Moss – Definition, Types, Life Cycle, Importance, Examples

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What is Moss?

  • Mosses, belonging to the division Bryophyta within the vast kingdom Plantae, are non-vascular plants that have evolved to thrive not only in the stereotypical damp and shaded habitats but also in sunnier and arid locales. With a staggering diversity, over 12,000 identified species of moss are distributed across 8 classes and 23 genera.
  • Scientifically referred to as Bryophyta sensu stricto, mosses are distinct from the broader group of bryophytes, which encompasses liverworts and hornworts. Characteristically, mosses manifest as dense green aggregations, often forming mats in moisture-rich or shaded terrains.
  • The individual moss plants are primarily composed of uncomplicated leaves, typically a single cell layer thick, affixed to a stem that may or may not exhibit branching. This stem plays a minimalistic role in the transportation of water and nutrients. Notably, while some moss species possess conducting tissues, these are rudimentary and differ structurally from the conducting tissues observed in vascular plants.
  • Mosses exhibit a unique reproductive strategy. They lack seeds and, post-fertilization, give rise to sporophytes characterized by unbranched stalks crowned with spore-containing capsules. Typically, these plants range in height from 0.2 to 10 cm, with exceptions like Dawsonia, which can attain a height of 50 cm, making it the tallest known moss.
  • It’s crucial to differentiate mosses from liverworts, hornworts, and lichens. While they might superficially resemble mosses, lichens are a result of fungal symbiosis and are not genetically related to mosses. Moreover, contrary to popular belief, several mosses do possess advanced vascular systems. In the life cycle of mosses, the haploid gametophyte generation is dominant, a pattern that contrasts with vascular plants where the diploid sporophyte generation prevails.
  • From a commercial perspective, mosses are primarily valued for their role in forming peat, especially the genus Sphagnum. Their decorative appeal is also leveraged in horticulture and floristry. Historically, their insulating properties and impressive liquid absorption capacity, up to 20 times their weight, have been utilized.
  • In the broader context, the kingdom Plantae encompasses all plant forms on Earth. These multicellular eukaryotes are distinguished by the presence of a cell wall and chlorophyll, vital for photosynthesis, enabling autotrophic nutrition. The vastness of the plant kingdom necessitates its categorization into multiple subdivisions.
  • Grasping the intricacies of the Kingdom Plantae is foundational to understanding the diverse plant forms, ranging from the diminutive algae to the towering Sequoia. While some plants are adorned with flowers, others exhibit basic thalloid structures or possess well-defined root, shoot, and leaf systems.

Definition of Moss

Mosses are small, non-vascular plants belonging to the division Bryophyta that typically form dense green mats or clumps in damp or shaded environments, lacking seeds and reproducing through spores.

Habitat of Mosses

Mosses, representing a diverse group within the plant kingdom, exhibit a wide range of habitat preferences. Their adaptability and resilience allow them to colonize various ecological niches. Here’s an overview of the habitats of mosses:

  1. Forest Environments:
    • Mosses frequently form dense colonies in cool coastal forests and high-altitude or latitude moss forests. The forest floor, especially beneath conifers, often showcases a lush carpet of mosses, providing a cool, moist environment conducive to their growth.
  2. Rocky Surfaces:
    • Mosses have the ability to colonize basalt flows, as observed in regions like Iceland. The adaptability of mosses allows them to thrive on rocks, often forming a green layer over them.
  3. Water-Rich Areas:
    • Mosses are commonly found growing along streams, seeps, springs, and in areas with travertine deposits. Their affinity for moisture-rich environments enables them to flourish in such habitats.
  4. Urban Environments:
    • Mosses can colonize various urban structures, from brick walls to concrete surfaces. Their resilience allows them to thrive even in disturbed, sunny areas of cities.
  5. Ecological Adaptations:
    • Mosses display a wide range of substrate preferences, from rocks, exposed mineral soil, and tree trunks to bogs and marshes. Their habitat choice can be influenced by factors such as soil acidity, moisture levels, and light availability.
    • While mosses often grow as epiphytes on trees, they are not parasitic and do not derive nutrients from the trees.
  6. Desiccation Tolerance:
    • Mosses possess the remarkable ability to survive desiccation. They can endure dry periods, only to revive and resume their metabolic activities upon rehydration.
  7. Geographical Orientation:
    • In the Northern Hemisphere, the north side of structures, such as trees and rocks, often supports more luxuriant moss growth due to reduced sunlight exposure and resultant moisture retention. Conversely, in the Southern Hemisphere, the south side may exhibit denser moss colonization.
  8. Aquatic Habitats:
    • Some moss species are adapted to completely aquatic environments or waterlogged conditions. These mosses can grow to considerable lengths, especially in slow-moving waterways or bogs.
  9. Symbiotic Relationships:
    • In boreal forests, certain moss species establish symbiotic relationships with nitrogen-fixing cyanobacteria. This association benefits the ecosystem by enhancing nitrogen availability.

