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BIOLOGICAL DIVERSITY:

Table of Contents

Gymnosperms are Seed Plants | | | | Gnetales | Angiosperms are Flowering Plants | | Trends in | Learning Objectives | Terms | Review Questions | Links

Gymnosperms are Seed Plants | Back to Top

Gymnosperms have but not or flowers. Gymnos means naked, means seed: thus the term = naked seeds. Gymnosperms developed during the Paleozoic Era and became the dominant seed plant group during the early Era, as shown in Figure 1. The ancestors of gymnosperms were some now-extinct type of heterosporous or related group. There are 700 living of gymnosperms placed into four divisions: conifers (such as and spruce), cycads (such as the palm, revoluta), ginkgos (the maidenhair , biloba), and gnetophytes (such as Mormon tea, ).

Figure 1. The records of some and plant groups. The width of the shaded space is an indicator of the number of species. Image from Purves et al., : The Science of , 4th Edition, by Sinauer Associates (www.sinauer.com) and WH Freeman (www.whfreeman.com), used with permission.

Gymnosperms are undoubtedly the group from which the angiosperms developed, although, as noted in Origin of Species, which group "remains an abominable mystery". Numerous gymnosperm groups have been proposed as ancestors over the past century.

Cycads | Back to Top

Cycads are placed in the Cycadophyta. They retain several fern-like features, notably pinnate and circinate vernation. However, they usually produce cones of nonphotosynthetic reproductive structures, a distinctively unfernlike feature. Cycads, like all seed plants, are also heterosporous, unlike the which are all homosporous. cones are unisexual, in fact the plants producing them are dioecious, having separate male and plants. Cycads also produce free-swimming sperm (a feature found only in ginkgoes among living seed plant groups).

Cycads were much more prominent in the of the Mesozoic than they are today. Presently, they are restricted to the . floridana is the only cycad occurring natively in the continental . Several species of Cycas, notably C. revoluta (shown in Figure 2), are commonly encountered cultivated plants in warm, moist areas. leaves are often used in Palm Sunday services in some churches, both for their feathery appearance and ease of obtaining from local greenhouses.

Figure 2. Cycas revoluta from Hawaii, Top: female cones bearing large, seeds. Image from http://www.botany.hawaii.edu/faculty/carr/images/cyc_rev_f.jpg; Bottom: male cone bearing numerous -producing sporangia. Image from http://www.botany.hawaii.edu/faculty/carr/images/cyc_rev_m.jpg.

Ginkgos | Back to Top

The ginkgoes also were a much more prominent group in the past than they are today. The sole survivor of this once robust and diverse group is , the maidenhair tree shown in Figure 3. Extensively used as an ornamental plant, Ginkgo was thought extinct in the wild until it was discovered growing natively in a remote area of . Ginkos are dioecious, with separate male and female plants. The males are more commonly planted since the produce seeds that have a nasty odor. is by wind. Recently, Ginkgo has become the current herbal rave, although scientific studies have debunked the claim that the herbal supplement made from ginkgoes improves . I think.

Figure 3. Ginkgo biloba, close-up of young . Image from http://www.dinoworld.net/ginkgo.JPG. Precise systematic placement of the ginkgoes has yet to bet determined. Ginkgoes have motile (swimming) sperm, a rarity among living seed plants (only ginkgoes and cycads have this feature today), although the vegetative anatomy of ginkgoes is more -like (long shoot and short shoot discussed below; structure of their ). Ginkgoes, like the cycads, are dioecious, and also have similar seed features to cycads.

Plants possibly allied to the modern ginkgoes have been found in -aged and later rocks. These plants have been classified in the -genera (shown in Figure 4) and , although recent studies suggest these genera are really morphological variants and that the modern Ginkgo should be used to include these . During the Mesozoic ginkgoes were worldwide in their distribution and important elements in the gymnosperm forests that dominated the land.

