WEEK 7: PLANT BIODIVERSITY

EARLY PLANT LIFE

The kingdom Plantae constitutes large and varied groups of organisms. There are more than 300,000 species of
catalogued plants. Of these, more than 260,000 are seed plants. Mosses, ferns, conifers, and flowering plants are
all members of the plant kingdom. Most biologists also consider green algae to be plants, although others exclude
all algae from the plant kingdom. The reason for this disagreement stems from the fact that only green algae,
the Charophytes, share common characteristics with land plants (such as using chlorophyll a and b plus carotene
in the same proportion as plants). These characteristics are absent in other types of algae.

Evolution Connection
Algae and Evolutionary Paths to Photosynthesis. Some scientists consider all algae to be plants, while others
assert that only the Charophytes belong in the kingdom Plantae. These divergent opinions are related to the
different evolutionary paths to photosynthesis selected for in different types of algae. While all algae are
photosynthetic—that is, they contain some form of a chloroplast—they didn’t all become photosynthetic via the
same path.
The ancestors to the green algae became photosynthetic by endosymbiosing a green, photosynthetic bacterium
about 1.65 billion years ago. That algal line evolved into the Charophytes, and eventually into the modern mosses,
ferns, gymnosperms, and angiosperms. Their evolutionary trajectory was relatively straight and monophyletic. In
contrast, the other algae—red, brown, golden, stramenopiles, and so on—all became photosynthetic by
secondary, or even tertiary, endosymbiotic events; that is, they endosymbiosed cells that had already
endosymbiosed a cyanobacterium. These latecomers to photosynthesis are parallels to the Charophytes in terms
of autotrophy, but they did not expand to the same extent as the Charophytes, nor did they colonize the land.
The different views on whether all algae are Plantae arise from how these evolutionary paths are viewed. Scientists
who solely track evolutionary straight lines (that is, monophyly), consider only the Charophytes as plants. To
biologists who cast a broad net over living things that share a common characteristic (in this case, photosynthetic
eukaryotes), all algae are plants.

Plant Adaptations to Life on Land

As organisms adapted to life on land, they had to contend with several challenges in the terrestrial environment.
Water has been described as “the stuff of life.” The cell’s interior is a watery soup: in this medium, most small
molecules dissolve and diffuse, and the majority of the chemical reactions of metabolism take place. Desiccation,
or drying out, is a constant danger for an organism exposed to air. Even when parts of a plant are close to a source
of water, the aerial structures are likely to dry out. Water also provides buoyancy to organisms. On land, plants
need to develop structural support in a medium that does not give the same lift as water. The organism is also
subject to bombardment by mutagenic radiation, because air does not filter out ultraviolet rays of sunlight.
Additionally, the male gametes must reach the female gametes using new strategies, because swimming is no
longer possible. Therefore, both gametes and zygotes must be protected from desiccation. The successful land
plants developed strategies to deal with all of these challenges. Not all adaptations appeared at once. Some
species never moved very far from the aquatic environment, whereas others went on to conquer the driest
environments on Earth.
To balance these survival challenges, life on land offers several advantages. First, sunlight is abundant. Water acts
as a filter, altering the spectral quality of light absorbed by the photosynthetic pigment chlorophyll. Second,
carbon dioxide is more readily available in air than in water, since it diffuses faster in air. Third, land plants evolved
before land animals; therefore, until dry land was colonized by animals, no predators threatened plant life. This
situation changed as animals emerged from the water and fed on the abundant sources of nutrients in the
established flora. In turn, plants developed strategies to deter predation: from spines and thorns to toxic
chemicals.
Early land plants, like the early land animals, did not live very far from an abundant source of water and developed
survival strategies to combat dryness. One of these strategies is called tolerance. Many mosses, for example, can
dry out to a brown and brittle mat, but as soon as rain or a flood makes water available, mosses will absorb it and
are restored to their healthy green appearance. Another strategy is to colonize environments with high humidity,
where droughts are uncommon. Ferns, which are considered an early lineage of plants, thrive in damp and cool
places such as the understory of temperate forests. Later, plants moved away from moist or aquatic environments
using resistance to desiccation, rather than tolerance. These plants, like cacti, minimize the loss of water to such
an extent they can survive in extremely dry environments.
The most successful adaptation solution was the development of new structures that gave plants the advantage
when colonizing new and dry environments. Four major adaptations are found in all terrestrial plants: the
alternation of generations, a sporangium in which the spores are formed, a gametangium that produces haploid
cells, and apical meristem tissue in roots and shoots. The evolution of a waxy cuticle and a cell wall with lignin also
contributed to the success of land plants. These adaptations are noticeably lacking in the closely related green
algae—another reason for the debate over their placement in the plant kingdom.

• Alternation of Generations
Alternation of generations describes a life cycle in which an organism has both haploid and diploid multicellular
stages (Figure).
Alternation of generations between the 1n gametophyte and 2n sporophyte
is shown.
Haplontic refers to a lifecycle in which there is a dominant haploid
stage, and diplontic refers to a lifecycle in which the diploid is the
dominant life stage. Humans are diplontic. Most plants exhibit
alternation of generations, which is described
as haplodiplodontic: the haploid multicellular form, known as a
gametophyte, is followed in the development sequence by a
multicellular diploid organism: the sporophyte. The gametophyte gives rise to the gametes (reproductive cells) by
mitosis. This can be the most obvious phase of the life cycle of the plant, as in the mosses, or it can occur in a
microscopic structure, such as a pollen grain, in the higher plants (a common collective term for the vascular
plants). The sporophyte stage is barely noticeable in lower plants (the collective term for the plant groups of
mosses, liverworts, and lichens). Towering trees are the diplontic phase in the lifecycles of plants such as sequoias
and pines.
Protection of the embryo is a major requirement for land plants. The vulnerable embryo must be sheltered from
desiccation and other environmental hazards. In both seedless and seed plants, the female gametophyte provides
protection and nutrients to the embryo as it develops into the new generation of sporophyte. This distinguishing
feature of land plants gave the group its alternate name of embryophytes.

