Club Mosses, Ferns & Horsetails: the Seed-free …...(pl.) sorus (sing.); a sorus may be protected...
Transcript of Club Mosses, Ferns & Horsetails: the Seed-free …...(pl.) sorus (sing.); a sorus may be protected...
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Club Mosses, Ferns & Horsetails: the Seed-free Vascular Plants
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Vascular Plants - a quick review Two unrelated groups within “cryptogams” – seed free vascular plants – are recognized as phyla:
1. Lycopodiophyta : lycopods 2. Polypodiophyta: ferns, horsetails, and whisk ferns
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Vascular Plants - a quick review
They produce free spores, the principal dispersal units, via meiosis. Spore: a reproductive cell, capable of developing into an adult without fusion with another cell.
spores
Why were the seed-free plants “grouped” together?
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spores
Vascular Plants - a quick review
Spores develop within a sporangium (pl. sporangia)
sporangium
Why were the seed-free plants “grouped” together?
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spores
Vascular Plants - a quick review
Spores germinate and develop into gametophytes that exist independently of the spore-producing plants. The gametophytes (haploid, n) tend to be inconspicuous and short-lived.
gametophyte
sporangium
Why were the seed-free plants “grouped” together?
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spores
Vascular Plants - a quick review
Like all plants, seed-free plants produce two kinds of gametes in their gametophytes: sperm and egg that unite to form a zygote (2n or diploid) via fertilization!
gametophyte
sporangium
zygote
Why were the seed-free plants “grouped” together?
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spores
Vascular Plants - a quick review
The sporophyte (2n) develops from the zygote and is more conspicuous, usually perennial and lives for an indefinite period!
gametophyte
sporangium
zygote
sporophyte
Why were the seed-free plants “grouped” together?
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Wisconsin Seed-free Plants The best website to identify and see images of Wisconsin’s seed-free plants is Gary Fewless’ at UW-Green Bay; links provided below
Key to Ferns and Fern Allies of Wisconsin List of Pteridophytes of Wisconsin Glossary of Fern Terminology also: Michigan Online Flora
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The best manual to identify lycopods and ferns is Ferns of Northeastern and Central North America (2nd ed.) in the Peterson Field Guides
Wisconsin Seed-free Plants
Warning: Families and genera (and thus species names) are changing quickly in the seed-free plants
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Phylum Lycopodiophyta���club mosses, spike mosses, quillworts!
Leaves microphylls: generally small, simple, one-veined leaves. Microphylls evolved by the process of enation and vascularization. An enation is a veinless, lateral protrubence on the stem. When the enation becomes vascularized it is termed a microphyll.!
Sporangia: the spore producers on the sporophytes are located singly on the upper surfaces or in axils of the bracts of a cone or of green leaves!
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Phylum Lycopodiophyta!
This group arose in the Later Silurian about 420 million years ago and was dominant in the Carboniferous and used to be much more diverse.!
Carboniferous forest from Illinois
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Phylum Lycopodiophyta!Lycopodiaceae - club mosses
Diphasiastrum complanatum
Ground cedar, crowfoot
15 genera and about 375 species Lycopodium now split into several genera cosmopolitan distribution, most diverse in tropics
Evergreen, stems elongate and dichotomously branching. Leaves often densely covering the stem.
Oily compounds in the cell walls ignite rapidly into a flash of light and were used by magicians and sorcerers in the Middle Ages. More recently they were used as a flash early in photography and in experimental photocopying machines. Many species over-collected for Christmas wreaths.
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Phylum Lycopodiophyta!Lycopodiaceae - club mosses
Homosporous: “same spore” - one kind of spore produced; cones terete (rounded)
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Phylum Lycopodiophyta!Lycopodiaceae - club mosses
other examples
Dendrolycopodium obscurum Ground pine
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Phylum Lycopodiophyta!Lycopodiaceae - club mosses
other examples
Lycopodiella inundata Bog club moss
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Phylum Lycopodiophyta!Lycopodiaceae - club mosses
other examples
Huperzia lucidula Shiny club moss
Alkaloid for Alzheimers?
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Phylum Lycopodiophyta!
Huperzia lucidula Shiny club moss
Polytrichum forest mosses
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Phylum Lycopodiophyta!Selaginellaceae - spike mosses
1 genus and about 750 species Mainly tropical family with some species extending into arctic regions of both hemispheres
Leaves spirally arranged and often 4-ranked on the secondary and ultimate branches. Spores borne in or near the axils of well-differentiated sprophylls, usually on 4 sided stroboli.