In essence, mosses, with their diverse habitat preferences, play crucial ecological roles, from nitrogen fixation to soil stabilization. Their presence across various environments underscores their adaptability and ecological significance.

Characteristics of Mosses

Mosses, classified under the order Bryophyta, are non-vascular plants characterized by several distinct features that set them apart from other plant groups.

  1. Life Cycle Dominance: Mosses predominantly exist in the haploid gametophyte stage, contrasting with vascular plants where the diploid sporophyte stage is more dominant. The sporophytes in mosses are ephemeral and rely heavily on the gametophyte for their nutritional and water needs.
  2. Leaf Structure: Moss leaves are simple, often composed of a single cell layer. They are spirally arranged, and some exhibit a single row of cells with pronounced midribs. Unlike vascular plants, their leaves lack internal air spaces.
  3. Absence of True Roots: Instead of conventional roots, mosses possess rhizoids – thread-like structures that anchor them to their substrate. Notably, these rhizoids do not absorb water or nutrients like roots in vascular plants. This feature differentiates mosses from liverworts, as mosses have multicellular rhizoids.
  4. Stem Characteristics: The stems of mosses are typically weak, free-standing, and exhibit colors ranging from green to brown. The gametophyte stems can manifest in various forms – straight or curved, simple or branched.
  5. Reproductive Features: Mosses lack flowers and reproduce through spores. Their spore-bearing capsules, or sporangia, are borne individually on elongated, unbranched stems. These capsules mature before the elongation of their supporting stalks, a feature distinguishing most mosses from liverworts.
  6. Non-vascular Nature: Mosses do not possess water-bearing xylem tracheids or vessels, setting them apart from vascular plants. They primarily absorb water and nutrients through their leaves, utilizing photosynthesis to synthesize food.
  7. Absence of Mycorrhizal Associations: With the exception of the class Takakiopsida, mosses do not form mycorrhizal relationships.
  8. Stomatal Presence: While early diverging classes like Takakiopsida, Sphagnopsida, Andreaeopsida, and Andreaeobryopsida may lack stomata or possess non-functional pseudostomata, stomata have been lost in the subsequent classes over 60 times.
  9. Sporophyte Dependency: The sporophytes, which are short-lived, may be capable of photosynthesis but are largely dependent on the gametophyte for their sustenance.
  10. Distinguishing Features: Mosses can be differentiated from polysporangiophytes and liverworts by their distinct stem, simple non-vascular leaves not arranged in three ranks, and the presence of capsules containing spores on unbranched stems.

In summary, mosses, with their unique characteristics, play a pivotal role in terrestrial ecosystems, contributing to soil formation, water retention, and serving as habitats for numerous microorganisms and invertebrates.

Classification of Mosses

Mosses, some of the most ancient terrestrial plants, have a rich taxonomic history. Historically, all mosses were grouped under the phylum Bryophyta. Within this classification, the true mosses, often referred to as Musci, constituted one of the three primary divisions of Bryophyta. However, recent taxonomic advancements, driven by distinctions in thallus and sporangia structures, have proposed a reclassification where Bryophyta is a subset of mosses. This contemporary classification delineates the following groups:

  1. Liverworts (Hepatophyta):
    • Liverworts are characterized by a flat thallus that rests on the substrate, anchored by rhizoids.
    • The thallus typically contains oil cells.
    • Distinctive reproductive structures, gametangiophores, manifest as umbrella-like structures. These can be male (antheridiophores) or female (archegoniophores). Notably, the female archegonia open downwards, while the male antheridia open upwards.
    • Upon maturation, the sporophyte’s sporangium slightly protrudes due to the elongation of its base stalk, the seta.
    • Spore dispersal is facilitated by grooves on the spore caps, which release spores within hours of opening.
    • A representative example is Marchantia.
  2. Hornworts (Anthocerotophyta):
    • Resembling liverworts, hornworts have a more plantlet-like appearance.
    • Each cell typically contains one or two large chloroplasts, and they lack oil cells.
    • Gametangia are embedded in the upper part of the thallus.
    • The mature sporangia, or sporophytes, rise above the thallus, resembling small horns. These are formed due to robust outgrowths from the seta.
    • The spores can be dispersed over several weeks once the sporangium grooves open.
    • With just over 100 identified species, hornworts are relatively rare, with limited presence in regions like the Netherlands.
  3. True Mosses (Bryophyta sensu stricto):
    • True mosses exhibit a distinct architecture, resembling stems and leaves, despite lacking genuine stems, leaves, roots, or specialized tissues.
    • Gametangia develop at the apex of both male and female gametophytes.
    • Post-fertilization, the zygote metamorphoses into a sporophyte, which remains attached to the female gametophyte.
    • The sporophyte comprises a seta and a capsule, topped with an operculum. As the capsule matures, the operculum detaches, revealing the peristome’s teeth at the capsule’s opening.
    • The elongation of the seta elevates the sporangium, exposing it to wind currents.
    • Within the capsule, meiosis occurs in spore-producing cells, yielding haploid spores, which can take days for dispersal.
    • True mosses are the most diverse and prevalent group within the Bryophytes, with over 500 species found in places like the Netherlands. Examples include Sphagnum, Polytrichum, and Funaria.

In summary, mosses, with their intricate classification and diverse forms, offer a fascinating glimpse into the evolutionary history of terrestrial plants.

Types of Mosses

Mosses, with their diverse forms and ecological roles, are classified into various types based on their morphological and habitat characteristics. Here, we delve into the primary types of mosses:

  1. Peat Moss (Sphagnopsida):
    • Commonly referred to as sphagnum or peat moss, members of the Sphagnopsida class predominantly inhabit moist, bog-like environments, often forming extensive, cushiony masses.
    • These mosses have applications in horticulture, especially as soil amendments, though they can acidify the soil. Consequently, acid-loving plants like blueberries and cranberries are often found in proximity to peat moss.
    • Peat moss is among the plants that can decompose into peat, a material historically used as an alternative fuel for cooking and heating in certain European regions.
  2. Rock Moss (Andreaeopsida):
    • Also termed lantern mosses, members of the Andreaeopsida class are typically found growing on rocky substrates.
    • Many species within this class form tufted growths and exhibit a distinctive black hue.
    • The nomenclature “lantern” is derived from certain plant parts resembling lanterns. These mosses are adapted to thrive in cold climates and are often found at high latitudes.
  3. Polytrichopsida:
    • Mosses of the Polytrichopsida class are recognized as pioneer species, capable of colonizing areas after disturbances like floods or fires.
    • This class boasts a diverse array of species, ranging from the towering Dawsonia superba, the world’s tallest moss, to other diminutive species. Their adaptability allows them to flourish in various challenging habitats.
  4. Bryopsida:
    • Encompassing a staggering 95% of all moss species, the Bryopsida class is the most diverse group of mosses.
    • A unifying characteristic of this class is the presence of a toothed spore capsule. This capsule safeguards the spores, which, upon dispersal, germinate to give rise to new moss plants.
    • The unique “ring of teeth” structure on the spore capsules facilitates the efficient dispersal of spores when conditions are conducive.

In conclusion, mosses, with their myriad types, play pivotal roles in various ecosystems, from bogs to rocky terrains, showcasing the adaptability and resilience of these ancient plants.

Life Cycle of Mosses

Mosses exhibit a distinctive life cycle characterized by the alternation of generations, oscillating between the haploid gametophyte stage and the diploid sporophyte stage.