• Sporangia in Seedless Plants

The sporophyte of seedless plants is diploid and results from syngamy (fusion) of two gametes. The sporophyte
bears the sporangia (singular, sporangium): organs that first appeared in the land plants. The term “sporangia”
literally means “spore in a vessel,” as it is a reproductive sac that contains spores Figure. Inside the multicellular
sporangia, the diploid sporocytes, or mother cells, produce haploid spores by meiosis, where the
2n chromosome number is reduced to 1n (note that many plant sporophytes are polyploid: for example, durum
wheat is tetraploid, bread wheat is hexaploid, and some ferns are 1000-ploid). The spores are later released by
the sporangia and disperse in the environment. Two different types of spores are produced in land plants,
resulting in the separation of sexes at different points in the lifecycle. Seedless non-vascular plants produce only
one kind of spore and are called homosporous. The gametophyte phase is dominant in these plants. After
germinating from a spore, the resulting gametophyte produces both male and female gametangia, usually on the
same individual. In contrast, heterosporous plants produce two morphologically different types of spores. The
male spores are called microspores, because of their smaller size, and develop into the male gametophyte; the
comparatively larger megaspores develop into the female gametophyte. Heterospory is observed in a
few seedless vascular plants and in all seed plants.
Spore-producing sacs called sporangia grow at the ends of long, thin stalks in this photo of the
moss Esporangios bryum.
When the haploid spore germinates in a hospitable environment, it generates a multicellular
gametophyte by mitosis. The gametophyte supports the zygote formed from the fusion of
gametes and the resulting young sporophyte (vegetative form). The cycle then begins anew.
The spores of seedless plants are surrounded by thick cell walls containing a tough polymer known
as sporopollenin. This complex substance is characterized by long chains of organic molecules related to fatty
acids and carotenoids: hence the yellow color of most pollen. Sporopollenin is unusually resistant to chemical and
biological degradation. In seed plants, which use pollen to transfer the male sperm to the female egg, the
toughness of sporopollenin explains the existence of well-preserved pollen fossils. Sporopollenin was once
thought to be an innovation of land plants; however, the green algae Coleochaetes forms spores that contain
sporopollenin.
 • Gametangia in Seedless Plants
Gametangia (singular, gametangium) are structures observed on multicellular haploid gametophytes. In the
gametangia, precursor cells give rise to gametes by mitosis. The male gametangium (antheridium) releases sperm.
Many seedless plants produce sperm equipped with flagella that enable them to swim in a moist environment to
the archegonia: the female gametangium. The embryo develops inside the archegonium as the sporophyte.
Gametangia are prominent in seedless plants, but are very rarely found in seed plants.

• Apical Meristems
Shoots and roots of plants increase in length through rapid cell division in a tissue called the apical meristem,
which is a small zone of cells found at the shoot tip or root tip (Figure). The apical
meristem is made of undifferentiated cells that continue to proliferate throughout the
life of the plant. Meristematic cells give rise to all the specialized tissues of the organism.
Elongation of the shoots and roots allows a plant to access additional space and
resources: light in the case of the shoot, and water and minerals in the case of roots. A
separate meristem, called the lateral meristem, produces cells that increase the diameter
of tree trunks.
Addition of new cells in a root occurs at the apical meristem. Subsequent enlargement of these cells causes the
organ to grow and elongate. The root cap protects the fragile apical meristem as the root tip is pushed through
the soil by cell elongation.

Additional Land Plant Adaptations

As plants adapted to dry land and became independent from the constant presence of water in damp habitats,
new organs and structures made their appearance. Early land plants did not grow more than a few inches off the
ground, competing for light on these low mats. By developing a shoot and growing taller, individual plants
captured more light. Because air offers substantially less support than water, land plants incorporated more rigid
molecules in their stems (and later, tree trunks). In small plants such as single-celled algae, simple diffusion
suffices to distribute water and nutrients throughout the organism. However, for plants to evolve larger forms,
the evolution of vascular tissue for the distribution of water and solutes was a prerequisite. The vascular system
contains xylem and phloem tissues. Xylem conducts water and minerals absorbed from the soil up to the shoot,
while phloem transports food derived from photosynthesis throughout the entire plant. A root system evolved to
take up water and minerals from the soil, and to anchor the increasingly taller shoot in the soil.
In land plants, a waxy, waterproof cover called a cuticle protects the leaves and stems from desiccation. However,
the cuticle also prevents intake of carbon dioxide needed for the synthesis of carbohydrates through
photosynthesis. To overcome this, stomata or pores that open and close to regulate traffic of gases and water
vapor appeared in plants as they moved away from moist environments into drier habitats.
Water filters ultraviolet-B (UVB) light, which is harmful to all organisms, especially those that must absorb light to
survive. This filtering does not occur for land plants. This presented an additional challenge to land colonization,
which was met by the evolution of biosynthetic pathways for the synthesis of protective flavonoids and other
compounds: pigments that absorb UV wavelengths of light and protect the aerial parts of plants from
photodynamic damage.
Plants cannot avoid being eaten by animals. Instead, they synthesize a large range of poisonous secondary
metabolites: complex organic molecules such as alkaloids, whose noxious smells and unpleasant taste deter
animals. These toxic compounds can also cause severe diseases and even death, thus discouraging predation.
Humans have used many of these compounds for centuries as drugs, medications, or spices. In contrast, as plants
co-evolved with animals, the development of sweet and nutritious metabolites lured animals into providing
valuable assistance in dispersing pollen grains, fruit, or seeds. Plants have been enlisting animals to be their
helpers in this way for hundreds of millions of years.

Evolution of Land Plants

No discussion of the evolution of plants on land can be undertaken without a brief review of the timeline of the
geological eras. The early era, known as the Paleozoic, is divided into six periods. It starts with the Cambrian
period, followed by the Ordovician, Silurian, Devonian, Carboniferous, and Permian. The major event to mark the
Ordovician, more than 500 million years ago, was the colonization of land by the ancestors of modern land plants.
Fossilized cells, cuticles, and spores of early land plants have been dated as far back as the Ordovician period in
the early Paleozoic era. The oldest-known vascular plants have been identified in deposits from the Devonian.
One of the richest sources of information is the Rhynie chert, a sedimentary rock deposit found in Rhynie, Scotland
(Figure), where embedded fossils of some of the earliest vascular plants have been identified.
 This Rhynie chert contains fossilized material from vascular plants. The area inside
the circle contains bulbous underground stems called corms, and root-like
structures called rhizoids.
Paleobotanists distinguish between extinct species, as fossils,
and extant species, which are still living. The extinct vascular plants, classified
as zosterophylls and trimerophytes, most probably lacked true leaves and
roots and formed low vegetation mats similar in size to modern-day mosses,
although some trimetophytes could reach one meter in height. The later
genus Cooksonia, which flourished during the Silurian, has been extensively studied from well-preserved
examples. Imprints of Cooksonia show slender branching stems ending in what appear to be sporangia. From the
recovered specimens, it is not possible to establish for certain whether Cooksonia possessed vascular tissues.
Fossils indicate that by the end of the Devonian period, ferns, horsetails, and seed plants populated the landscape,
giving rising to trees and forests. This luxuriant vegetation helped enrich the atmosphere in oxygen, making it
easier for air-breathing animals to colonize dry land. Plants also established early symbiotic relationships with
fungi, creating mycorrhizae: a relationship in which the fungal network of filaments increases the efficiency of the
plant root system, and the plants provide the fungi with byproducts of photosynthesis.

