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Chapter 15 PROTISTA:
ALGAE AND HETEROTROPHIC PROTISTS
Life began about 3.5 billion years ago in the oceans with the appearance of prokaryotes. The oldest reliable date for the appearance of the eukaryotes is about 1.9 billion years ago, when the first members of a group of unicellular organisms called acritarchs appear in the fossil record in China. Acritarchs … Are probably the remains of a group of ancient eukaryotes Were plankton Some resemble dinoflagellates while others resemble green algae Their relationship among living organisms is uncertain http://www.ucl.ac.uk/GeolSci/micropal/acritarch.html http://www.geo.arizona.edu/palynology/ppacrtrc.html The kingdom Protista contains eukaryotes that cannot be assigned with certainty to other kingdoms The kingdom Protista is an artificial grouping and classification does not represent evolutionary relationships. This kingdom is also known as Protoctista. Fungi, plants and animals are all probably derived from protists. Protists covered in this course are those photosynthetic organisms that function like plants in ecosystems. They are the "grass of the ocean". Protists to be studied include:
- Algae: photosynthetic organisms studied by phycologists.
- Slime molds and oomycetes: heterotrophic organisms that are traditionally studied by mycologists , although these organisms are not fungi. Another group of protists not included in this course are the ciliates, flagellates, and other heterotrophs. The phylogenetic relationship among the different groups of protists is controversial, e.g. the relationship between the green and brown algae.
ECOLOGY OF THE ALGAE
Algae are dominant in salt and fresh water habitat. Everywhere they grow, they play a role similar to that of plants in terrestrial habitats.
Along rocky shores, the large and more complex members of the brown, red and green algae grow forming bands that reflect the ability of the seaweeds to withstand exposure. Seaweeds in this intertidal zone are exposed twice a day to large fluctuations of humidity, salinity and light, in addition to pounding action of the surf and forceful, abrasive water motions. Polar seaweeds endure months of darkness under the sea ice. Seaweeds are the food source to a host of herbivores and parasites. Large beds of seaweeds provide a safe habitat for many aquatic organisms, e.g. kelp beds off the coast of California. Plankton refers to all suspended drifting organisms found in all bodies of water. Planktonic algae and cyanobacteria constitute the phytoplankton found in oceans and fresh water. Heterotrophic plankton and usually swimming microorganisms are called zooplankton. Bacteria and some heterotrophic protists form the bacterioplankton. Phytoplankton is found at the base of the food chain. Colonial and single-celled chrysophytes, dinoflagellates, diatoms and green algae are the most important organisms at the base of the food chain in freshwater habitats. Unicellular and colonial haptophytes, dinoflagellates and diatoms are the primary producers of the ocean. In both marine and freshwater habitats, phytoplankton populations are kept in check by seasonal climatic changes, nutrient limitation and predation. Phytoplankton is the major producers of oxygen in the atmosphere. Phytoplankton reduces the amount of CO 2 in the atmosphere by fixing it during photosynthesis. Phytoplankton is important in the deposition of CaCO 3 deposits on the ocean floor. The CO 2 fixed by photosynthesis and the calcification process is replaced by atmospheric CO 2 Several types of multicellular algae are important members of coral reefs and deposit a substantial amount of calcium compound important in coral building. Some haptophyte protists produce substantial amounts of sulfur oxides that are added to the atmosphere and reflect sunlight helping to maintain a cooler temperature.
Phylum Dinophyta
The dinophyta are also known as dinoflagellates.