Life cycle of a typical moss (Polytrichum commune)
Life cycle of a typical moss (Polytrichum commune) | Image Credit: LadyofHats, Public domain, via Wikimedia Commons
  1. Gametophyte Phase:
    • The gametophyte, being haploid, differentiates into structures resembling stems and leaves. At the apex of these structures, the sex organs develop.
    • The female reproductive organ, termed the archegonium, is reminiscent of a bottle-like structure, composed of a single cell layer. Surrounding the archegonium are specialized leaves known as the perichaetium, which offer protection.
    • Conversely, the male reproductive organ, the antheridium, is a diminutive, stalked, and club-shaped entity. It is safeguarded by modified leaves termed the perigonium.
    • Upon maturation, the antheridium releases biflagellate sperm cells, or antherozoids. These motile cells navigate through water to reach and fertilize the egg housed within the archegonium.
    • This fertilization event culminates in the formation of a diploid zygote, marking the inception of the sporophyte phase. Subsequent cellular divisions in the archegonium lead to the creation of the calyptra, a protective sheath for the emerging sporophyte’s capsule.
  2. Sporophyte Phase:
    • The zygote evolves into the sporophyte, comprising a foot, an elongated stalk termed the seta, and a spore-containing capsule capped by the operculum.
    • Intriguingly, the sporophyte remains parasitically tethered to the gametophyte, relying on it for sustenance and hydration.
    • Within the capsule, specialized cells undergo meiosis to yield haploid spores.
    • The capsule’s aperture is fortified with tooth-like structures, the peristomes, which deter spore release during damp conditions.
    • Upon maturation, the operculum and peristome detach, facilitating spore dispersal.
    • Landing on conducive substrates, these spores germinate into protonema, filamentous entities that serve as precursors to the gametophyte, thus completing the cyclical process.
  3. Asexual Reproduction:
    • Mosses also possess the capability for asexual reproduction. Fragmentation of stems or leaves can lead to the formation of independent entities, which subsequently develop into new moss plants.

In summation, the life cycle of mosses is a complex interplay between two distinct phases, each with its unique morphological and reproductive attributes, ensuring the perpetuation and diversity of these ancient plants.

Cultivation of Mosses

Mosses, with their delicate and intricate structures, have been revered for their aesthetic appeal, especially in Japanese gardening traditions. Their cultivation, however, is not as straightforward as that of many other plants, and their growth requirements can be quite specific.

  1. Aesthetic Appreciation:
    • Mosses are often encouraged to grow in gardens, particularly in Japanese settings, where they impart a sense of tranquility, antiquity, and stillness. They can carpet forest scenes in old temple gardens, enhancing the serene ambiance.
    • In bonsai culture, moss is used to cover the soil, further emphasizing the impression of age and timelessness.
  2. Cultivation Techniques:
    • While there aren’t universally established cultivation rules, moss collections often commence with samples transplanted from the wild, preserved in moisture-retaining bags.
    • Cultivating moss from spores is a less controlled process. Given the right conditions, moss spores can colonize hospitable surfaces within a few years. Porous and moisture-retentive materials, such as brick and wood, are particularly conducive to moss growth. Acidic preparations, including mixtures of buttermilk, yogurt, and moss samples, can also promote moss colonization.
  3. Moss Lawns:
    • In regions like the Pacific Northwest, where the climate is cool, humid, and cloudy, moss is sometimes cultivated as a lawn alternative. Such moss lawns are low-maintenance, requiring minimal mowing, fertilizing, or watering.
  4. Green Roofs and Walls:
    • Mosses are employed in green roofs due to their numerous benefits, including reduced weight, enhanced water absorption, and drought tolerance. Their lack of true roots means they need less planting medium, making them a sustainable choice for green roofs and walls.
  5. Mossery:
    • The 19th-century fascination with moss-collecting led to the creation of “mosseries” in gardens. These structures, open to the north to ensure shade, housed moss samples, which were regularly moistened to promote growth.
  6. Aquascaping:
    • Aquatic mosses are popular in aquascaping. They thrive in environments with low nutrients, light, and temperature. Their slow growth and hardiness make them suitable for aquarium settings, where they also help maintain optimal water chemistry for aquatic life.
  7. Growth Inhibition:
    • While mosses are appreciated for their beauty, they can be problematic, especially in nurseries and greenhouses. Moss growth can be curtailed by methods such as improving drainage, increasing sunlight exposure, and enhancing soil pH. Chemical treatments, like ferrous sulfate, can also be used to combat moss growth. However, it’s essential to modify the growth conditions to prevent moss regrowth.

In conclusion, the cultivation of mosses requires a nuanced understanding of their specific needs and growth conditions. Whether they’re being grown for their aesthetic appeal or other purposes, mosses offer unique opportunities and challenges for gardeners and landscapers.

