plant species. There are also methods of cultural control. • Prescribed Fire -- The use of spot burning and wider-area prescribed ground fires can kill some invasive species, though growth or germinaon of others may be smulated. There are significant pros and cons associated with prescribed burns relave to invasive plant control. Knowledge of longterm fire effects on the community and species’ responses are crical. • Water Level -- In areas where water levels can be manipulated, flooding can kill species unable to withstand submergence or saturated soil, and water drawdowns can expose aquac species to drying condions or allow effecve herbicide treatment. • Seed Bank -- Soil solarizaon and mulching can kill or minimize seed germinaon.

Early Detecon and Rapid Response (EDRR) Early detecon of an infestaon opmizes chances for successful eradicaon using the least damaging control opons. It minimizes disturbance to the natural community, which in turn suppresses the opportunity for the invasive plant or other non-nave species to establish. EDRR is oen an important component of Cooperave Weed Management Areas (CWMA). An associaon of landowners, land managers, and local, state, and federal partners, a CWMA develops prevenon and management plans for a specified geographical area and an agreement regarding roles, finances, and other issues among the partners. CWMA plans typically include weed surveying and mapping components as well as integrated weed

62 management, and some may include educaon and training, EDRR, monitoring, revegetaon, and annual evaluaon and adaptaon of the weed management plan.

Mapping - Early Detecon and Distribuon Mapping System (EDDMapS) Mapping the locaons of invasive plant infestaons provides a more accurate picture of the species’ spread. These map data can be used to rank species’ invasive potenal and recommend problemac species for state regulaon through the Tennessee Department of Agriculture’s Pest Plant Rule 0080-06-24. Mapping can be done by individuals online or with mobile applicaons such as SEEDN (Southeast Early Detecon Network). hp://www.eddmaps.org/ hp://apps.bugwood.org/seedn.html

Prevenon The best tool is prevenon. USDA’s Animal and Plant Health Inspecon Service (APHIS) monitors import and export of biological organisms. Their Plant Protecon and Quaranne (PPQ) “regulates the importaon of plants and plant products under the authority of the Plant Protecon Act. PPQ maintains its import program to safeguard U.S. agriculture and natural resources from the risks associated with the entry, establishment, or spread of animal and plant pests and noxious weeds.” There are similar regulatory protecons for the importaon of animal species. hp://www.aphis.usda.gov/wps/portal/aphis/home/

What You Can Do

• Remove invasive plant species on personal property. • Volunteer for invasive plant pulls in public parks and natural areas. • Assist mapping documentaon of invasive species through EDDMapS. • Clean shoes, clothing, pets, or gear aer vising an infested area to minimize unintenonal spread of seeds, etc. • Encourage local gardeners and nurseries to avoid problemac invasive plant species. • Encourage the use of nave species in private and public landscape planngs. • Volunteer to assist rare plant monitoring. • Support public and private natural areas and other habitat conservaon efforts. • Volunteer with nave revegetaon and restoraon projects.

63 VII. Resources

Fungi

Mushrooms of the Southeastern United States Alan E. Bessee, William C. Roody, Arleen R. Bessee, Dail L. Dunaway Syracuse University Press, 2007 Mushrooms of West Virginia and the Central Appalachians William C. Roody, University Press of Kentucky, 2003 The Kingdom Fungi: The Biology of Mushrooms, Molds, and Lichens Steven L. Stephenson, Timber Press, 2010 The Fih Kingdom (Third Edion) Bryce Kendrick, Focus Publishing, 1992, 2000 Images and info on North American species -- hp://www.mushroomexpert.com/ WWW Virtual Library: Mycology -- hp://mycology.cornell.edu/ Comprehensive list of internet mycological resources, includes secon on teaching and learning fungi Tom Volk’s Fungi -- hp://bot.botany.wisc.edu/toms_fungi/ General mycology info Fungi Images on the Net -- hp://fungi.fvlmedia.dk Metadirectory locate/view 3500 photos General informaon on fungi – hp://faculty.clintoncc.suny.edu/faculty/michael.gregory/files/bio%20102/ bio%20102%20lectures/fungi/fungi.htm Online copy of M.C. Rayner’s out-of-print book, Trees and Toadstools, Rodale Press, 1947. Documents early discovery of mycorrhizal associaons. hp://journeytoforever.org/farm_library/rayner/rayner_intro.html Cumberland Mycological Society -- hp://www.cumberlandmycology.com/index.htm

Lichens

Lichens William Purvis, Smithsonian Books, 2000 Lichens of North America Irwin M. Brodo, Sylvia Duran Sharnoff, Stephen Sharnoff; Yale University Press, 2001 Lichens of North America hp://www.sharnoffphotos.com/lichen_info/index.html Lichen Links and Resources, US Forest Service hp://www.fs.fed.us/wildflowers/interesng/lichens/resources.shtml An overview of the biology of lichens by Frank Bungartz, Arizona State Univ. hp://nhc.asu.edu/lherbarium/lichen_info/index.php