Mostly unicellular fresh water organisms; one colonial genus. Molecular evidence indicates that earlier euglenoids were phagocytic. About one third of euglenoids contain chloroplasts; their chloroplasts resemble those of the green algae and suggest that they were formed from endosymbiotic green algae. About two thirds of the genera are colorless heterotrophs that depend on particle feeding and absorption of dissolved organic compounds. They are mostly freshwater organisms living in waters rich in organic compounds and particles. Cell structure: Cell membrane, with pellicle immediately beneath the membrane. Lack cell wall; one genus has a wall-like covering made of manganese and iron minerals. The pellicle is made of protein strips arranged in the form of a helix; it may be rigid or flexible. Single flagellum for movement coming from the reservoir , and a second non-emergent flagellum. Flagellar swelling and the stigma or eyespot make the light-sensing system. Contractile vacuol e used in maintaining water balance. Pyrenoids are found in chloroplasts. It is a region where rubisco is found and paramylon , a polysaccharide is stored. Pigments present: chlorophylls a and b, carotenoids and several xanthophylls. Euglenoids grown in absence of light have been known to lose their chloroplasts and become heterotrophic. Reproduction in euglenoids is asexual, by mitotic cell division. Sexual reproduction is unknown. The nuclear membrane remains intact during mitosis in a way similar to the fungi. About 900 species are known. An intact mitotic nuclear envelope is probably a primitive condition. The break down of the nuclear membrane is probably a derived condition that appeared after euglenoids separated from the main stack of protists. http://botit.botany.wisc.edu/courses/botany_130/Diversity/Euglena/Euglena.html http://www.life.umd.edu/labs/delwiche/PSlife/lectures/Euglenophyta.html http://www.csupomona.edu/~jcclark/classes/bot125/resource/survey/euglenophyta.html
Phylum Cryptophyta
Members of this phylum are known as cryptomonads. Fresh and salt-water unicellular flagellates found usually in cold or subsurface waters. Cells are small, 3 to 50 micrometers. Cryptomonads are important components of the phytoplankton and are readily ingested by zooplankton.
Cryptomonads are important components of the phytoplankton and are readily ingested by zooplankton. Cryptomonads are rich in polyunsaturated fatty acids, which are essential for the growth and development of zooplankton. Species may be photosynthetic or phagocytic. Photosynthetic species contain in their chloroplast chlorophyll a and c, α-carotene, a phycobilin (either phycoerythrin or phycocyanin). Unlike other algae, cryptomonads lack β-carotene. The chloroplast has four bounding membranes: The outer of the four membranes is continuous with the nuclear envelope and is called the chloroplast ER. The space between the second and third membrane contains starch grains and the remains of a nucleus, the nucleomorph , complete with three chromosomes and a nucleolus with typical eukaryotic RNA. It is theorized that a unicellular red alga was ingested and became an endosymbiont of a heterotroph. The chloroplast in the red alga is the remnant of a primary endosymbiont. The red alga is a secondary endosymbiont in the heterotroph cell, the cryptomonad. The cells have two anterior slightly unequal flagella of about the same length as the cell. About 200 species are known.
Phylum Haptophyta
Haptophytes are mostly marine phytoplankton but terrestrial and freshwater species are known. About 300 known species in 80 genera found mostly in the tropics. Distinguished by a haptonema , a threadlike structure often coiled used for attachment. Haptonema microtubules do not have the 9 + 2 arrangement typical of eukaryotic cilia and flagella. The haptonema can be bent and coiled but cannot beat. It helps in catching prey or may in sensing and avoiding obstacles. There are two flagella of equal length present. Coccoliths present on the outer surface of the cell of 12 or more families. Coccoliths are calcified scales made of organic material. There are two types of coccoliths, one is made in the Golgi apparatus and transported to the surface of the cell, and the other is formed outside the cell. Coccolithoforids appeared in the Late Triassic some 230 million years ago.
Their food reserve is in the form of glycogen. Asexual reproduction is by means of motile zoospores , which have the characteristic two flagella of heterokonts. Sexual reproduction is oogamous : one gamete large and nonmotile, the other small and motile. Eggs are produced in the oogonia. The antheridium contains many male nuclei. The fertilized egg forms a thick-walled zygote called the oospore. The oospore serves as a resting stage during stressful conditions. Oomycetes are also called water molds, white rusts and downy mildew. Water Molds Are Aquatic Oomycetes. Abundant in fresh water. Mostly saprophytic and a few parasitic including species that cause diseases to fish and fish eggs. Species may be homothallic or heterothallic. Saprolegnia and Achlya are common water molds that reproduce sexually and asexually. Some Terrestrial Oomycetes Are Important Plant Pathogens Terrestrial oomycetes produce motile zoospores when water is available. Terrestrial oomycetes are important plant pathogens; the genus Phytophthora is particularly destructive to plants. They attack important crops like grapes, pineapples, onions, strawberries, apples, citrus fruits, cacao, etc. Phytophthora cinnamomi killed millions of avocado trees in southern California, and destroyed thousands of hectares of Eucalyptus timberland in Australia. Phytophthora ramorum was the cause of the disease called “the sudden oak death.” It attacks many species of oaks and also 26 other species of plants including firs and coastal redwoods. The great potato famine in Ireland (1846) was caused by the oomycete Phytophthora infestans. A gene has been found in a species of wild potato, Solanum ulbocastanum , from Mexico, that is resistant to potato blight. The resistant gene has now been inserted in the commercial potatoes, Solanum tuberosum. The genus Pythium attacks and rot seeds in the wild (preemergence damping-off) and seedling (postemergence damping-off)
Phylum Bacillariophyta
An ancient group that appeared in the fossil record about 250 million years ago, and became abundant in the fossil record about 100 million years ago during the Cretaceous.
