III. Structural classification. Plants enjoy a structural classification system, in addition to the genetic taxonomies above. It is called the life form classification. A. It has to do with the appearance or structure of an individual plant (and nothing to do with its ancestry per se). The reason for using a structural classification is that many biogeographers are interested in how plants affect the appearance of a particular landscape, giving a certain character to an area or region. Mastering the taxonomy of all the species in the area (not to mention dealing with the controversies in taxonomy) is not going to give you that sense as well as knowing the vegetation structurally. B. I'll just run through the categories in a list here: 1. Vascular plants are plants that have an internal system of tubular cells or vessel cells for conducting water and dissolved minerals around within the plants. This internal system has two basic parts: Xylem (tubular cells or vessel cells, which are themselves dead) and phloem (or sieve cells for moving food around, which are fully alive). Because of this internal division of labor, the vascular model allows for the growth of really large plants (e.g., those giant sequoias in the Sierra). Non-vascular plants are limited to small size. So, vascular plants tend to dominate most landscapes visually. a. Tree (awesome scientific word, hard to pronounce <G>): i. Trees are woody vascular plants. ii. They generally have a single main trunk, with branching away from this main trunk. iii. Examples: a. Pines b. Oaks c. Sycamores (which often branch off pretty low to the ground) d. Palms (the tree definition maxxed out!) b. Shrub: i. Shrubs are also woody vascular plants. ii. Unlike trees, however, they have several major stems branching out from the ground or very close to it. iii. Most shrubs are smaller than most trees. iv. Examples: a. Rose b. Manzanita (common on the hills of California) c. Gardenia d. Hibiscus e. Chamise (another common California hillside native) c. Liana: i. Lianas are also woody vascular plants. ii. They have a climbing habit, seeking support from trees, rocks, hillsides, fences, what have you. iii. Examples: a. Poison Oak (leaflets three, let it be!)
b. Star Jasmine (often used as an aromatic ground cover in landscaping) c. Lilac d. Herb: i. Herbs are tender vascular plants, with little to no wood development. ii. Without the support of a woody internal framework, they are small in size. iii. There are some sub-types: a. Forbs are broadleafed forbs, such as dandelions, pothos, philodendron (and a lot of common indoor house plants) b. Graminoids, a fancy way of describing grasses, which are also herbs: linear-bladed, the lower part of their leaves wrapping around the stem. Examples include wheat, barley, corn, rye, oats, fescues, and other common lawn grasses. c. Ferns or pteridophytes are herbs. Their leaves share a lot of characteristics with branches and are called fronds. They reproduce with spores, not seeds, and do not flower. Spores have only half the genetic material that a seed would (kind of like sperm and eggs in humans). What spore reproduction means is that the fern plant we're all familiar with generates a fine dust of spores. In the right conditions, the spore develops into a tiny plant-like entity (0.6 cm) called a prothallium that generates eggs and sperm. This prothallium is the sexually-reproducing phase of the fern life cycle. The eggs and sperm find one another in water and their union produces the fern plant we know, with a complete set of genes, which gets their spores out and about. d. Club mosses or microphyllophytes (micro-dinky leaf plants, rendered loosely) are very small plants a few centimeters high, though they have horizontal stems that may sprawl out something like 15 meters. They look almost like miniature pine trees, complete with scaly leaves, stems, and "cones" (filled with spores not seeds). e. Horsetails or equisetophytes have jointed stems that makes them look a little like miniature bamboo. There are small scaly leaves that grow in whorls around the stem at the joints. Like the club mosses, the horsetails have spore-producing cones at the tops of their stems. 2. Byrophytes are non-vascular and, thus, are unable to pull water and nutrients up from the ground for any significant distance. It is for this reason that they are considered to be rather primitive plants, transitional from aquatic plants (such as kelp) and land plants (such as trees). Given their lack of a vascular system, they are really dependent on a very moist environment. Like the ferns, club mosses, and horsetails (which are vascular), they reproduce by spores, alternating generations the way I described for ferns. a. Given their lack of a vascular system, bryophytes have to be pretty small, a few centimeters, maybe up to a meter at most, and so they live close to the ground or on tree trunks. b. Like the vascular plants, however, they do have the capacity to photosynthesize (they are autotrophs). c. They can be subdivided: i. True mosses (including peat moss or sphagnum, much loved by gardeners for keeping moisture in the soil), with structures that resemble leaves, stems, and roots but no vascular tissue to conduct materials around within the plant. So, like all bryophytes, they are pretty small, ranging from microsopic in size to maybe one meter tall. Interesting geotrivia: Mosses are very old land plants (going back about 280,000,000 years) and were the first green plants to stand erect on the land. ii. Liverworts or hepatophyta are weird plants that sort of look like leaves or chains of leaves on the ground. They have no roots or stems, just these leaf-like structures. Photosynthesis goes on at the top surface of the plant, while storage of food goes on directly below. iii. Hornworts or anthocerotophytes resemble liverworts but they have irregular and misshapen greasy-looking blue-green "leaves." From this structure rises a long, slender stalk within which the spores are developing: This structure is the "horn" in "hornwort. It can get up to half a meter in height. 