Paleobotanist How organisms acquired traits that allow them to colonize new environments—and how the
contemporary ecosystem is shaped—are fundamental questions of evolution. Paleobotany (the study of extinct
plants) addresses these questions through the analysis of fossilized specimens retrieved from field studies,
reconstituting the morphology of organisms that disappeared long ago. Paleobotanists trace the evolution of
plants by following the modifications in plant morphology: shedding light on the connection between existing
plants by identifying common ancestors that display the same traits. This field seeks to find transitional species
that bridge gaps in the path to the development of modern organisms. Fossils are formed when organisms are
trapped in sediments or environments where their shapes are preserved. Paleobotanists collect fossil specimens
in the field and place them in the context of the geological sediments and other fossilized organisms surrounding
them. The activity requires great care to preserve the integrity of the delicate fossils and the layers of rock in which
they are found.
One of the most exciting recent developments in paleobotany is the use of analytical chemistry and molecular
biology to study fossils. Preservation of molecular structures requires an environment free of oxygen, since
oxidation and degradation of material through the activity of microorganisms depend on its presence. One
example of the use of analytical chemistry and molecular biology is the identification of oleanane, a compound
that deters pests. Up to this point, oleanane appeared to be unique to flowering plants; however, it has now been
recovered from sediments dating from the Permian, much earlier than the current dates given for the appearance
of the first flowering plants. Paleobotanists can also study fossil DNA, which can yield a large amount of
information, by analyzing and comparing the DNA sequences of extinct plants with those of living and related
organisms. Through this analysis, evolutionary relationships can be built for plant lineages.
Some paleobotanists are skeptical of the conclusions drawn from the analysis of molecular fossils. For example,
the chemical materials of interest degrade rapidly when exposed to air during their initial isolation, as well as in
further manipulations. There is always a high risk of contaminating the specimens with extraneous material,
mostly from microorganisms. Nevertheless, as technology is refined, the analysis of DNA from fossilized plants
will provide invaluable information on the evolution of plants and their adaptation to an ever-changing
environment.

The Major Divisions of Land Plants

The green algae and land plants are grouped together into a subphylum called the Streptophytina, and thus are
called Streptophytes. In a further division, land plants are classified into two major groups according to the
absence or presence of vascular tissue, as detailed in Figure. Plants that lack vascular tissue, which is formed of
specialized cells for the transport of water and nutrients, are referred to as non-vascular plants. Liverworts,
mosses, and hornworts are seedless, non-vascular plants that likely appeared early in land plant evolution.
Vascular plants developed a network of cells that conduct water and solutes. The first vascular plants appeared
in the late Ordovician and were probably similar to lycophytes, which include club mosses (not to be confused
with the mosses) and the pterophytes (ferns, horsetails, and whisk ferns). Lycophytes and pterophytes are
referred to as seedless vascular plants, because they do not produce seeds. The seed plants, or spermatophytes,
form the largest group of all existing plants, and hence dominate the landscape. Seed plants include
gymnosperms, most notably conifers (Gymnosperms), which produce “naked seeds,” and the most successful of
all plants, the flowering plants (Angiosperms). Angiosperms protect their seeds inside chambers at the center of
a flower; the walls of the chamber later develop into a fruit.
This table shows the major divisions of green plants.
Which of the following statements about plant divisions is false?
1. Lycophytes and pterophytes are seedless vascular plants.
2. All vascular plants produce seeds.
3. All nonvascular embryophytes are
bryophytes.
4. Seed plants include angiosperms and
gymnosperms.

Section Summary
Land plants acquired traits that made it possible to
colonize land and survive out of the water. All land
plants share the following characteristics:
alternation of generations, with the haploid plant
called a gametophyte, and the diploid plant called a
sporophyte; protection of the embryo, formation of haploid spores in a sporangium, formation of gametes in a
gametangium, and an apical meristem. Vascular tissues, roots, leaves, cuticle cover, and a tough outer layer that
protects the spores contributed to the adaptation of plants to dry land. Land plants appeared about 500 million
years ago in the Ordovician period.

BRYOPHYTES
Bryophytes are the group of plants that are the closest extant relative of early terrestrial plants. The first
bryophytes (liverworts) most likely appeared in the Ordovician period, about 450 million years ago. Because of
the lack of lignin and other resistant structures, the likelihood of bryophytes forming fossils is rather small. Some
spores protected by sporopollenin have survived and are attributed to early bryophytes. By the Silurian period,
however, vascular plants had spread through the continents. This compelling fact is used as evidence that non-
vascular plants must have preceded the Silurian period.

More than 25,000 species of bryophytes thrive in mostly damp habitats, although some live in deserts. They
constitute the major flora of inhospitable environments like the tundra, where their small size and tolerance to
desiccation offer distinct advantages. They generally lack lignin and do not have actual tracheids (xylem cells
specialized for water conduction). Rather, water and nutrients circulate inside specialized conducting cells.
Although the term non-tracheophyte is more accurate, bryophytes are commonly called nonvascular plants.

In a bryophyte, all the conspicuous vegetative organs—including the photosynthetic leaf-like structures, the
thallus, stem, and the rhizoid that anchors the plant to its substrate—belong to the haploid organism or
gametophyte. The sporophyte is barely noticeable. The gametes formed by bryophytes swim with a flagellum, as
do gametes in a few of the tracheophytes. The sporangium—the multicellular sexual reproductive structure—is
present in bryophytes and absent in the majority of algae. The bryophyte embryo also remains attached to the
parent plant, which protects and nourishes it. This is a characteristic of land plants.

The bryophytes are divided into three phyla: the liverworts or Hepaticophyta, the hornworts or Anthocerotophyta,
and the mosses or true Bryophyta.
• Liverworts
Liverworts (Hepaticophyta) are viewed as the plants most closely related to the
ancestor that moved to land. Liverworts have colonized every terrestrial habitat on Earth
and diversified to more than 7000 existing species (Figure). Some gametophytes form
lobate green structures, as seen in Figure. The shape is similar to the lobes of the liver,
and hence provides the origin of the name given to the phylum. This 1904 drawing shows
the variety of forms of Hepaticophyta. A liverwort, Lunularia cruciata, displays its lobate, flat
thallus. The organism in the photograph is in the gametophyte stage.
 Openings that allow the movement of gases may be observed in liverworts. However,
these are not stomata, because they do not actively open and close. The plant takes
up water over its entire surface and has no cuticle to prevent
desiccation. Figure represents the lifecycle of a liverwort. The cycle starts with the
release of haploid spores from the sporangium that developed on the sporophyte.
Spores disseminated by wind or
water germinate into flattened
thalli attached to the substrate by thin, single-celled filaments.
Male and female gametangia develop on separate, individual
plants. Once released, male gametes swim with the aid of their
flagella to the female gametangium (the archegonium), and
fertilization ensues. The zygote grows into a small sporophyte
still attached to the parent gametophyte. It will give rise, by
meiosis, to the next generation of spores. Liverwort plants can
also reproduce asexually, by the breaking of branches or the
spreading of leaf fragments called gemmae. In this latter type
of reproduction, the gemmae—small, intact, complete pieces
of plant that are produced in a cup on the surface of the thallus
(shown in Figure)—are splashed out of the cup by raindrops.
The gemmae then land nearby and develop into gametophytes.
The life cycle of a typical liverwort is shown.