Diatoms are unicellular or colonial organisms that form an important component of the phytoplankton. They may count for as much as 25% of the primary production of the earth. There may be as many as 100,000 species, some of the most diverse and abundant algae on earth. Diatoms are the primary source of food for many marine animals; they provide essential carbohydrates, fatty acids, sterols, and vitamins to the consumers. Diatoms live in both freshwater and marine habitats, but are especially abundant in cold marine waters. Diatoms can also inhabit terrestrial habitats such as damp cliff faces, moist tree trunks and on the surfaces of buildings. The Walls Of Diatoms Consist Of Two Halves Cell wall in two parts known as frustules , are made of polymerized silica (SiO 2 H 2 O, 95%) and carbohydrates especially pectin (5%). The shell is composed of an upper and lower half, with the lower half fitting neatly within the upper, like a Petri dish. The shell is highly ornamented and perforated with microscopic holes so precisely spaced that they are used commercially to test the resolution of expensive microscope lenses. These holes connect the living protoplast with the external environment. Freshwater forms are usually cylindrical in shape: pennate. Marine species are usually spherical or circular: centric. Reproduction In Diatoms Is Mainly Asexual Reproduction is usually asexual. Changes in the environment or critical small size triggers sexual reproduction. Before a diatom can undergo mitosis, it must be living in an environment with sufficient silicon to allow it to construct a new shell. The diploid protoplast undergoes typical mitosis within the shell, and then the two-shell halves separate. One protoplast gets the top half, and the other gets the bottom half. In either case, the protoplast then secretes a new "bottom" to the "Petri dish"(i.e., a new half fitting inside the old half). This means that after every mitotic division, one of the resulting diatoms is smaller than the original. This can go on for several generations. Eventually, the protoplast inside the tiny shell undergoes meiosis rather than mitosis. Four haploid gametes are released from the shell, which is discarded. When two gametes meet and fuse, the resulting diploid cell is called an auxospore (zygote). The auxospore grows into a normal size of the species. It then secretes a silica case of the original size...and the cycle begins anew.
Phylum Phaeophyta
Phaeophytes are also known as brown algae It is an entirely marine group especially abundant in temperate and cold waters. Common in the intertidal and subtidal zones; dominant alga of rocky shores. About 1,500 species. The Thallus Size - few are microscopic, most much larger - up to 60 m. Larger forms with complex structure. There are no known unicellular or colonial representatives of this group. The simplest form of plant is a branched, filamentous thallus (pl. thalli): a relatively undifferentiated vegetative body. The thalli range in complexity from simple branched filaments to aggregation of branched filaments called pseudoparenchyma. Adjacent cells are connected by plasmodesmata without desmotubules connecting the ER. Pigments Cells contain numerous disk-shaped, golden-brown plastids that are similar both biochemically and structurally to those of chrysophytes and diatoms. Chlorophyll a and c (no Chlorophyll b), ß-carotene, fucoxanthin and other xanthophylls. Food reserves are typically complex polysaccharides, sugars and higher alcohols and sometimes fats. Glucose and mannitol are polymerized together as laminarin. Mannitol is a six-carbon sugar-alcohol; it is linked together with glucose in a beta-1, linkage. The principal carbohydrate reserve is laminarin and true starch is absent. There are two groups based on the presence or absence of pyrenoids. Kelps Kelps ( Macrocystis and Nereocystis ) and rockweeds have a highly differentiated bodies The walls are made of cellulose and algin , an alginic acid, a long-chained heteropolysaccharide. Some have stem-like, root-like, leaf-like organs. Since they do not have vascular systems, these structures are not true stems, roots, or leaves. Termed rhizoid , holdfast , stalk or stipe , and blade.