Figure 4. Ginkgoites leaf from the of North Korea. Although this fossil is not the typical fan shape of most modern ginkgo leaves, modern ginkgoes do produce leaves of this general shape. Image from http://www.dges.tohoku.ac.jp/museum/fosgal19.html.

Conifers | Back to Top

The conifers remain a major group of gymnosperms that include the pines, spruce, , bald cypress and Norfolk Island (). The division Pinophyta contains approximately 550 species of conifers. The conifers are cone producing and that usually have needle-like leaves. Needles have a thick cuticle, sunken stomates, and a reduced surface area. The conifers, as a group, are well adapted to withstand extremes in and occur in nearly all habitats from the equator to the subpolar regions. The biome consists largely of various conifer species.

Auracarias

Members of this group of conifers have numerous small, scale-like leaves spiraling around their stems. Araucaria, a major genus that gives its name to the group, is a common ornamental because of the symmetry and beauty of its growth form. The monkey puzzle tree, shown in Figure 5, is a species of Araucaria.

Figure 5. Image of Araucaria sp. Note the large female comes at the tips of branches. Image from http://www.botany.hawaii.edu/faculty/carr/images/araucar_sp3.jpg.

The fossil record of Auracarias and similar plants is quite good. The fossil genus Auracarioxylon that grew in Arizona during the Triassic Period comprises the largest group of petrified wood in the Petrified National Park of Arizona.

Taxodiaceae: Sequoias and more

Members of this group include some of the largest trees, and have been significant members of the forests of the world since the Mesozoic. Sequoia, shown in Figure 6, and Sequoiadendron are major genera in this group.

Figure 6. Top: photograph of Sequoia sempervirens. Note the tall, woody stem with leaves borne at the top of the stem. Image from http://www.botany.hawaii.edu/faculty/carr/images/seq_sem_hab.jpg. Bottom: Close-up of Sequoia sempervirens leaf and cone. Image from http://www.botany.hawaii.edu/faculty/carr/images/seq_sem_cu.jpg.

The Pine Life Cycle

Pines have an interesting life cycle, shown in Figure 7, that takes two to complete. Not all seed plants have such a long time span to complete their life history: some flowering plants manage to do it in as little as a few weeks.

The , as in all other groups, is the dominant, photosynthetic part of the life cycle: when you are holding pine needles in your hand you are holding sporophyte parts. Pines have specialized reproductive structures in which occurs: pine cones. Pollen grains are produced in the male cones, and contain the male (which consists of only a very few cells). Pollen released from the male cones is carried by wind to the female cones, where it lands. The cones close and the next the pollen grain germinates to produce a that grows into the female gametophyte. The sperm (from the pollen grain) and fuse, forming the next generation sporophyte. The sporophyte develops into an encased within a seed. The seed is later released to be transported by the wind to where (hopefully) it lands and germinates. If you have seen a large pine tree you realize there are hundreds or more female cones on such a tree. Pine pollen has been noted to travel great distances from the plant that produced it, if the wind is strong enough. To aid this transport pine pollen has two air sacs, and thus is quite distinctive, as shown in Figure 8.

Figure 7. The pine life cycle. Note: to view these four segments in correct sequence you will need to increase your browser window width as much as possible. Images from Purves et al., Life: The Science of Biology, 4th Edition, by Sinauer Associates (www.sinauer.com) and WH Freeman (www.whfreeman.com), used with permission.

Figure 8. Pine pollen from 38,000 year old sediments in the White Mountains of eastern Arizona. Image from http://www.smu.edu/geology/palynolab.htm.