• Hornworts
The hornworts (Anthocerotophyta) belong to the broad
bryophyte group. They have colonized a variety of habitats on land, although they are never far from a source of
moisture. The short, blue-green gametophyte is the dominant phase of the lifecycle of a hornwort. The narrow,
pipe-like sporophyte is the defining characteristic of the group. The sporophytes emerge from the parent
gametophyte and continue to grow throughout the life of the plant (Figure).
Hornworts grow a tall and slender sporophyte.

Stomata appear in the hornworts and are abundant on the sporophyte.

Photosynthetic cells in the thallus contain a single chloroplast. Meristem cells at the
base of the plant keep dividing and adding to its height. Many hornworts establish
symbiotic relationships with cyanobacteria that fix nitrogen from the environment.

The lifecycle of hornworts (Figure) follows the general pattern of alternation of generations. The gametophytes
grow as flat thalli on the soil with embedded gametangia. Flagellated sperm swim to the archegonia and fertilize
eggs. The zygote develops into a long and slender sporophyte that eventually splits open, releasing spores. Thin
cells called pseudoelaters surround the spores and help propel them further in the environment. Unlike the
elaters observed in horsetails, the hornwort pseudoelaters are single-celled structures. The haploid spores
germinate and give rise to the next generation of gametophyte.
The alternation of generation in hornworts is shown.

• Mosses

More than 10,000 species of mosses have been catalogued.

Their habitats vary from the tundra, where they are the main
vegetation, to the understory of tropical forests. In the tundra,
the mosses’ shallow rhizoids allow them to fasten to a substrate
without penetrating the frozen soil. Mosses slow down erosion,
store moisture and soil nutrients, and provide shelter for small
animals as well as food for larger herbivores, such as the musk
ox. Mosses are very sensitive to air pollution and are used to
monitor air quality. They are also sensitive to copper salts, so
these salts are a common ingredient of compounds marketed to
eliminate mosses from lawns.
Mosses form diminutive gametophytes, which are the dominant phase of the lifecycle. Green, flat structures—
resembling true leaves, but lacking vascular tissue—are attached in a spiral to a central stalk. The plants absorb
water and nutrients directly through these leaf-like structures. Some mosses have small branches. Some primitive
traits of green algae, such as flagellated sperm, are still present in mosses that are dependent on water for
reproduction. Other features of mosses are clearly adaptations to dry land. For example, stomata are present on
the stems of the sporophyte, and a primitive vascular system runs up the sporophyte’s stalk. Additionally, mosses
are anchored to the substrate—whether it is soil, rock, or roof tiles—by multicellular rhizoids. These structures
are precursors of roots. They originate from the base of the gametophyte, but are not the major route for the
absorption of water and minerals. The lack of a true root system explains why it is so easy to rip moss mats from
a tree trunk. The moss lifecycle follows the pattern of alternation of generations as shown in Figure. The most
familiar structure is the haploid gametophyte, which germinates from a haploid spore and forms first
a protonema—usually, a tangle of single-celled filaments that hug the ground. Cells akin to an apical meristem
actively divide and give rise to a gametophore, consisting of a photosynthetic stem and foliage-like structures.
Rhizoids form at the base of the gametophore. Gametangia of both sexes develop on separate gametophores.
The male organ (the antheridium) produces many sperm, whereas the archegonium (the female organ) forms a
single egg. At fertilization, the sperm swims down the neck to the venter and unites with the egg inside the
archegonium. The zygote, protected by the archegonium, divides and grows into a sporophyte, still attached by
its foot to the gametophyte.
This illustration shows the life cycle of mosses.

The slender seta (plural, setae), as seen in Figure, contains

tubular cells that transfer nutrients from the base of the
sporophyte (the foot) to the sporangium or capsule.

This photograph shows the

long slender stems, called
setae, connected to capsules
of the moss Thamnobryum
alopecurum.

A structure called
a peristome increases the spread of spores after the tip of
the capsule falls off at dispersal. The concentric tissue
around the mouth of the capsule is made of triangular, close-
fitting units, a little like “teeth”; these open and close
depending on moisture levels, and periodically release
spores.

Section Summary

Seedless nonvascular plants are small, having the

gametophyte as the dominant stage of the lifecycle. Without
a vascular system and roots, they absorb water and
nutrients on all their exposed surfaces. Collectively known as
bryophytes, the three main groups include the liverworts,
the hornworts, and the mosses. Liverworts are the most
primitive plants and are closely related to the first land plants. Hornworts developed stomata and possess a single
chloroplast per cell. Mosses have simple conductive cells and are attached to the substrate by rhizoids. They
colonize harsh habitats and can regain moisture after drying out. The moss sporangium is a complex structure
that allows release of spores away from the parent plant.

SEEDLESS VASCULAR PLANTS

The vascular plants, or tracheophytes, are the dominant and most conspicuous group of land plants. More than
260,000 species of tracheophytes represent more than 90 percent of Earth’s vegetation. Several evolutionary
innovations explain their success and their ability to spread to all habitats.
Bryophytes may have been successful at the transition from an aquatic habitat to land, but they are still dependent
on water for reproduction, and absorb moisture and nutrients through the gametophyte surface. The lack of roots
for absorbing water and minerals from the soil, as well as a lack of reinforced conducting cells, limits bryophytes
to small sizes. Although they may survive in reasonably dry conditions, they cannot reproduce and expand their
habitat range in the absence of water. Vascular plants, on the other hand, can achieve enormous heights, thus
competing successfully for light. Photosynthetic organs become leaves, and pipe-like cells or vascular tissues
transport water, minerals, and fixed carbon throughout the organism.