Kelps have a meristematic region between the stipe and the blade. Sargassum and Fucus grow from repeated divisions from a single apical cell. Some species have floatation bladders. Some free-floating species have lost the holdfast. Some of the kelps have modified elongated cells in the center of the stipe that are capable of conducting carbohydrates from the blades near the water surface to the lower parts of the alga. Some brown algae have evolved sieve tubes comparable to those found in food-conducting tissue of vascular plants. Sieve tube elements are joined end-on-end by the sieve plates. Of great economic importance: fertilizer, food especially in Japan, source of algin - stabilizer & moisture retainer in many products such as ice cream, cake frosting, paint, pharmaceuticals, processing of natural and synthetic rubber. Life Cycle Their life cycle involves an alternation of generation, and meiosis occurs during spore formation ( sporic meiosis ). The gametophytes of the primitive brown algae produce reproductive structures called plurilocular gametangia. They may function as male or female gametangia or produce flagellated haploid spores that give rise to new gametophytes. The diploid sporophyte produces both plurilocular and unilocular sporangia. The plurilocular sporangia produce diploid zoospores that produce diploid sporophytes. Meiosis takes place in the unilocular sporangia producing haploid zoospores that germinate to produce haploid gametophytes. Zoospores have tinsel and whip flagella. Some groups (e.g. Fucus ) do not form spores and have a gametic life cycle without alternation of generations.
Phylum Rhodophyta
Red algae are mostly marine organisms found in tropical and warm waters. Fewer than 100 species occur in fresh water. Some occur in cooler regions of the world. Many species are found in very deep water. There are 4100 to 6000 known species. Red algae are mostly structurally complex multicellular organisms with very few species unicellular or microscopic filaments. They may grow attached to the substrate, submerged vegetation and a few are free floating.
The resultant diploid zygote then produces a few diploid carpospores , which are release into the water. Carpospores produce sporophytes that form haploid spores, which in turn produce new gametophytes. In some red algae, the zygote produces a carposporophyte generation, which remains attached to the parent gametophyte. The carposporophyte divides mitotically and eventually produces carpospores. The carpospores are released and settle onto a substrate, and grow into separate diploid sporophytes. In many red algae, the diploid zygote is transferred to another cell of the gametophyte called the auxiliary cell where it proliferates into many carpospores. The carpospores produce a new generation called the tetrasporophyte. Meiosis occurs I in specialized cells of the tetrasporophyte , called the tetrasporangia. Each tetraspore germinates into a gametophyte.
Phylum Chlorophyta
Also known as green algae. A diverse group of about 17,000 species. Most chlorophytes are aquatic, but some green algae can live on the surface of snow, on tree trunks, in soils, or symbiotically with protozoans, hydras or lichen-forming fungi. Chlorophytes range in size from microscopic to quite large: unicellular, colonies, branched and unbranched filaments, thalloid. Green algae have chlorophylls a and b and store starch as a food reserve inside their plastids. Most green algae have firm cell walls composed of cellulose, hemicellulose and peptic substances. The flagellated reproductive cells of some green algae resemble that of plant sperm. Based on studies of mitosis, cytokinesis, reproductive cells and molecular similarities, the green algae have been divided into several classes. Three of these classes will be studied here: Cytokinesis in the Chlorophyta The classes Chlorophyceae and Ulvophyceae form a phycoplast during cell division, which is system of microtubules parallel to the plane of cell division. Nuclear envelope persists during mitosis. Mitotic spindle forms and then disappears at telophase. Daughter nuclei are separated by the phycoplast in which the microtubules lie perpendicular to the axis of division.