Uses of Mosses

Mosses, with their unique properties and characteristics, have been harnessed for various applications throughout history. Here are some of the notable uses of mosses:

  1. Floral and Home Decorations:
    • Mosses, owing to their aesthetic appeal, are frequently employed by florists and interior designers. Their lush green texture and ability to thrive in various conditions make them ideal for enhancing the visual appeal of indoor spaces and floral arrangements.
  2. Medical Applications during World War I:
    • Sphagnum moss, renowned for its exceptional water-absorbing capacity, was utilized as a makeshift bandage during World War I. Its ability to retain moisture and provide a degree of cushioning made it a valuable resource in emergency medical situations on the battlefield.
  3. Historical Fire Safety:
    • In earlier times, mosses were employed as rudimentary fire extinguishers. Their inherent moisture retention capability allowed them to smother and suppress small fires effectively.
  4. Fuel Source:
    • Peat, formed from the accumulated and compacted layers of moss over time, serves as a combustible material. In various regions, especially those with abundant peat bogs, peat has been traditionally extracted and used as a fuel source for heating and cooking.

In essence, mosses, beyond their ecological roles, have found diverse applications in human endeavors, ranging from decorative purposes to practical uses in energy and medicine.

Economic Importance of Mosses

Mosses, though seemingly inconspicuous, have significant economic implications. Their unique properties and adaptability have made them valuable in various sectors. Here’s an overview of the economic importance of mosses:

  1. Horticulture and Gardening:
    • Mosses are used to improve the soil’s water retention capacity, especially in potting mixes.
    • They are also employed as decorative elements in gardens, terrariums, and as ground covers in shaded areas.
  2. Medical Applications:
    • Historically, during World War I, Sphagnum moss was used as a bandage due to its high absorbency, antiseptic properties, and ability to keep wounds clean.
  3. Fuel Source:
    • Peat moss, which is decomposed moss layers, is harvested and used as a fuel in certain regions, especially in parts of Europe. It’s burned to produce heat in residential and industrial settings.
  4. Water Filtration:
    • Due to their capacity to absorb and retain water, mosses have been explored for use in water filtration systems, helping to remove impurities and contaminants from water.
  5. Erosion Control:
    • Mosses play a crucial role in preventing soil erosion. Their dense growth helps bind the soil, especially in areas prone to erosion like riverbanks and hillsides.
  6. Bio-indicators:
    • Mosses are sensitive to environmental changes, especially air pollution. They are used as bio-indicators to monitor air quality and the presence of heavy metals in the environment.
  7. Research and Biotechnology:
    • Mosses are used as model organisms in research due to their simple structure and ease of genetic manipulation. They provide insights into plant evolution and development.
    • In biotechnology, mosses are being explored for the production of complex biopharmaceuticals.
  8. Cultural and Recreational Value:
    • Moss gardens, particularly in countries like Japan, have cultural and aesthetic significance. They offer serene environments for relaxation and meditation.
  9. Insulation:
    • Traditionally, mosses have been used for insulation in buildings, especially in indigenous communities in the Arctic regions. They provide a natural insulating layer against cold temperatures.
  10. Agricultural Uses:
    • Peat moss is used as a soil conditioner, enhancing the soil’s capacity to retain water and nutrients. It’s beneficial for crops that require well-aerated and moist soil conditions.

In summary, mosses, with their multifaceted applications, contribute significantly to the economy, ranging from traditional uses to modern biotechnological applications. Their ecological roles further underscore their importance in maintaining environmental balance and sustainability.

Examples of Mosses

Mosses encompass a diverse range of species, each with its unique characteristics and habitats. Here are some notable examples of mosses:

  1. Sphagnum Moss (Peat Moss):
    • Found in bogs and wetlands, Sphagnum moss is renowned for its water-retention capabilities. It’s also the primary component of peat.
  2. Polytrichum commune (Haircap Moss or Common Hair Moss):
    • Recognizable by its tall, erect growth and hair-like structures on the leaf tips, it’s commonly found in various habitats, from forests to grasslands.
  3. Bryum argenteum (Silver Moss):
    • Often found in urban areas, this moss has a silvery appearance and grows on pavements, walls, and roofs.
  4. Funaria hygrometrica (Bonfire Moss or Water Measuring Moss):
    • Typically found in recently burned areas, this moss is known for its ability to indicate humidity changes.
  5. Tortula muralis (Wall Screw Moss):
    • As the name suggests, this moss is commonly found on walls and can tolerate dry conditions.
  6. Dicranum scoparium (Broom Fork Moss):
    • Recognized by its tufted growth and long, curved leaves, it’s often found in woodlands and forests.
  7. Marchantia polymorpha:
    • Although technically a liverwort, it’s commonly mistaken for moss. It has a flat, thalloid structure and is known for its umbrella-like reproductive structures.
  8. Ceratodon purpureus (Purple Horn Toothed Moss):
    • This moss is adaptable and can be found in various habitats, from urban areas to forests. It’s recognized by its purplish setae.
  9. Atrichum undulatum (Wavy Starburst Moss):
    • Found in woodlands, this moss is known for its wavy leaves and starburst-like appearance.
  10. Mnium hornum (Swan’s Neck Thyme Moss):
    • Commonly found in damp woodlands, it’s recognized by its glossy leaves and elongated setae.