In seedless vascular plants, the diploid sporophyte is the dominant phase of the lifecycle. The gametophyte is now
an inconspicuous, but still independent, organism. Throughout plant evolution, there is an evident reversal of
roles in the dominant phase of the lifecycle. Seedless vascular plants still depend on water during fertilization, as
the sperm must swim on a layer of moisture to reach the egg. This step in reproduction explains why ferns and
their relatives are more abundant in damp environments.
Vascular Tissue: Xylem and Phloem

The first fossils that show the presence of vascular tissue date to the Silurian period, about 430 million years ago.
The simplest arrangement of conductive cells shows a pattern of xylem at the center surrounded by
phloem. Xylem is the tissue responsible for the storage and long-distance transport of water and nutrients, as
well as the transfer of water-soluble growth factors from the organs of synthesis to the target organs. The tissue
consists of conducting cells, known as tracheids, and supportive filler tissue, called parenchyma. Xylem conductive
cells incorporate the compound lignin into their walls, and are thus described as lignified. Lignin itself is a complex
polymer that is impermeable to water and confers mechanical strength to vascular tissue. With their rigid cell
walls, the xylem cells provide support to the plant and allow it to achieve impressive heights. Tall plants have a
selective advantage by being able to reach unfiltered sunlight and disperse their spores or seeds further away,
thus expanding their range. By growing higher than other plants, tall trees cast their shadow on shorter plants
and limit competition for water and precious nutrients in the soil.
Phloem is the second type of vascular tissue; it transports sugars, proteins, and other solutes throughout the
plant. Phloem cells are divided into sieve elements (conducting cells) and cells that support the sieve elements.
Together, xylem and phloem tissues form the vascular system of plants.
Roots: Support for the Plant

Roots are not well preserved in the fossil record. Nevertheless, it seems that roots appeared later in evolution
than vascular tissue. The development of an extensive network of roots represented a significant new feature of
vascular plants. Thin rhizoids attached bryophytes to the substrate, but these rather flimsy filaments did not
provide a strong anchor for the plant; neither did they absorb substantial amounts of water and nutrients. In
contrast, roots, with their prominent vascular tissue system, transfer water and minerals from the soil to the rest
of the plant. The extensive network of roots that penetrates deep into the soil to reach sources of water also
stabilizes trees by acting as a ballast or anchor. The majority of roots establish a symbiotic relationship with fungi,
forming mycorrhizae, which benefit the plant by greatly increasing the surface area for absorption of water and
soil minerals and nutrients.
Leaves, Sporophylls, and Strobili

A third innovation marks the seedless vascular plants. Accompanying the prominence of the sporophyte and the
development of vascular tissue, the appearance of true leaves improved their photosynthetic efficiency. Leaves
capture more sunlight with their increased surface area by employing more chloroplasts to trap light energy and
convert it to chemical energy, which is then used to fix atmospheric carbon dioxide into carbohydrates. The
carbohydrates are exported to the rest of the plant by the conductive cells of phloem tissue.

The existence of two types of morphology suggests that leaves evolved independently in several groups of plants.
The first type of leaf is the microphyll, or “little leaf,” which can be dated to 350 million years ago in the late Silurian.
A microphyll is small and has a simple vascular system. A single unbranched vein—a bundle of vascular tissue
made of xylem and phloem—runs through the center of the leaf. Microphylls may have originated from the
flattening of lateral branches, or from sporangia that lost their reproductive capabilities. Microphylls are present
in the club mosses and probably preceded the development of megaphylls, or “big leaves”, which are larger
leaves with a pattern of branching veins. Megaphylls most likely appeared independently several times during the
course of evolution. Their complex networks of veins suggest that several branches may have combined into a
flattened organ, with the gaps between the branches being filled with photosynthetic tissue.
In addition to photosynthesis, leaves play another role in the life of the plants. Pine cones, mature fronds of ferns,
and flowers are all sporophylls—leaves that were modified structurally to bear sporangia. Strobili are cone-like
structures that contain sporangia. They are prominent in conifers and are commonly known as pine cones.
Ferns and Other Seedless Vascular Plants
By the late Devonian period, plants had evolved vascular tissue, well-defined leaves, and root systems. With these
advantages, plants increased in height and size. During the Carboniferous period, swamp forests of club mosses
and horsetails—some specimens reaching heights of more than 30 m (100 ft)—covered most of the land. These
forests gave rise to the extensive coal deposits that gave the Carboniferous its name. In seedless vascular plants,
the sporophyte became the dominant phase of the lifecycle.

Water is still required for fertilization of seedless vascular plants, and most favor a moist environment. Modern-
day seedless tracheophytes include club mosses, horsetails, ferns, and whisk ferns.

• Phylum Lycopodiophyta: Club Mosses

The club mosses, or phylum Lycopodiophyta, are the earliest group of
seedless vascular plants. They dominated the landscape of the Carboniferous,
growing into tall trees and forming large swamp forests. Today’s club mosses
are diminutive, evergreen plants consisting of a stem (which may be branched)
and microphylls (Figure). The phylum Lycopodiophyta consists of close to 1,200
species, including the quillworts (Isoetales), the club mosses (Lycopodiales), and
spike mosses (Selaginellales), none of which are true mosses or bryophytes.
Lycophytes follow the pattern of alternation of generations seen in the
bryophytes, except that the sporophyte is the major stage of the lifecycle. The
gametophytes do not depend on the sporophyte for nutrients. Some gametophytes develop underground and
form mycorrhizal associations with fungi. In club mosses, the sporophyte gives rise to sporophylls arranged in
strobili, cone-like structures that give the class its name. Lycophytes can be homosporous or heterosporous.
In the club mosses such as Lycopodium clavatum, sporangia are arranged in clusters called strobili.

• Phylum Monilophyta: Class Equisetopsida (Horsetails)

Horsetails, whisk ferns and ferns belong to the phylum Monilophyta,
with horsetails placed in the Class Equisetopsida. The single genus Equisetum is
the survivor of a large group of plants, known as Arthrophyta, which produced
large trees and entire swamp forests in the Carboniferous. The plants are
usually found in damp environments and marshes (Figure).
Horsetails thrive in a marsh.

The stem of a horsetail is characterized by the

presence of joints or nodes, hence the name
Arthrophyta (arthro- = "joint"; -phyta = "plant"). Leaves and branches come out as
whorls from the evenly spaced joints. The needle-shaped leaves do not contribute
greatly to photosynthesis, the majority of which takes place in the green stem (Figure).
Thin leaves originating at the joints are noticeable on the horsetail plant. Horsetails were
once used as scrubbing brushes and were nicknamed scouring brushes.

Silica collects in the epidermal cells, contributing to the stiffness of horsetail plants.
Underground stems known as rhizomes anchor the plants to the ground. Modern-day
horsetails are homosporous and produce bisexual gametophytes.