The role of the phycoplast is presumed to ensure that the cleavage furrow will pass between the two daughter nuclei. Cytokinesis is by cell plate formation or development of a furrow. The Chlorophyceae form four narrow bands of microtubules known as flagellar roots, which are associated with the flagellar basal bodies (centrioles) of the flagella. The Ulvophyceae have a persistent spindle but do not develop a phragmoplast or cell plate. The class Charophyceae does not form a phycoplast but develop a phragmoplas t like land plants. Formation of a phragmoplast, which is parallel-aligned microtubules and microfilaments at right angles to the forming cell plate , is to generate a guiding and supporting matrix for the deposition of new cell plate. The phragmoplast is a system of microtubules, microfilaments and ER vesicles that is oriented perpendicular to the plane of division. It serves in the assembling of the cell plate and the cell wall. As the cell plate matures in the center of the phragmoplast, the phragmoplast and developing cell plate grow outward until they reach the of the dividing cell. See pages 64-67in your textbook. Spindle is persistent through mitosis. Cytokinesis is by cell plate formation or furrowing, just like bryophytes and vascular plants. The flagellar root system of microtubules provides anchorage to the flagellum. The multilayered structure is often associated with one of the flagellar roots. The type of multilayered structure is often an important taxonomic character. The flagellar root had multilayered structure of the Charophyceae is very similar to that found in the sperm of bryophytes and some vascular plants. Class Chlorophyceae There are approximately 350 genera and 2650 living species of chlorophyceans. Mostly freshwater species. They come in a wide variety of shapes and forms, including free-swimming unicellular species, colonies, non-flagellate unicells, filaments, and more. Cytokinesis may be by furrowing or by cell plate formation. When flagellate, the flagella are apical and equal in length, and directed forward. They also reproduce in a variety of ways, though all have a haploid life cycle, in which only the zygote cell is diploid. The zygote will often serve as a resting spore, able to lie dormant though potentially damaging environmental changes such as desiccation. Chlamydomonas is motile unicellular chlorophyte. Two equal flagella.
Majority has one plane of division, unlike the Ulva with three planes Spindle and nuclear envelope persist through mitosis. Flagellated cells may have two, four or many flagella directed forward Alternation of generations with a haploid gametophyte and diploid sporophyte. They have sporic meiosis or a diploid, dominant life history involving gametic meiosis. Cladophora is a filamentous septate ulvophyte. It forms large blooms in fresh water. There are both marine and fresh water species of Cladophora. Each cell is multinucleated and has one single, peripheral, net-like chloroplast with many pyrenoids. Marine species have an alternation of isomorphic generations. Most of the fresh water species do not have an alternation of generations. Ulva consists of a two-cell thick flat thallus that may grow up to a meter in length. It is known as sea lettuce. Ulva is anchored to the substrate by a holdfast produced by extensions of the cells at its base. The cells of the thallus are uninucleate and have one chloroplast. Ulva is anisogamous and has an alternation of isomorphic generations. See the Life Cycle of Ulva on page 335. Codium and Halimeda are examples of siphonous marine algae. Very large, coenocytic cells that are rarely septate characterize siphonous algae. Cell walls are only produced during reproduction. Siphonous green algae are diploid, with gametes being the only haploid stage. Halimeda has calcified cell walls. Examples to study: Thalloid: Ulva. Siphonous: Acetabularia, Codium, Ventricaria, Halimeda. Filamentous septate: Cladophora. Class Charophyceae Growth habit may be unicellular, filamentous, colonial or thalloid (parenchymatous). Considered closely related to plants due to structural, biochemical and genetic similarities. The orders Coleochaetales and Charales have plant-like characteristics. These include: Asymmetrical flagellated cells always have two flagella. Breakdown of the nuclear envelope at mitosis Persistent spindles or phragmoplast at cytokinesis. Presence of phytochrome, flavonoids and chemical precursors of the cuticle.
Other molecular features. Spirogyra is an unbranched, filamentous charophyte. Found in fresh water, often forming blooms. Cells uninucleate. Filaments are surround by a watery sheath. Chloroplasts one or more, flat ribbon-like with numerous pyrenoids. Asexual reproduction occurs by fragmentation. There are no flagellated cells at any stage of its life cycle. Sexual reproduction takes place through the formation of a conjugation tube. The cytoplasm of one cells migrates to the other cell and function as isogametes. The zygote becomes surrounded by a thick wall of sporopollenin. Meiosis is zygotic. Desmids are a large group of fresh water charophytes. Lack flagellated cells. Desmid cells consist of two sections of semi-cells joined by a narrow constriction. Sexual reproduction is similar to Spirogyra. Two orders of Charophyceae, the Coleochaetales and the Charales , resemble bryophytes and vascular plants. They have plant-like microtubular phragmoplast operating during cytokinesis. They are oogamous and their sperm are ultrastructurally similar to those of bryophytes. Morphological and molecular studies indicate that an early basal split in the green algae gave rise to a chlorophyte clade containing most of the green algae, and a streptophyte clade that includes the Coleochaetales and Charales, zygnematalean green algae, and land plants (bryophytes and vascular plants). Coleochaetales Include branched filamentous and discoid genera. Growth occurs at the apex or peripheral cells, and the plant is anchored in mud or silt by translucent rhizoids. Coleochaete has uninucleate vegetative cells that each contains one large chloroplast with an embedded pyrenoid. It reproduces asexually by zoospores that are formed singly within cells. Sexual reproduction is oogamous. The zygotes remain attached to the parental thallus, which stimulate the growth of a layer of cells that covers the zygotes. These parental cells have wall ingrowths are believed to function in nutrient transport between gametophyte and sporophyte. Charales The thallus in some stoneworts is encrusted with white lime, giving a crusty texture (hence the name brittlewort). The Charales exhibit apical growth.