3. Epiphytes may be either vascular or non-vascular, so I couldn't put them in the tidy classes above. What differentiates them is that they live on top of other plants without soil contact for at least part of their lives. Very commonly, you'll see them on tree branches, especially in tropical rainforests, where small plants would not get enough sunshine on the forest floor to photosynthesize. So, small plants take up life on top of other plants to get access to the sunshine they need. a. Most epiphytes seek only mechanical support from other plants to have access to sunshine. They may extract water from the pools of water that may form in their own leaves or perch sites or from moist litter that accumulates there. Other epiphytes are parasitical, sinking their roots into the vascular tissue of their hosts to extract water and nutrients brought up by their hosts. b. Epiphytes can be subdivided by just how long they live up on top of another plant: i. Full epiphytes spend their entire lives as arboreal species, never once coming in contact with the ground. ii. Hemi-epiphytes spend only part of their lives without soil contact in a completely arboreal stage. There are two variants on this: a. A primary hemi-epiphyte starts life as an epiphyte until it can get aerial roots down from the treetops to the ground where they become rooted. The resulting rooted plant still leans on the plants they started out on and now sort of resemble lianas. Note the Tarzanesque appearance of this forest because of these aerial roots! b. A secondary hemi-epiphyte starts life on the ground and then climbs up on a nearby tree like a liana but, unlike a liana, once part of it reaches a suitable perch, it abandons its soil-touching roots and spends the rest of its life as an epiphyte. c. Some examples: i. Bromeliads have many epiphytic species. ii. Orchids also have many epiphytic species. iii. Mistletoe is a parasitical epiphyte that grows on oaks and other trees. iv. Strangler fig is a primary hemi-epiphyte that starts out as an epiphyte on a tropical rainforest tree and then sends down aerial roots in such profusion that they strangle the host tree. The tree dies and slowly rots out, leaving behind the huge, hollow cylinder formed by all those tangled fig roots, and the strangler fig is left for all intents and purposes a tree of sorts. v. Dodder weed in California is an obligate parasite on several chaparral brush species. Though related to morning glory, it is incapable of photosynthesis, so it looks like this bright orange netting! It has to sink modified roots into the stems of its host to extract water and food from it. 4. Fungi are no longer, strictly speaking, classified as plants! They are incapable of photosynthesis and do not descend from a photosynthetic ancestor. In fact, genetic evidence suggests that they are, in fact, more closely related to animals than to plants, probably sharing a common protist ancestor with animals! Subjectively, we tend to think of them as plants because they are immobile and because they have structures that vaguely resemble stems and roots, sort of, and they do reproduce with spores. a. Fungi, incapable of making their own food like plants and unable to move and hunt for food like animals, make their livings as saprophytes (they are very important detritus feeders), parasites (think: athlete's foot, jock itch, yeast infections, and ringworm, all commonly caused by fungi), and symbionts (more on that in a minute). b. Examples: i. Mushrooms and toadstools (toadstools are poisonous mushrooms), morels and truffles ii. Yeasts, including those used to make bread rise iii. Powdery mildew iv. Molds, including bread mold v. An assortment of blights, rusts, and diseases like the ones I mentioned above. 5. Algae (except blue-green algae, which are also called cyanobacteria) are very simple eukaryotic organisms that can perform photosynthesis with chlorophyll. Because of this, they are often lumped in with plants and gave rise to plants. They are a very ancient lineage, going back about three billion years. a. Most are one-celled organisms, such as that slime that forms in toilets if you're not into the Suzy Homemaker routine (science comes alive in the privacy of your dwelling unit -- isn't Nature beautiful!?). b. Sometimes they group together in colonies, such as algal mats. c. Some versions can group together to form organisms with simple tissues, such as kelp (seaweed, which can get up to 65 m in length!). Here's an image of an underwater "forest" off the coast of Alaska, made up of kelp -- isn't it gorgeous? Looks just like a forest on land, doesn't it? 7. Blue-green algae (aka cyanobacteria) and bacteria are prokaryotic one-celled creatures, meaning their DNA is not housed in a nucleus. Blue-green "algae" is now seen as a misnomer but, interestingly enough, cyanobacteria are relevant to regular algae and plants: The chloroplasts inside plants and ordinary algae are basically endosymbiotic cyanobacteria. This means that the eukaryotic cells that make up algae and plants have internal structures and organelles, one of which is the chloroplast. The chloroplast so strongly resembles existing cyanobacteria that we think an ancient symbiosis was struck up between a cyanobacterium and some eukaryotic cell, wherein the cyanobacterium photosynthesized food for both cells in exchange for a safe shelter from the harsh outside world, complete with a flow of nutrients needed for photosynthesis, and this union became the template for regular eukaryotic algae and plants. 