Gnetales | Back to Top

The Gnetales, shown in Figure 9, are an odd group: they have some angiosperm-like features but are not themselves angiosperms. Cladistic analyses support placement of the gnetales (or some portion of them) as outgroups for the flowering plants. Three distinctive genera comprise this group: , , and Ephedra. Ephedra occurs in the western United States where it has the common name "Mormon tea". It is a natural source for the chemical ephedrine, although there is no evidence the Mormons in Utah (where the plant is extremely common) ever used it for tea. Welwitschia is limited to coastal in South , although fossil leaf, cuticle and pollen evidence indicates plants of this type were widespread during the Mesozoic Era. Welwitschia is noted for its two long, prominent leaves. Gnetum has leaves that look remarkably like those in angiosperms, as well as vessels in the , generally considered an angiosperm characteristic.

Figure 9. Representative gnetalean plants. Top: Ephedra female plant. Image from http://www.botany.hawaii.edu/faculty/carr/images/eph_sp_f.jpg. Middle: Welwitschia habit shot of a cultivated specimen. Image from http://www.botany.hawaii.edu/faculty/carr/images/wel_mir.jpg. Bottom: Gnetum gnemon habit shot showing pinnate leaves bearing female strobili. Image from http://www.botany.hawaii.edu/faculty/carr/images/gne_gne.jpg. Among the gnetalean plants, Ephedra is perhaps the best known. One folkloric name for the plant is "Mormon tea". This is a misnomer as there appears little or no evidence that members of a religion that bans such as ever brewed a tea from the plant. However, the plant does produce the drug ephedrine, a lately linked to deaths of athletes.

Welwistchia is a very bizarre plant natively growing only in the coastal deserts of South Africa. The plant produces two long leaves and a crown of reproductive cones rimming a brown, central body. Pollen resembling Welwitschia has been found in many parts of the world, indicating a formerly more widespread distribution of this enigmatic plant.

Angiosperms are Flowering Plants | Back to Top

Flowering plants, the angiosperms, were the last of the seed plant groups to evolve, appearing during the later part of the of the Age of (the beginning of the , 140 million years ago). All flowering plants produce flowers. Within the female parts of the angiosperms produce a diploid and triploid . Fertilization is accomplished by a variety of , including wind, , and water. Two sperm are released into the female gametophyte: one fuses with the egg to produce the zygote, the other helps form the nutritive known as endosperm.

The angiosperms (angios = hidden) produce modified leaves grouped into flowers that in turn develop fruits and seeds. There are presently 235,000 known living species. Most angiosperms also have larger xylem cells known as vessels that improve the efficiency of their vascular systems.

Whence came the angiosperms? This was Darwin's "abominable mystery". Clearly angiosperms are descended from some group of Mesozoic-aged gymnosperm seed plant....but which one? Click here to view an online lab exercise in phylogeny and try to figure things out!

The classical view of flowering suggests early angiosperms were evergreen trees that produced large -like flowers. Click here to view an illustration of suggested paths of floral evolution. However, this view has recently been contradicted by the oldest fossil yet found, a 140 million year old plant found by David Dilcher and his associates.

The angiosperms underwent an during the Cretaceous, and for the most part escaped the major at the end of the Cretaceous.

Flowers | Back to Top

Flowers are collections of reproductive and sterile tissue arranged in a tight whorled array having very short internodes. Sterile parts of flowers are the and . When these are similar in size and shape, they are termed . Reproductive parts of the flower are the (male, collectively termed the androecium) and carpel (often the carpel is referred to as the pistil, the female parts collectively termed the ). Lily flowers (shown in Figure 10) demonstrates these concepts. Figure 10. Flower of lily () illustrating a complete flower. The image is reduced from gopher://wiscinfo.wisc.edu:2070/I9/.image/.bot/.130/Angiosperm/Lilium/Flower_dissection/Flower. Follow that link to view a larger image of this flower.

Flowers may be complete, where all parts of the flower are present and functional, or incomplete, where one or more parts of the flower are absent. Many angiosperms produce a single flower on the tip of a shoot (like the lily pictured in Figure 10, or ). Other plants produce a stalk bearing numerous flowers, termed an , such as is seen in many orchids. Many flowers show for pollination, bearing numerous white or yellow petals. Others, like the grasses, , and , are wind pollinated and have their petals reduced and often inconspicuous.