• Phylum Monilophyta: Class Psilotopsida (Whisk Ferns)

While most ferns form large leaves and branching roots, the whisk ferns,
Class Psilotopsida, lack both roots and leaves, probably lost by reduction.
Photosynthesis takes place in their green stems, and small yellow knobs
form at the tip of the branch stem and contain the sporangia. Whisk ferns
were considered an early pterophytes. However, recent comparative DNA
analysis suggests that this group may have lost both vascular tissue and
roots through evolution, and is more closely related to ferns.
The whisk fern Psilotum nudum has conspicuous green stems with knob-
shaped sporangia.
 • Phylum Monilophyta: Class Psilotopsida (Ferns)
With their large fronds, ferns are the most readily recognizable seedless vascular
plants. They are considered the most advanced seedless vascular plants and
display characteristics commonly observed in seed plants. More than 20,000
species of ferns live in environments ranging from tropics to temperate forests.
Although some species survive in dry environments, most ferns are restricted to
moist, shaded places. Ferns made their appearance in the fossil record during
the Devonian period and expanded during the Carboniferous.
The dominant stage of the lifecycle of a fern is the sporophyte, which consists of
large compound leaves called fronds. Fronds fulfill a double role; they are photosynthetic organs that also carry
reproductive organs. The stem may be buried underground as a rhizome, from which adventitious roots grow to
absorb water and nutrients from the soil; or, they may grow above ground as a trunk in tree ferns
(Figure). Adventitious organs are those that grow in unusual places, such as roots growing from the side of a
stem. Some specimens of this short tree-fern species can grow very tall.
The tip of a developing fern frond is rolled into a crozier, or fiddlehead (Figure
a and Figure b). Fiddleheads unroll as the frond develops. Croziers, or fiddleheads, are the
tips of fern fronds.
The lifecycle of a fern is depicted in Figure.
This life cycle of a fern shows alternation of generations with a
dominant sporophyte stage.
Most ferns produce the same type of spores and are therefore
homosporous. The diploid sporophyte is the most
conspicuous stage of the lifecycle. On the underside of its
mature fronds, sori (singular, sorus) form as small clusters
where sporangia develop (Figure).
Sori appear as small bumps on the
underside of a fern frond.

Inside the sori, spores are produced by meiosis and released into the air. Those that land
on a suitable substrate germinate and form a heart-shaped gametophyte, which is attached to the ground by thin
filamentous rhizoids (Figure).
Shown here are a young sporophyte (upper part of image) and a heart-shaped
gametophyte (bottom part of image).

The inconspicuous gametophyte harbors both sex gametangia. Flagellated sperm

released from the antheridium swim on a wet surface to the archegonium, where
the egg is fertilized. The newly formed zygote grows into a sporophyte that emerges
from the gametophyte and grows by mitosis into the next generation sporophyte.

Landscape Designer. Looking at the well-laid parterres of flowers and fountains in the grounds of royal castles
and historic houses of Europe, it’s clear that the gardens’ creators knew about more than art and design. They
were also familiar with the biology of the plants they chose. Landscape design also has strong roots in the United
States’ tradition. A prime example of early American classical design is Monticello: Thomas Jefferson’s private
estate. Among his many interests, Jefferson maintained a strong passion for botany. Landscape layout can
encompass a small private space, like a backyard garden; public gathering places, like Central Park in New York
City; or an entire city plan, like Pierre L’Enfant’s design for Washington, DC.
A landscape designer will plan traditional public spaces—such as botanical gardens, parks, college campuses,
gardens, and larger developments—as well as natural areas and private gardens. The restoration of natural places
encroached on by human intervention, such as wetlands, also requires the expertise of a landscape designer.

With such an array of necessary skills, a landscape designer’s education includes a solid background in botany,
soil science, plant pathology, entomology, and horticulture. Coursework in architecture and design software is
also required for the completion of the degree. The successful design of a landscape rests on an extensive
knowledge of plant growth requirements, such as light and shade, moisture levels, compatibility of different
species, and susceptibility to pathogens and pests. Mosses and ferns will thrive in a shaded area, where fountains
provide moisture; cacti, on the other hand, would not fare well in that environment. The future growth of
individual plants must be taken into account, to avoid crowding and competition for light and nutrients. The
appearance of the space over time is also of concern. Shapes, colors, and biology
must be balanced for a well-maintained and sustainable green space. Art,
architecture, and biology blend in a beautifully designed and implemented
landscape.
This landscaped border at a college campus was designed by students in the horticulture
and landscaping department of the college.

The Importance of Seedless Vascular Plants

Mosses and liverworts are often the first macroscopic organisms to colonize an area, both in a primary
succession—where bare land is settled for the first time by living organisms—or in a secondary succession, where
soil remains intact after a catastrophic event wipes out many existing species. Their spores are carried by the
wind, birds, or insects. Once mosses and liverworts are established, they provide food and shelter for other
species. In a hostile environment, like the tundra where the soil is frozen, bryophytes grow well because they do
not have roots and can dry and rehydrate rapidly once water is again available. Mosses are at the base of the food
chain in the tundra biome. Many species—from small insects to musk oxen and reindeer—depend on mosses for
food. In turn, predators feed on the herbivores, which are the primary consumers. Some reports indicate that
bryophytes make the soil more amenable to colonization by other plants. Because they establish symbiotic
relationships with nitrogen-fixing cyanobacteria, mosses replenish the soil with nitrogen.

At the end of the nineteenth century, scientists observed that lichens and mosses were becoming increasingly
rare in urban and suburban areas. Since bryophytes have neither a root system for absorption of water and
nutrients, nor a cuticle layer that protects them from desiccation, pollutants in rainwater readily penetrate their
tissues; they absorb moisture and nutrients through their entire exposed surfaces. Therefore, pollutants dissolved
in rainwater penetrate plant tissues readily and have a larger impact on mosses than on other plants. The
disappearance of mosses can be considered a bioindicator for the level of pollution in the environment.

Ferns contribute to the environment by promoting the weathering of rock, accelerating the formation of topsoil,
and slowing down erosion by spreading rhizomes in the soil. The water ferns of the genus Azolla harbor nitrogen-
fixing cyanobacteria and restore this important nutrient to aquatic habitats.
Seedless plants have historically played a role in human life through uses as tools, fuel, and medicine. Dried peat
moss, Sphagnum, is commonly used as fuel in some parts of Europe and is considered a renewable
resource. Sphagnum bogs (Figure) are cultivated with cranberry and blueberry bushes. The ability of Sphagnum to
hold moisture makes the moss a common soil conditioner. Florists use blocks of Sphagnum to maintain moisture
for floral arrangements.
Sphagnum acutifolium is dried peat moss and can be used as fuel. (credit: Ken Goulding)

The attractive fronds of ferns make them a favorite ornamental plant. Because they
thrive in low light, they are well suited as house plants. More importantly, fiddleheads
are a traditional spring food of Native Americans in the Pacific Northwest, and are
popular as a side dish in French cuisine. The licorice fern, Polypodium glycyrrhiza, is part
of the diet of the Pacific Northwest coastal tribes, owing in part to the sweetness of its
rhizomes. It has a faint licorice taste and serves as a sweetener. The rhizome also figures in the pharmacopeia of
Native Americans for its medicinal properties and is used as a remedy for sore throat.
By far the greatest impact of seedless vascular plants on human life, however, comes from their extinct
progenitors. The tall club mosses, horsetails, and tree-like ferns that flourished in the swampy forests of the
Carboniferous period gave rise to large deposits of coal throughout the world. Coal provided an abundant source
of energy during the Industrial Revolution, which had tremendous consequences on human societies, including
rapid technological progress and growth of large cities, as well as the degradation of the environment. Coal is still
a prime source of energy and also a major contributor to global warming.