8. Lichens are that crusty or fuzzy stuff you find growing on rocks and tree trunks. a. They are tiny compound creatures, representing the union of a fungus with an alga and/or a cyanobacterium (sometimes all three). b. This is about the perfect symbiosis: A total win-win situation for everyone involved: i. The alga and/or cyanobacterium provides food for everyone through photosynthesis, allowing the fungus to live in harsh, sterile environments (e.g., rocks) that it could not colonize before for lack of food (most fungi are parasites or detritivores and rocks can't play hosts and there's little dead organic matter to be had on them). ii. The fungus provides protection from drying out and is able to draw water and minerals into the compound, allowing the alga or cyanobacterium to live in dry habitats normally off limits. iii. Lichenologist Trevor Goward put it this way: "Lichens are fungi that have discovered agriculture," which is a rather cool analogy. c. This symbiosis is so close that it models the far more ancient symbioses between mitochondria and chloroplasts with some other ancient bacterium to form the eukaryotic cell. i. Eukaryotic cells have this internal division of labor among organelles that strongly resemble existing species of bacteria and cyanobacteria. ii. This internal division of labor was later extended to coördinating an external division of labor among different cell types and tissues, which is the basis for all complex multicellular organisms today, including human beings. iii. So, have some respect for that obscure crusty stuff on rocks, as it helps us understand the process by which the eukaryotic cell evolved, on which we're based!
C. These basic life forms are often refined for particular purposes with the following considerations: 1. Size, relative to each life form category, of course, e.g. a. In trees: tall might be >25 m; medium might be 8-25m; while small might be <8 m. b. For grasses, it would be more appropriate to define tall as > 1 m, medium as 15 cm-1 m, and small as <15 cm. 2. Coverage, that is, the percentage of ground covered by a given life form and its pattern of coverage. a. You could describe a woodland as having 25% tree coverage, 25% shrub coverage, 65% grass coverage, and 50% forb coverage. Yes, the percentages add up to more than 100% because the forb coverage, for example, might well be below the tree coverage, double-counting the ground cover. b. Pattern of coverage refers to its spatial distribution. i. Barren is the pattern that means members of a given life form are not present at all in a given vegetation (so ground coverage would be 0%). Examples might be the tree layer in a short-grass prairie or in a desert or in a tundra or in chaparral. ii. Discontinuous means that a given life form IS present in a vegetation, but it is rather sparse and open, with ground coverage below, say, 60%. More than that, the spatial distribution of the individual plants is random or even close to uniform. We're talking a thin but even representation of that life form. a. An excellent example is the shrub layer in a desert: They are found there, maybe with ground coverage of, oh, 10-15%, but far apart from one another (several meters). b. When you see a discontinuous pattern in vegetation, this sparse and random to uniform pattern, you are probably looking at intense competition for a scarce and unpredictable resource (water, in the desert). c. This competition can even be played out with chemical warfare among the plants, which is called "allelopathy." Plants can release chemicals that kill or suppress competitors for water: Volatile, aromatic compounds, such as terpenes, are used to pollute the local air against seedlings in an adult plant's root zone or the chemicals can be released from fallen leaves or from their roots. And here you thought plants just sat there passively: They are actually pretty active players in their environments. iii. Grouped or clumped means that members of a given life form are present in a vegetation, with, say, 10-50% ground coverage, but they occur in clusters, groups, or clumps. a. An example would be the oak tree coverage in a California oak woodland or oak park: Groups of oak trees cluster here and there in a gently hilly landscape, with grasses in between. b. This is a very common pattern, and it reflects the concentration of plants where a necessary resource tends to concentrate, such as water in uneven terrain. iv. Continuous means that a life form is present in great density and forms a pretty much even pattern: random or uniform. The coverage is very high, over 90%, so that there are hardly any breaks in the life form's ground coverage. a. An example would be the shrub cover in chaparral or the trees in a tropical rainforest or the grass in tall- grass prairie. b. This pattern and luxuriance indicates that the landscape provides a pretty even exposure to all resources needed by the species of a given life form. 3. Periodicity is sometimes used to modify life form descriptions. It refers to the duration of active photosynthesis each year. Are the plants evergreen or are they deciduous? a. Plants face a basic cost and benefit dilemma in maintaining the ability to photosynthesize all year round, when the declination of the sun shifts from the northern to the southern hemispheres each year, which affects sun angle, which affects the efficiency of photosynthesis. i. Plants "want" to photosynthesize all year round, to feed themselves. ii. The problem is they may face a lean season of inefficient photosynthess. This is a real problem, since their photo- synthetic structures (e.g., leaves) transpire water and burn energy during