The tubules are composed of slightly solidified protoplasm through which more liquefied protoplasm flows rapidly. The foremost edge of the plasmodium consists of a thin film of gel separated from the substrate by only a plasma membrane and a slime sheath. Myxomycetes in the feeding stage are most often found under the bark of decaying logs or between layers of leaf litter. Plasmodia may be white or bright red, orange or yellow. When the food supply is exhausted or other unfavorable conditions occur, the plasmodium changes, taking on the appearance of a fungus. These structures, called fruiting bodies, contain the reproductive spores, which, when released, germinate and begin the life cycle anew. The protoplasm of the young fruiting body contains many nuclei that increase in number by mitosis. Progressively, the protoplasm is cleaved into a large number of spores, each containing a single diploid nucleus. Meiosis occurs giving rise to four haploid nuclei, three of which disintegrate leaving a single haploid nucleus in each spore. There are three types of fruiting bodies: sporangium , aethalium and plasmodiocarp. Spores are very resistant to environmental extremes and are long-lived. Spores germinate to produce amoebas and flagellated cells. Amoebas and flagellated cells are interconvertible, multiply by mitosis and eventually act as gametes (plasmogamy). Amoebas may secrete a thin wall during unfavorable periods and become dormant for a year or more. This resting stage is called a microcyst. During dry periods the plasmodium becomes encysted and forms a sclerotium. See the Life Cycle on page 341. http://www.botany.hawaii.edu/faculty/wong/Bot201/Myxomycota/Myxomycota.htm http://www.cs.cuc.edu/~tfutcher/Slimemolds.html http://www.uoguelph.ca/~gbarron/myxoinde.htm
Phylum Dictyosteliomycota
Dictyostelids are also known as cellular slime molds. There are about 50 known species in four genera. They may be most closely related to amoebas, phylum Rhizopoda, than to any other group.
Members in this phylum exist in a uninucleated haploid myxamoeba form, deriving nutrients via phagotrophic assimilation. Dictyostelids have cellulose-rich cell wall during a stage of their life cycle. They undergo mitosis similar to plant and animals in which the nuclear envelope breaks down. Dictyostelids have centrioles. When their food grows scarce, the myxamoebas agglomerate into a single "colonial'' organism, resembling an amoeba, which feeds for a while before fixing itself in place and growing a stalk. A chemical called acrasin , now known to be cyclic AMP that they themselves give off attracts the amoebas to each other; their cell walls become sticky and contribute to the formation of the cellular mass of pseudoplasmodium. The uninucleate, haploid myxamoebas retain their individuality in the mass of 10,000 to 120, cells. This mass is called a pseudoplasmodium (also called a grex or slug ), which in a cooperative effort between individual cells, produces a fruiting body called a sorocarp. The sorocarp then gives rise to spores. The eventual fate of an individual cell is determined at an early stage by its position in the aggregation, which appears to be controlled by the stage of the cll cycle the individual cell was in when aggregation began. Cells in the anterior end of the slug develop into the stalk of the sporangium; these cells become covered with cellulose and undergo apoptosis, programmed cell death. The posterior cells of the pseudoplasmodium move to the top of the stalk and become dormant spores. Eventually the spores are dispersed and each releases a single myxamoeba. The fusion of the sex cells (individual amoebas) produces a large resistant body called a macrocyst , which upon germination produces new cells capable of continuous asexual reproduction. http://www.csupomona.edu/~jcclark/classes/bot125/resource/survey/dictyosteliomycota.html