Section Summary
Vascular systems consist of xylem tissue, which transports water and minerals, and phloem tissue, which
transports sugars and proteins. With the development of the vascular system, there appeared leaves to act as
large photosynthetic organs, and roots to access water from the ground. Small uncomplicated leaves are
microphylls. Large leaves with vein patterns are megaphylls. Modified leaves that bear sporangia are sporophylls.
Some sporophylls are arranged in cone structures called strobili.
The seedless vascular plants include club mosses, which are the most primitive; whisk ferns, which lost leaves and
roots by reductive evolution; and horsetails and ferns. Ferns are the most advanced group of seedless vascular
plants. They are distinguished by large leaves called fronds and small sporangia-containing structures called sori,
which are found on the underside of the fronds.

Mosses play an essential role in the balance of the ecosystems; they are pioneering species that colonize bare or
devastated environments and make it possible for a succession to occur. They contribute to the enrichment of
the soil and provide shelter and nutrients for animals in hostile environments. Mosses and ferns can be used as
fuels and serve culinary, medical, and decorative purposes.

EVOLUTION OF SEED PLANTS

The first plants to colonize land were most likely closely related to modern day mosses (bryophytes) and are
thought to have appeared about 500 million years ago. They were followed by liverworts (also bryophytes) and
primitive vascular plants—the pterophytes—from which modern ferns are derived. The lifecycle of bryophytes
and pterophytes is characterized by the alternation of generations, like gymnosperms and angiosperms; what
sets bryophytes and pterophytes apart from gymnosperms and angiosperms is their reproductive requirement
for water. The completion of the bryophyte and pterophyte life cycle requires water because the male
gametophyte releases sperm, which must swim—propelled by their flagella—to reach and fertilize the female
gamete or egg. After fertilization, the zygote matures and grows into a sporophyte, which in turn will form
sporangia or "spore vessels." In the sporangia, mother cells undergo meiosis and produce the haploid spores.
Release of spores in a suitable environment will lead to germination and a new generation of gametophytes.

In seed plants, the evolutionary trend led to a dominant sporophyte generation, and at the same time, a
systematic reduction in the size of the gametophyte: from a conspicuous structure to a microscopic cluster of cells
enclosed in the tissues of the sporophyte. Whereas lower vascular plants, such as club mosses and ferns, are
mostly homosporous (produce only one type of spore), all seed plants, or spermatophytes, are heterosporous.
They form two types of spores: megaspores (female) and microspores (male). Megaspores develop into female
gametophytes that produce eggs, and microspores mature into male gametophytes that generate sperm.
Because the gametophytes mature within the spores, they are not free-living, as are the gametophytes of other
seedless vascular plants. Heterosporous seedless plants are seen as the evolutionary forerunners of seed plants.
Seeds and pollen—two critical adaptations to drought, and to reproduction that doesn’t require water—
distinguish seed plants from other (seedless) vascular plants. Both adaptations were required for the colonization
of land begun by the bryophytes and their ancestors. Fossils place the earliest distinct seed plants at about 350
million years ago. The first reliable record of gymnosperms dates their
appearance to the Pennsylvanian period, about 319 million years ago
(Figure). Gymnosperms were preceded by progymnosperms, the first
naked seed plants, which arose about 380 million years ago.
Progymnosperms were a transitional group of plants that superficially
resembled conifers (cone bearers) because they produced wood from
the secondary growth of the vascular tissues; however, they still
reproduced like ferns, releasing spores into the environment.
Gymnosperms dominated the landscape in the early (Triassic) and
middle (Jurassic) Mesozoic era. Angiosperms surpassed gymnosperms
by the middle of the Cretaceous (about 100 million years ago) in the late
Mesozoic era, and today are the most abundant plant group in most
terrestrial biomes.
Various plant species evolved in different eras.

Pollen and seed were innovative structures that allowed seed plants to break their dependence on water for
reproduction and development of the embryo, and to conquer dry land. The pollen grains are the male
gametophytes, which contain the sperm (gametes) of the plant. The small haploid (1n) cells are encased in a
protective coat that prevents desiccation (drying out) and mechanical damage. Pollen grains can travel far from
their original sporophyte, spreading the plant’s genes. The seed offers the embryo protection, nourishment, and
a mechanism to maintain dormancy for tens or even thousands of years, ensuring germination can occur when
growth conditions are optimal. Seeds therefore allow plants to disperse the next generation through both space
and time. With such evolutionary advantages, seed plants have become the most successful and familiar group
of plants, in part because of their size and striking appearance.
Evolution of Gymnosperms

The fossil plant Elkinsia polymorpha, a "seed fern" from the Devonian period—about 400 million years ago—is
considered the earliest seed plant known to date. Seed ferns (Figure) produced their seeds along their branches
without specialized structures. What makes them the first true seed plants is that they developed structures called
cupules to enclose and protect the ovule—the female gametophyte and associated tissues—which develops into
a seed upon fertilization. Seed plants resembling modern tree ferns became more numerous and diverse in the
coal swamps of the Carboniferous period.
This fossilized leaf is from Glossopteris, a seed fern that thrived during the
Permian age (290–240 million years ago).

Fossil records indicate the first gymnosperms (progymnosperms) most

likely originated in the Paleozoic era, during the middle Devonian
period: about 390 million years ago. Following the wet Mississippian and
Pennsylvanian periods, which were dominated by giant fern trees, the
Permian period was dry. This gave a reproductive edge to seed plants,
which are better adapted to survive dry spells. The Ginkgoales, a group
of gymnosperms with only one surviving species—the Gingko biloba—
were the first gymnosperms to appear during the lower Jurassic. Gymnosperms expanded in the Mesozoic era
(about 240 million years ago), supplanting ferns in the landscape, and reaching their greatest diversity during this
time. The Jurassic period was as much the age of the cycads (palm-tree-like
gymnosperms) as the age of the dinosaurs. Gingkoales and the more familiar
conifers also dotted the landscape. Although angiosperms (flowering plants)
are the major form of plant life in most biomes, gymnosperms still dominate
some ecosystems, such as the taiga (boreal forests) and the alpine forests at
higher mountain elevations (Figure) because of their adaptation to cold and dry
growth conditions.
This boreal forest (taiga) has low-lying plants and conifer trees.

• Seeds and Pollen as an Evolutionary Adaptation to Dry Land

Unlike bryophyte and fern spores (which are haploid cells dependent on moisture for rapid development of
gametophytes), seeds contain a diploid embryo that will germinate into a sporophyte. Storage tissue to sustain
growth and a protective coat give seeds their superior evolutionary advantage. Several layers of hardened tissue
prevent desiccation, and free reproduction from the need for a constant
supply of water. Furthermore, seeds remain in a state of dormancy—induced
by desiccation and the hormone abscisic acid—until conditions for growth
become favorable. Whether blown by the wind, floating on water, or carried
away by animals, seeds are scattered in an expanding geographic range, thus
avoiding competition with the parent plant.