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Phylum Lycopodiophyta!Selaginellaceae - spike mosses
1 genus and about 750 species Mainly tropical family with some species extending into arctic regions of both hemispheres
Leaves spirally arranged and often 4-ranked on the secondary and ultimate branches. Spores borne in or near the axils of well-differentiated sprophylls, usually on 4 sided stroboli. Heterosporous: “different” spores !!Unlike Lycopodiaceae, Selaginellaceae are heterosporous with different types of spores: microspores and megaspore (not related to size) the micro give rise to male gametophytes and the mega give rise to the female gametophytes.
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Phylum Lycopodiophyta!Selaginellaceae - spike mosses
Heterosporous: “different” spores !!Unlike Lycopodiaceae, Selaginellaceae are heterosporous with different types of spores: microspores and megaspore (not related to size) the micro give rise to male gametophytes and the mega give rise to the female gametophytes.
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Phylum Lycopodiophyta!Selaginellaceae - spike mosses
Selaginella eclipes
meadow spike moss
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Phylum Lycopodiophyta!Selaginellaceae - spike mosses
Selaginella ruprestis
Rock spike moss Selaginella selaginoides Northern spike moss
Endangered!
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Phylum Lycopodiophyta!Isoetaceae - quillworts
1 genus and about 150 species; worldwide; 2 spp. in WI - Isoetes echinospora most common Aquatic or semi-aquatic plants with corm-like stems with secondary growth. Absorb carbon through their roots.
Leaves with ligules and quite long (15+ cm) for microphylls.
Isoetes echinospora I. butleri - not native
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Phylum Lycopodiophyta!Isoetaceae - quillworts
Heterosporous plants with sporangia borne at the base of sporophylls which are similar to vegetative leaves. Megasporangia + microsporangia
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Phylum Polypodiophyta - ferns & horsetails!Equisetaceae - horsetails, scouring rushes [previously placed in own phylum - Equisetophyta]
1 genus, Equisetum, 15 species with a cosmopolitan distribution except for Australia or New Zealand [9 species in Wisconsin]
Plants are primarily colonizers of unforested areas, lake margins and wetlands
Shoots monomorphic or dimorphic (see right). Often highly branched (horsetails) appearing like leaves. Others not branched (scouring rushes).
Internodes with conspicuous vertical ridges; jointed stems; stems hollow (both important taxonomic features for keying species)
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Equisetaceae - horsetails, scouring rushes
Leaves in whorls; leaves one veined, verticillate, united to form a sheath around the stem; these leaves are probably reduced megaphylls: Megaphylls are larger than microphylls have a blade that has a complex system of veins.
megaphyll
sheath
Megaphyll theory is supported by placement of Equisetum among primitive ferns using DNA.
branch
Phylum Polypodiophyta!
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Equisetaceae - horsetails, scouring rushes
Sporangia clustered terminally in cones composed of polygonal, umbrella-like structures with sporangia beneath.
Phylum Polypodiophyta!
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Equisetaceae - horsetails, scouring rushes
Horsetails are homosporous, have green spores with hygroscopic elators, and form conspicuous green gametophytes
Phylum Polypodiophyta!
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Equisetaceae - some examples of native horsetails & scouring rushes
Equisetum arvense Field horsetail
Equisetum sylvaticum Woodland horsetail
Phylum Polypodiophyta!
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Equisetaceae - some examples of native horsetails & scouring rushes
Equisetum laevigatum Smooth scouring rush
Equisetum scirpoides Dwarf horsetail
Phylum Polypodiophyta!
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Phylum Polypodiophyta���the true ferns!
Most diverse of the vascular cryptogams, both in species number (ca. 11,000 worldwide) and form (small aquatics to trees). Like the horsetails (which may be thought of as specialized ferns), leaves are megaphylls; blade is called a frond and the petioles as stipes.
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Phylum Polypodiophyta���ferns!
Circinate vernation: vernation is the arrangement of folded leaves in a bud, forming a crozier or fiddlehead, i.e. coiled or rolled up at the tip and unfolding lengthwise when emerging - due to auxin and differential growth of tissue. Protects young bud.
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Phylum Polypodiophyta���ferns!