64 Mosses and Liverworts

Common Mosses of the Northeast and Appalachians Karl B. McKnight, Joseph R. Rohrer, Kirsten McKnight Ward, and Warren J. Perdrizet Princeton Field Guide, Princeton University Press, 2013 A Trailside Guide to Mosses and Liverworts of the Cherokee Naonal Forest Paul G. Davison (with Mark J. Pistrang) 2008 hp://www.blurb.com/bookstore/detail/422248 Outstanding Mosses and Liverworts of Pennsylvania and Nearby States Susan Munch, Albright College, 2006 [email protected] Gathering Moss: A Natural and Cultural History of Mosses Robin Wall Kimmerer, Oregon State University Press, 2003 Bryophytes info and images, Southern Illinois Univ. Carbondale hp://bryophytes.plant.siu.edu/index.html

Vascular Plants

Guide to the Vascular Plants of Tennessee Edward W. Chester, B. Eugene Wofford, Joey Shaw, Dwayne Estes, and David H. Webb, Eds. (Addional authors: Claude Bailey, Andrea Bishop, Hal DeSelm, Dennis Horn, Chris Fleming, Aaron Floden, William Marn, Mary Priestley, and Ed Schilling) University of Tennessee Press, 2015 A Fih Checklist of Tennessee Vascular Plants Edward W. Chester, B. Eugene Wofford, Dwayne Estes, and Claude Bailey BRIT Press, 2009 hp://www.brit.org/brit-press/books/sbm-31/ Guide to the Trees, Shrubs, and Woody Vines of Tennessee B. Eugene Wofford and Edward W. Chester, University of Tennessee Press, 2002 Plant Life of Kentucky: An Illustrated Guide to the Vascular Flora Ronald L. Jones, University Press of Kentucky, 2005 Plant Idenficaon Terminology: An Illustrated Glossary [2nd Ed.] James G. Harris and Melinda Woolf Harris, Spring Lake Publishing, 2001 Botany in a Day: The Paerns Method of Plant Idenficaon Thomas J. Elpel, HOPS Press, 2004 Manual of Vascular Plants of Northeastern United States and Adjacent Canada Henry A. Gleason and Arthur Cronquist, The New York Botanical Garden, 1991 Illustrated Companion to Gleason & Cronquist’s Manual Noel H. Holmgren, The New York Botanical Garden, 1998 Woody Plants of Kentucky and Tennessee: Complete Winter Guide to Idenficaon and Use Ronald L. Jones and B. Eugene Wofford, University Press of Kentucky, 2013 Woody Plants of the Southeastern United States: A Winter Guide Ron Lance, University of Georgia Press, 2004

65 Guide to the Vascular Plants of the Blue Ridge B. Eugene Wofford, University of Georgia Press, 1989 Nave Trees of the Southeast: An Idenficaon Guide L. Katherine Kirkman, Claud L. Brown, and Donald J. Leopold, Timber Press, 2007 Forest Plants of the Southeast and Their Wildlife Uses James H. Miller, Karl V. Miller, and Ted Bodner, University of Georgia Press, 2005 Ferns of Northeastern and Central North America, 2nd Ed. Boughton Cobb, Elizabeth Farnsworth and Cheryl Lowe, Peterson Field Guides, 2005 A Natural History of North American Trees Donald Culross Peae, Houghton Mifflin Co., 2007 Edible Wild Plants, Eastern/Central North America Lee Allen Peterson, Peterson Field Guides, Houghton Mifflin, 1977 Stearn’s Diconary of Plant Names for Gardeners William Stearn, Timber Press, 1996 New Pronouncing Diconary of Plant Names American Nurseryman Publishing Company, 1964, (800) 621-5727 ISBN 1-887632-50-6 Tennessee Department of Environment and Conservaon (TDEC) Division of Natural Areas hps://www.tn.gov/environment/program-areas/na-natural-areas.html University of Tennessee Vascular Plant Herbarium (photos and distribuon) hps://herbarium.utk.edu/vascular/index.php USDA PLANTS Database, Natural Resources Conservaon Service -- hp://plants.usda.gov/java/ Vanderbilt Bioimages – hp://bioimages.vanderbilt.edu/ Plant Conservaon Alliance – hp://www.nps.gov/plants/ Tennessee Nave Plant Society -- hp://www.tnps.org

Invasive Plants

A Management Guide for Invasive Plants in Southern Forests James H. Miller, Steven T. Manning, and Stephen F. Enloe Forest Service Publicaon GTR-SRS-131 hp://www.srs.fs.usda.gov/pubs/36915 Bringing Nature Home: How Nave Plants Sustain Wildlife in Our Gardens Douglas W. Tallamy, Timber Press, 2007 Center for Invasive Species and Ecosystem Health – hp://www.invasive.org/ Photos of invasive species Tennessee Invasive Plant Council – hp://www.tnipc.org Non-nave invasive plant species list, informaon on nave plant landscaping, etc.