respiration no matter what season it is. b. So, if you're a plant, you have to "decide" whether to dump your photosynthetic structures (e.g., your leaves) during the lean season to avoid drying out or starving. The problem is that, if you do so, it's going to cost you a lot to rebuild all your leaves at the beginning of the good season. What to do? You have to "decide" whether the benefit of escaping dehydration and starvation is higher than the cost of rebuilding all your leaves each year. c. Some plants are evergreen: They photosynthesize (or remain capable of photosynthesis) all year round. i. Plants can get away with this if they live in the humid tropics and subtropics, where moisture, temperature, and sun angle don't vary all that badly over the course of the year. In this situation, there is no point to incurring the cost of dumping your leaves, and anyone who does is going to be a less effective competitor in a vegetation filled with other plants efficiently photosynthesizing and growing all year. So the humid tropics have almost exclusively broadleaf evergreen vegetation, and broadleaf evergreens and some needleleaf evergreens dominate the humid subtropics. ii. Other plants are evergreen, even in climates with rough seasons, but they have to come up with some way of coping with the hazards of variations in photosynthetic efficiency while respiring all year round. a. Many plants have small leaf surfaces as a way of retarding water loss during the rough season. The downside of this is their photosynthesis is also reduced in efficiency because there's less surface to photosynthesize with. This is less of a cost when everyone else in your climate is also dealing with the rough season by reducing leaf size. 1. We can see this effect in local oak trees, which have small leaves maybe up to 4 or 5 cm long; if you've traveled Down South, you may have noticed the oak trees there have humongous leaves, like 10-20 cm long! 2. Conifer trees are the experts in leaf surface area reduction: They've adopted the needleleaf form. They are needleleaf evergreens. b. Leafless evergreens take this photosynthetic-surface idea to an extreme: They have no leaves or just some vestigial leaves and they do the bulk of their photosynthesis in their stems, which stay green all year (a local desert example is a plant called Mormon tea, or Ephedra nevadensis). Note the green stems. c. Then there are succulent evergreens: They can conduct photosynthesis all year round in leaves and stems, whenever conditions are right, becaue they are protected from dehydration by tough, waxy surfaces and water storage tissues (e.g. jade plants, cacti). Of course, all that water might attract thirsty herbivores, so many succulents are fairly bristling with spines and thorns to discourage that. d. Other plants are deciduous: Leaves are shed seasonally, originally to avoid dehydration (by respiring and transpiring their water out of their leaf surfaces when little water is available in their root zones). i. Dry season deciduousness is the older form of deciduousness: Leaves are dropped to avoid respiration water loss and photosynthetic water need during drought. The deciduous habit probably evolved in tropical wet and dry climates and was then found adaptive to cold season conditions, after successful deciduous plants migrated poleward. ii. Icy winter deciduousness: Plants cannot absorb ice through roots, so leaves are dropped to avoid dehydration due to photosynthesis and respiration/transpiration. iii. Low sun angle deciduousness: Even if the ground is not frozen, low sun angle and short winter days make photosynthesis too inefficient to support leaf respiration 24/7, so leaves are shed to avoid "starving." iv. Now, California has a rough SUMMER season, but our native vegetations are dominated by EVERGREEN plants. This is because deciduousness originally evolved in a winter drought climate, and plants are primed to recognize the onset of winter. It would make sense to dump your leaves in the summer here, which is a lot like the winter in the tropical wet and dry climate, but plants can't reason and switch their behavior. So, winter deciduous plants are at a disadvantage here, because they dump their leaves during the only season that photosynthesis is possible in this subtropical climate: You only see them in washes (e.g., sycamores and alders) and near springs (e.g., a few deciduous oak species) where they can get water during the summer. The rest of the California vegetation is evergreen, and they get through the rough summer by other "xerophytic" adaptations ("xero" means "dry"): Small leaves, tough leathery leaves with few stomata ("sclerophyllous" leaves, meaning "tough" leaves), oils and resins in their leaves to retard water loss, allelopathic chemicals to protect their water supplies from competitors. Enough of the life form classification for plants! You need to be familiar with the major categories (i.e., trees; shrubs; lianas; herbs, including forbs, grasses, and ferns; bryophytes in general; and epiphytes). You should also remember how lichens illustrate symbiosis and that very ancient symbioses between various bacteria and cyanobacteria gave rise to the eukaryotic cell. Rememer that algae, fungi, plants, and animals are based on eukaryotic cells, so those ancient symbioses are important to understanding how all of us came to be. Also, familiarize yourself with common modifications to the basic categories of the life form system: size, coverage, and periodicity. This classification of individual plant types will be important later, when I use them to describe some of the common vegetation landscapes of the world.
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Rodrigue
First placed on web: 10/30/00
Last revised: 06/28/07