These are just a few examples from the vast world of mosses. Each species has its unique adaptations and ecological roles, contributing to the biodiversity and health of various ecosystems.

Quiz

Which stage is dominant in the life cycle of mosses?
a) Diploid sporophyte
b) Haploid gametophyte
c) Zygote
d) Spore

Mosses are classified under which division?
a) Pteridophyta
b) Bryophyta
c) Gymnospermae
d) Angiospermae

Which of the following is NOT a characteristic of mosses?
a) Vascular tissues
b) Non-woody plants
c) Reproduce by spores
d) Lack true roots

What are the thread-like structures that help mosses attach to the substrate?
a) Stems
b) Rhizomes
c) Rhizoids
d) Roots

Which moss is commonly known as peat moss?
a) Polytrichopsida
b) Bryopsida
c) Sphagnopsida
d) Andreaeopsida

In which habitat would you most likely find mosses growing?
a) Dry deserts
b) Sandy beaches
c) Moist, shaded forests
d) Open fields with direct sunlight

What is the primary function of the capsule in mosses?
a) Photosynthesis
b) Storage of nutrients
c) Protection and dispersal of spores
d) Absorption of water

Mosses are primarily found in which type of environments?
a) Hot and arid
b) Cold and dry
c) Cool, humid, and cloudy
d) Warm and sunny

Which part of the moss plant is responsible for sexual reproduction?
a) Sporophyte
b) Rhizoid
c) Gametophyte
d) Capsule

Which of the following mosses is known to grow on rocks?
a) Sphagnopsida
b) Polytrichopsida
c) Bryopsida
d) Andreaeopsida

FAQ

What are mosses?

Mosses are small, non-vascular plants that belong to the division Bryophyta. They are typically found in damp and shaded locations.

How do mosses reproduce?

Mosses reproduce both sexually, through the fusion of gametes, and asexually, through the dispersal of spores.

Do mosses have roots?

No, mosses do not have true roots. Instead, they have thread-like structures called rhizoids that anchor them to the substrate.

Why are mosses considered non-vascular plants?

Mosses lack specialized tissues like xylem and phloem for the transport of water and nutrients, which is why they are termed non-vascular.

Can mosses grow in direct sunlight?

While some moss species can tolerate direct sunlight, most prefer shaded or partially shaded environments.

What is the significance of mosses in an ecosystem?

Mosses play crucial roles in ecosystems, including soil formation, water retention, and providing habitats for microorganisms and small animals.

How can mosses survive without true roots?

Mosses absorb water and nutrients directly through their leaves. The rhizoids anchor them to the substrate but do not function like true roots in absorption.

Why are mosses often found in moist environments?

Mosses require a moist environment for the movement of sperm cells during sexual reproduction. Their lack of vascular tissues also makes them dependent on external moisture.

Do mosses have flowers and seeds?

No, mosses do not produce flowers or seeds. Instead, they reproduce using spores.

Can mosses be used in gardening?

Yes, mosses are often used in Japanese gardening for aesthetic purposes, in moss lawns, and even in green roofs due to their water retention properties.

References

  1. Hartwell, L. H., Hood, L., Goldberg, M. L., Reynolds, A. E., & Silver, L. M. (2011). Genetics: From Genes to Genomes. Boston: McGraw Hill.
  2. McMahon, M. J., Kofranek, A. M., & Rubatzky, V. E. (2011). Plant Science: Growth, Development, and Utilization of Cultivated Plants (5th ed.). Boston: Prentince Hall.
  3. Rubinstein, C. V., Gerrienne, P., de la Puente, G., Astini, R. A., & Steemans, P. (2010). Early Middle Ordovician evidence for land plants in Argentina (eastern Gondwana). New Phytologist, 188(2).

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