66 VIII. Review Quesons

1. Important physical character(s) for fern idenficaon is/are a. frond branching paern b. gill aachment to the stalk c. shape of the sorus d. both a and c

2. Dichotomous keys a. fit two different locks b. are only used by experts c. systemacally determine species idenficaon d. are only available for plants

3. A thalloid liverwort a. features a flat sheet of green ssue not differenated into leaves and stems b. never produces sporophytes c. is easy to confuse with a moss d. grows in very dry habitats

4. Which statement about fungi is true? a. Fungi manufacture their own food. b. Fungi only reproduce asexually. c. Fungi cannot break down lignin in wood. d. Fungi are heterotrophs feeding on other organisms.

5. The underside of a young mushroom cap may be protected by thin ssue called the ______which oen leaves a ring on the stem. a. mycelium b. paral veil c. universal veil d. skirt

6. The term gymnosperm a. includes all green plants b. means “naked seed” c. applies only to palm trees d. references broad-leaved trees

7. Frucose lichens a. adhere ghtly to the substrate and cannot be removed without harming both b. produce big fleshy fruits in bright colors c. are three-dimensional and oen pendulous d. look like jelly

67 8. Because lichens receive nutrion from atmospheric mineral deposion, a. some species are bioindicators of air quality b. they do not have true roots c. they do not really need algae for photosynthesis d. both a and b

9. In a flower, stamens are the male reproducve structures producing a. spores b. spules c. ovules d. pollen

10. Mosses and leafy liverworts can be disnguished by examining a. the presence vs. absence of a leaf midrib b. leaf ps that are pointed vs. rounded and lobed c. leaves spiraling the stem vs. leaves in two, overlapping ranks d. all of the above

11. Which statement about fungi is false? a. Fungi are most closely related to plants. b. Saprotrophic fungi feed on dead organic maer. c. Mycorrhizal fungi form mutually beneficial relaonships with plants. c. Sac fungi (Ascomycota) are the main fungal component in lichens.

12. Some invasive pest species a. hitchhike into the U.S. on other products and organisms b. have been deliberately introduced for specific uses c. are bought and sold as pets d. were inadvertently released or escaped e. all of the above

13. Aside from the lack of true vascular ssue, what primary difference separates nonvascular plants from all vascular plants including ferns? a. all vascular plants reproduce through seeds b. nonvascular plants grow very tall c. the gametophyte generaon is dominant and free living in nonvascular plants d. nonvascular plants only reproduce every ten years

14. Clubmosses are relaves of a. true mosses or bryophytes b. club fungi c. true ferns or pteridophytes d. frucose lichens

68 15. An obligate wetland species a. usually occurs in non-wetlands b. almost always occurs in wetlands c. is a free-floang pond plant d. depends on another species for survival

16. Fungi most commonly associated with woody species and important to the health of temperate forests a. are ectomycorrhizal b. are harmful to herbaceous plants c. produce typical “mushroom” fruing bodies d. are endomycorrhizal e. both a and c

17. Key developments in the evoluon of plant species in order are a. vascular ssue, spores, and flowers b. seeds, waxy cucle, and flowers c. vascular ssue, seeds, and flowers d. waxy cucle, flowers, and seeds

18. Which statement accurately describes one difference between seeds and spores? a. Spores contain nutrious endosperm. b. Seeds have a protecve coat. c. Seeds are mainly dispersed by wind. d. Spores of ferns, mosses, and fungi readily survive inhospitable condions.

19. Harnessing energy from the sun, plants make their ‘food’ from these raw materials a. chemical ferlizers applied to the soil b. sugar and water c. carbon dioxide and water d. leaf ssue and insect frass

20. Before DNA studies became common, plant classificaon was based primarily on a. the size of a plant b. the habitat of a plant c. the flower of a plant d. the leaf of a plant

21. In scienfic nomenclature, each organism is given a two-word binomial. The first name is the ______and the second is the ______. a. specific epithet, genus b. family, genus c. genus, specific epithet d. family, specific epithet

69 22. A lichen represents the symbioc relaonship between fungi and ______. a. bacteria b. algae c. protozoa d. tardigrade