Angiosperm Life Cycle

Flowering plants also exhibit the typical plant alternation of generations, shown in Figure 11. The dominant phase is the sporophyte, with the gametophyte being much reduced in size and wholly dependant on the sporophyte for nutrition. The is not a unique angiosperm condition, but occurs in all seed plants as well. What makes the angiosperms unique is their flowers and the "" that occurs. Technically this is not double fertilization, but rather a single egg-sperm fusion (fertilization proper) plus a fusion of the second of two sperm cells with two haploid cells in the female gametophyte to p[produce triploid (3n) endosperm, a nutritive tissue for the developing embryo. More details on this aspect of the flowering plants are available in the FLOWERING PLANT : Fertilization and Fruits chapter.

Figure 11. Life cycle of corn, a typical monocot angiosperm. Note the formation of endosperm by "double fertilization". Images from Purves et al., Life: The Science of Biology, 4th Edition, by Sinauer Associates (www.sinauer.com) and WH Freeman (www.whfreeman.com), used with permission.

Angiosperm Systematics | Back to Top

The flowering plants, the division Magnoliophyta, contain more than 235,000 species, six times the number of species of all other plants combined. The flowering plants divide into two large groups, informally named the monocots and the dicots. The techjnical names for these groups are the for dicots and the class for monocots.

The are in the class Magnoliopsida and have these features: either woody or herbaceous, flower parts usually in fours and fives, leaves usually net-veined, vascular bundles arranged in a circle within the stem, and produce two (seed leaves) at . Prominent dicot families include the mustards, , cacti, and . Several dicot families are noteworthy because of the illegal drugs (shown in Figure 12) derived from them: the Cannabinaceae (marijuana) and ( from which opium and heroin are derived). (in the dicot ) is the plant from which the illegal drug is extracted.

Figure 12. Some dicot plants bused for making illegal drugs. Top: , left image, the plant that is the source of cocaine, from http://www.drugs.indiana.edu/graphics/photographs/cocaine1.gif, right image, cocaine, from http://www.drugs.indiana.edu/graphics/photographs/cocaine3.gif; Bottom left, sativa, the marijuana plant; image from http://www.drugs.indiana.edu/graphics/photographs/marij6.gif, right: somniferum, the opium being harvested; image from http://www.drugs.indiana.edu/graphics/photographs/opium7.gif.

Not all dicot plants are misused to produce illegal drugs. Notable dicot families with legitimate uses include the family, which includes the crop plants , , and peas as well as many ornamental landscape plants such as . Beans are an excellent source of nonanimal protein as well as fiber. Another dicot of enormous use is , , shown in Figure 13. Chocolate and are products of the plant family . is produced from arabica, a plant in the family , while tea comes from Camelia sinensis (), a plant native to China.

Figure 13. Economic dicots. Top: Gossypium sp., cotton., image from http://botit.botany.wisc.edu/images/401/Magnoliophyta/Magnoliopsida/Dilleniidae//Gossypium/Cotton_boll_leaves_MC.html; Bottom: annuum, the ; Image from http://www.botany.hawaii.edu/faculty/carr/images/cap_ann_233.jpg.

The class Liliopsida has plants that are herbaceous (a majority are, only palms and stand out as monocot trees), flower parts are in threes, leaves are usually parallel-veined, vascular bundles are scattered within the stem, and produce one (seed leaf) at germination. Monocot families include lilies, palms, orchids, irises, and grasses.

The monocot family (known previously as the Gramineae) includes the grasses such as corn, , , , and that are staple food products as well as ornamental plants such as crabgrass and tiff grass. The importance of this plant family to modern civilization cannot be overstated, as the first six plants mentioned in the previous sentence provide 75% of our food, either directly as food we eat or indirectly as food for animals we eat. Trends in Plant Evolution | Back to Top

Several evolutionary trends within the plant have been noted. The monophyletic nateure of this kingdom is not in dispute, with the first major division being between vascular and nonvascular plants. Wihin the vascular plants we see increasing changes in the relationship between sporophyte and gametophyte, culminating in flowering plants.