Sporangia borne on the margin or the lower surface of the leaf; often grouped in sori (pl.) sorus (sing.); a sorus may be protected by a flap-like structure called the indusium - an important feature for identification of ferns due to its remarkable variation in size, shape, and position.
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Phylum Polypodiophyta���ferns!
The sprorangium is often associated with an annulus a cluster or row of cells with thick walls that open the sporangium and catapult the spores into the air.
annulus
Ferns can be either homosporous or heterosporous.
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Phylum Polypodiophyta���ferns!
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Phylum Polypodiophyta!There is major disagreement on what are the fern families, but 30 are generally recognized worldwide. In Wisconsin there are 9 families. Characters used to circumscribe families often relate to cryptic features such as the presence or absence of annulus and its orientation, the length and diameter (number of cells) of the sporangial stalk, spore shape, stem and petiole cross-section anatomy. The primitive ferns include horsetails and whiskferns.
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Phylum Polypodiophyta!Ophioglossaceae - adder’s tongue family
Primitive ferns; 2 genera in Wisconsin: Ophioglossum and Botrychium (now 3 genera)
Sporangia are on an erect axis with a green blade attached below the sporangia bearing part
Ophioglossum pusillum Adder’s tongue fern
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Phylum Polypodiophyta!Ophioglossaceae - adder’s tongue family
Botrypus virginianum Rattlesnake fern
Botrychium lunaria Moonwort fern
Botrychium mormo Goblin fern
Primitive ferns; 4 genera in Wisconsin: Ophioglossum and Botrychium (now 3 genera)
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Phylum Polypodiophyta!Osmundaceae - royal fern family
Primitive ferns; 1 genus in Wisconsin: Osmunda
Fertile and sterile leaves dimorphic and the fertile portion is non-green at maturity. Sporangia not clustered in sori, but naked.
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Phylum Polypodiophyta!Osmundaceae - royal fern family
3 species of Osmunda in Wisconsin; easily separated by position of fertile portions
Osmunda cinnamomea Cinnamon fern
Osmunda claytoniana Interrupted fern
Osmunda regalis Royal fern
All wide-spread & Bot 401 exam plant!
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Phylum Polypodiophyta!Dennstaedtiaceae - bracken family
2 genera in Wisconsin. Includes one of the most widespread of all vascular plants.
Pteridium aquilinum Bracken fern
Clonal with rhizome; large compound leaves. Ubiquitous in Wisconsin.
Marginal sori with no indusia but with revolute (rolled over) leaf edge protecting sori.
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Phylum Polypodiophyta!Pteridaceae - maidenhair fern family
4 genera in Wisconsin
Adiatum pedatum Maidenhair fern
Distinctively compound frond with dark purple stipe and rachis
Sori that lack indusia or are protected by a reflexed or revolute margins
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Phylum Polypodiophyta!Aspleniaceae - spleenwort family
1 genus in Wisconsin. Defined by linear or kidney shaped sori.
Asplenium viride (A. trichomanes-ramosum)
Green spleenwort Asplenium platyneuron
Ebony spleenwort
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Phylum Polypodiophyta!
2 genera in Wisconsin (also Matteuccia – ostrich fern). Sensitive ferns has dimorphic fronds. Sterile frond pinnately lobed. Fertile frond turning black.
Onoclea sensibilis Sensitive fern
Onocleaceae- sensitive fern family
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Phylum Polypodiophyta!Dryopteridaceae - woodfern family
Large and diverse group of ferns; often broadly defined to include other smaller families. 2 genera and 13 species in Wisconsin.
Polystichum acrostichoides Christmas fern
Dryopteris intermedia Shield fern
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Phylum Polypodiophyta!Polypodiaceae - rockcap family
40 genera and over 500 species in tropics and subtropics, but a single species in Wisconsin.
Polypodium virginianum Rockcap fern
Distinctive fronds: simple, pinnately lobed; leathery. Colonizes bare rock.
Sori in two rows on each lobe.
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Phylum Polypodiophyta!Salviniaceae (Azollaceae) - mosquito fern family
2 species in Wisconsin. Floating aquatic, reduced ferns with 2 ranked leaves, each 2 lobed. Symbiotic relationship with N2 fixing blue green bacteria Anabaena azollae. Heterosporous!
Azolla mexicana & A. caroliniana Floating fern
Western & SE coastal elements