23. Angiosperms are flowering plants whose seeds are enclosed in a. ovaries b. sperm sacs c. anthers d. sepals

24. Mosses, liverworts, and hornworts, known collecvely as bryophytes, a. lack true vascular ssue b. generally prefer moist, sheltered habitats near the ground c. only reproduce asexually d. all of the above e. both a and b

25. Which characterisc best fits a monocot? a. two seed leaves b. flower parts in mulples of three c. vascular bundles in a ring d. woody trees with a taproot

26. For reproducon, ferns require a moist to wet environment because a. high humidity smulates spore release b. flagellated male sperm must swim to female egg c. baby ferns need lots of water d. all ferns are obligate wetland species

27. A calciphile is a plant that would most likely be found growing a. in soils derived from limestone b. at elevaons above 5,500 feet c. in highly leached, upland soils d. on the upper branches of trees

28. Mycelium is the vegetave body of a a. lichen b. fern c. fungus d. liverwort

70 29. Which trait is oen typical of non-nave invasive plants? a. slow growing b. produce flowers/fruit at a young age c. sterile d. fruits unpalatable to wildlife

30. In the wild, non-nave invasive plant species a. are nutrious sources of food for nave insects b. provide valuable habitat for rare wildlife populaons c. oen outcompete and displace nave plants d. are only found in disturbed areas

31. Why might it be beneficial for organisms like fungi, lichens, and non-vascular plants to reproduce asexually? a. Genec diversity serves no purpose for lower life forms. b. It is a quick way to colonize a suitable area. c. Sexual reproducon is boring. d. Genecally disnct individuals are rare.

Answer Key 1. d 2. c 3. a 4. d 5. b 6. b 7. c 8. d 9. d 10. d 11. a 12. e 13. c 14. c 15. b 16. e 17. c 18. b 19. c 20. c 21. c 22. b 23. a 24. e 25. b. 26. b 27. a 28. c 29. b 30. c 31. b

71 Appendix A: Common Mosses, Liverworts, Hornworts Compiled by Keith Bowman for the Spring Wildflower Pilgrimage, Great Smoky Mountains

72 Appendix B: Botanical Lan Pronunciaon Guide Compiled by Margie Hunter 1. Vowel sounds are based on English long and short sounds for a, e, i, o, u. 2. Each vowel constutes its own syllable. - even when joined – acroschoides (uh-kros-h-koh-'eye-deez) frankii ('frank-ee-eye) - even at the end – canadense (kan-uh-'den-see), Silene (sih-'lee-nee) - except for diphthongs ae and oe – as long "e" laevis ('lee-vis) au – like August australis (au-'stray-lis) eu – like yew Heuchera ('hyew-keh-ruh), unless Greek - Rheum ('ree-um) 3. Consonants mostly reflect English pronunciaons. c and g are hard in front of a, o, u c and g are so in front of e, i, y, ae, oe ch is hard 'k', typically 4. Emphasis is placed on the stressed vowel syllable. 5. Stressed vowels followed by two consonants have a short sound. Canna ('kan-uh) Hemerocallis (hem-er-uh-'kal-is) reptans ('rep-tanz) unless the consonants are br, cr, dr, fr, gr, pr, tr, cl, pl – then the stressed vowel is long rubrum ('roo-brum) 6. If the next to the last syllable (penult) is stressed, it is a long vowel sound. Hamamelis (ham-uh-'mee-lis) Agerana (uh-jeh-ruh-'ty-nuh) occidentalis (oks-ih- den-'tay-lis) unless the two consonant rule comes into play – see Hemerocallis, canadense 7. If the second to the last syllable (antepenult) is stressed, it is a short vowel sound. Erigeron (eh-'ridge-er-on) Clemas ('clem-uh-s) unless the last two vowels are together, then it's long Geranium (jer-'ay-nee-um) Pulmonaria (pul-muh-'nay-ree-uh) unless it's Delphinium [Greek] (Trillium, Trollius – rule #5 applies) 8. Unstressed syllables are typically short or neutral. 9. Words of Greek origin may differ in pronunciaon from words of Lan origin. Achillea (ak-ih-'lee-uh) giganteus (jy-gan-'tee-us) 10. If based on a person's name, either pronounce as the name would be or Lanize/anglicize. Stokesia ('stokes-ee-uh or stoh-'kee-see-uh)

Pronunciaon References These references indicate which vowel is stressed and whether it is long or short: • Standard Cyclopedia of Horculture – L.H. Bailey • Gray's Manual of Botany • New York Botanical Garden Illustrated Encyclopedia of Horculture – T.H. Evere Long syllables are accented with the grave symbol ` Erythrònium (#7) Short syllables are accented with the acute symbol ´ Coreópsis (#5) ** New Pronouncing Diconary of Plant Names, American Nurseryman Pub. Co. $5.00 Source: Horculture 97 (1) 2000.

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