Developing from green algal ancestors, plants show a trend for reduction of the complexity, size, and dominance of the gametophyte generation. In nonvascular plants the gametophyte is the conspicuous, photosynthetic, free-living phase of the life cycle. Conversely, the angiosperm gametophyte is reduced to between three and eight cells (hence it is very inconspicuous) and is dependent on the free-living, photosynthetic sporophyte for its nutrition.

Plants also developed and refined the -shoot-leaf axis with its specialized conducting cells of the xylem and . The earliest vascular plants, such as Cooksonia and Rhynia, were little more than naked (unleafed) photosynthetic stems. Some plants later developed that produced wood. Numerous leaf modifications are known, including "carnivorous" plants such as the , as well as plants that have reduced or lost leaves, such as Psilotum and the cacti.

A third trend is the development of the seed to promote the dormancy of the embryo. The seed allows the plant to wait out harsh environmental conditions. With the development of the seed during the Paleozoic era plants became less prone to mass extinctions.

The fourth trend in plant evolution is the encasing of a seed within a . The only plant group that produces true fruit is the flowering plants, the angiosperms. Fruits serve to protect the seed, as well as aid in . Table 1. Plant classification.

Division Characteristics Examples and Uses nonvascular plants, gametophyte Bryophyta , liverworts, dominanted life cycle naked photosynthetic stems, no Psilophyta Psilotum nudum, Tmesipteris;no commercial uses leaves, free-sporing microphyll leaves, sporangia in Lycopodium, a homosporous plant, , a Lycophyta (the strobili; more significant in the heterosporous plant; no commercial uses except as club mosses) Paleozoic ground cover in some areas Sphenophyta megaphyll leaves, sporangia in (= tight cones; plants with jointed , the scouring rush or horsetail; no Equisetophyta; stems; more significant in the commercial applications the horsetails) Paleozoic megaphyll pinnately compound leaves, sporangia on underside of Ornamental plants such as the Boston fern; tree ferns; Filicophyta leaves, clumped in sori; leaves commercial applications as ornamentals; ecologically (ferns) arise by circinnate vernation important plants in some areas of the world (uncoiling to look initially like a crozier or sheperd's crook) long, pinnately compound megaphyll leaves with a leathery feel arising from soft woode4d Cycas revoluta, a common ornamental known as the Cycadophyta stems by circinnate vernation; new Sago palm;Zamia floridana, only cycad native to the (the cycads) leaves arise as a crown or ; 48 contiguous United States; commercial uses as reproduction by seeds produced in ornamental plants female cones, pollen produced in male cones fan shaped, usually bilobed leaves on a woody tree with long anmd Ginkgo biloba, the maidenhair tree; commercial uses Ginkgophyta short shoot anatomy; seeds as ornamental plants and allegedly as a memory aid in (the ginkoes) produce a foul odor when mature; herbal form pollen elliptical, monoaperturate needle-like or scale-like leaves with thick cuticles and sunken stomata; soft wooded plants; seeds Pinus(wood, resin, pine straw, pine nuts, ); Coniferophyta lacking fruit, dispersede from (yew) a natural source for the anticancer drug taxol; hardened cones; pollen can be Douglas fir (wood); ornamental plants bisaccate (Pinus) or lacking sacs (Taxodium) living plant group most closely related to flowering plants; vessels in some members of group; Ephedra (Mormon tea) source of ephedrine; Gnetum; reproductive structures close to Welwitschia have no commercial uses flowers; pollen elliptical eith usually one aperture flowering plants (angiosperms) Monocots: cereal grains form staple of economy and that encase their seeds within a diet; palms are used in many places as building ripened wall known as a material; ecologicalimpact as pioneer species in fruit; fruits may be dry or fleshy, disturbed habitats; single or multiple; two major Magnoliophyta classes the monocots and dicots; Dicots: food crops such as beans, peas; fibers from xylem may include vessels as well cotton used to make clothing and paper; illegal drugs as ; range from such as marijuana, cocaine, heroin; legal products such herbaceous annuals to periennel as chocolate, tea, and coffee; leaves make trees tobacco a dangerous legal "drug"; ornamental plants

Learning Objectives | Back to Top

Be able to compare and contrast the characteristics of seed, nonseed, and nonvascular plants. To do this complete the table below. Characteristic Nonvascular Plants Nonseed Plants Seed Plants Dominant Phase of Plant Life Cycle Swimming Sperm Pollen Tube Pollen or Homospory or or Mixed Examples of Plants in this Group Be able to list the uses of some common gymnosperm plants, such as pines and ginkgoes. Be able to list some of the uses of flowering plants. It has been stated that the ancestor of flowering plants most likely was a gnetalean plant (or at least a plant closely related to them). What evidence supports this? How might the fact that a species in monoecious or dioecious affect your decision to use it as a crop plant? As a landscape plant? Terms | Back to Top

androecium angiosperms Auracarioxylon carpel cones conifers cycads dioecious endosperm Ephedra fertilization ginkgoes Gnetales Gnetum gymnosperms gynoecium homosporous internodes Mesozoic Era Paleozoic Era petals pistil pollen grains pollen tube sepals sporophyte stamen Welwitschia zygote

Review Questions | Back to Top

1. Endosperm tissue is unique to which of these groups? a) cycads; b) conifers; c) flowering plants; d) ferns 2. Seed plants have ___. a) pollen grains with male developing on the inside; b) female gametophytes developing inside an retained on the sporophyte; c) ; d) all of these 3. Which of these plants is not a gymnopserm? a) corn; b) pine; c) redwood; d) ginkgo 4. Which of these plants is not a flowering plant? a) corn; b) wheat; c) lily; d) douglass fir 5. Cycads and ginkgoes were important plants to the world's terrestrial ecosystems during the ____. a) Paleozoic era; b) era; c) Mesozoic era; d) 6. The presence of long shoot/short shoot anaytomy characterizes which of these pants? a) Ginkgo; b) Cycas; d) Zamia; d) ferns

Answers: 1. c; 2. d; 3. a; 4. d; 5. c; 6. a; Links | Back to Top

Non-Flowering Plant Family Access Page Sorted by family on the non-flowering plants. Thumbnail photos are linked to larger versions. This site is a great educational resource maintained by Gerald D. Carr. Encyclopedia of Plants Scientific and common names for garden plants. The Botanical Society of America The official website of the plant folks. Cycad Pages This Australian website is a great stepping stone for more information about a favorite plant group of mine: cycads. This page is maintained by Ken Hill of the Royal Botanic Gardens Sydney and Dennis Stephenson of the New York . Ginkgo biloba History, uses, and other information about this living "fossil tree" whose leaves are essentially unchanged since the Tertiary period. The Ginkgo Pages Cor Kwant has produced a great resource on a fascinating plant. A great resource for images and information! Gymnosperm Database Christopher J. Earle has developed a nifty resource for digging deeper into the gymnosperms. A series of links from this page take you into images and descriptions of the various gymnosperm groups. Erythroxylum: The Coca Plant This ethnobotanical page, by April Rottman, describes the systematics and uses of the plant as well as its product, cocaine. : The Versatile Crop This page, part of a series of ethnobotanical leaflets, is by Bryan Young, and describes the natural history and economic uses of barley, the 4th most important crop.

All text contents ©1995, 1999, 2000, 2001, 2003, 2004, by M.J. Farabee. Use of text for educational purposes is encouraged.

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