Pollen grains (Figure) are male gametophytes and are carried by wind, water,
or a pollinator. The whole structure is protected from desiccation and can
reach the female organs without dependence on water. Male gametes reach
female gametophyte and the egg cell gamete though a pollen tube: an
extension of a cell within the pollen grain. The sperm of modern
gymnosperms lack flagella, but in cycads and the Gingko, the sperm still
possess flagella that allow them to swim down the pollen tube to the female
gamete; however, they are enclosed in a pollen grain.
This fossilized pollen is from a Buckbean fen core found in Yellowstone National
Park, Wyoming. The pollen is magnified 1,054 times.

Evolution of Angiosperms
Undisputed fossil records place the massive appearance and diversification of angiosperms in the middle to late
Mesozoic era. Angiosperms (“seed in a vessel”) produce a flower containing male and/or female reproductive
structures. Fossil evidence (Figure) indicates that flowering plants first appeared in the Lower Cretaceous, about
125 million years ago, and were rapidly diversifying by the Middle Cretaceous, about 100 million years ago. Earlier
traces of angiosperms are scarce. Fossilized pollen recovered from Jurassic geological material has been
attributed to angiosperms. A few early Cretaceous rocks show clear imprints of leaves resembling angiosperm
leaves. By the mid-Cretaceous, a staggering number of diverse flowering plants crowd the fossil record. The same
geological period is also marked by the appearance of many modern groups of insects, including pollinating
insects that played a key role in ecology and the evolution of flowering plants.
Although several hypotheses have been offered to explain this sudden profusion and variety of flowering plants,
none have garnered the consensus of paleobotanists (scientists who study ancient plants). New data in
comparative genomics and paleobotany have, however, shed some light on the evolution of angiosperms. Rather
than being derived from gymnosperms, angiosperms form a sister clade (a species and its descendents) that
developed in parallel with the gymnosperms. The two innovative structures of flowers and fruit represent an
improved reproductive strategy that served to protect the embryo, while increasing genetic variability and range.
Paleobotanists debate whether angiosperms evolved from small woody bushes, or were basal angiosperms
related to tropical grasses. Both views draw support from cladistics studies, and the so-called woody magnoliid
hypothesis—which proposes that the early ancestors of angiosperms were shrubs—also offers molecular
biological evidence.

The most primitive living angiosperm is considered to be Amborella trichopoda, a small plant native to the
rainforest of New Caledonia, an island in the South Pacific. Analysis of the genome of A. trichopoda has shown that
it is related to all existing flowering plants and belongs to the oldest confirmed branch of the
angiosperm family tree. A few other angiosperm groups called basal angiosperms, are viewed
as primitive because they branched off early from the phylogenetic tree. Most modern
angiosperms are classified as either monocots or eudicots, based on the structure of their
leaves and embryos. Basal angiosperms, such as water lilies, are considered more primitive
because they share morphological traits with both monocots and eudicots.
This leaf imprint shows a Ficus speciosissima, an angiosperm that flourished during the
Cretaceous period.

• Flowers and Fruits as an Evolutionary Adaptation

Angiosperms produce their gametes in separate organs, which are usually housed in a flower. Both fertilization
and embryo development take place inside an anatomical structure that provides a stable system of sexual
reproduction largely sheltered from environmental fluctuations. Flowering plants are the most diverse phylum on
Earth after insects; flowers come in a bewildering array of sizes, shapes, colors, smells, and arrangements. Most
flowers have a mutualistic pollinator, with the distinctive features of flowers reflecting the nature of the pollination
agent. The relationship between pollinator and flower characteristics is one of the great examples of coevolution.

Following fertilization of the egg, the ovule grows into a seed. The surrounding tissues of the ovary thicken,
developing into a fruit that will protect the seed and often ensure its dispersal over a wide geographic range. Not
all fruits develop from an ovary; such structures are “false fruits.” Like flowers, fruit can vary tremendously in
appearance, size, smell, and taste. Tomatoes, walnut shells and avocados are all examples of fruit. As with pollen
and seeds, fruits also act as agents of dispersal. Some may be carried away by the wind. Many attract animals that
will eat the fruit and pass the seeds through their digestive systems, then deposit the seeds in another location.
Cockleburs are covered with stiff, hooked spines that can hook into fur (or clothing) and hitch a ride on an animal
for long distances. The cockleburs that clung to the velvet trousers of an enterprising Swiss hiker, George de
Mestral, inspired his invention of the loop and hook fastener he named Velcro.

Evolution Connection
Building Phylogenetic Trees with Analysis of DNA Sequence Alignments. All living organisms display patterns
of relationships derived from their evolutionary history. Phylogeny is the science that describes the relative
connections between organisms, in terms of ancestral and descendant species. Phylogenetic trees, such as the
plant evolutionary history shown in Figure, are tree-like branching diagrams that depict these relationships.
Species are found at the tips of the branches. Each branching point, called a node, is the point at which a single
taxonomic group (taxon), such as a species, separates into two or more species.
 This phylogenetic tree shows the evolutionary
relationships of plants.

Phylogenetic trees have been built to

describe the relationships between species
since Darwin’s time. Traditional methods
involve comparison of homologous
anatomical structures and embryonic
development, assuming that closely related
organisms share anatomical features
during embryo development. Some traits
that disappear in the adult are present in
the embryo; for example, a human fetus, at
one point, has a tail. The study of fossil
records shows the intermediate stages that
link an ancestral form to its descendants.
Most of these approaches are imprecise
and lend themselves to multiple
interpretations. As the tools of molecular
biology and computational analysis have been developed and perfected in recent years, a new generation of tree-
building methods has taken shape. The key assumption is that genes for essential proteins or RNA structures,
such as the ribosomal RNA, are inherently conserved because mutations (changes in the DNA sequence) could
compromise the survival of the organism. DNA from minute amounts of living organisms or fossils can be
amplified by polymerase chain reaction (PCR) and sequenced, targeting the regions of the genome that are most
likely to be conserved between species. The genes encoding the ribosomal RNA from the small 18S subunit and
plastid genes are frequently chosen for DNA alignment analysis.

Once the sequences of interest are obtained, they are compared with existing sequences in databases such as
GenBank, which is maintained by The National Center for Biotechnology Information. A number of computational
tools are available to align and analyze sequences. Sophisticated computer analysis programs determine the
percentage of sequence identity or homology. Sequence homology can be used to estimate the evolutionary
distance between two DNA sequences and reflect the time elapsed since the genes separated from a common
ancestor. Molecular analysis has revolutionized phylogenetic trees. In some cases, prior results from
morphological studies have been confirmed: for example, confirming Amborella trichopoda as the most primitive
angiosperm known. However, some groups and relationships have been rearranged as a result of DNA analysis.

Section Summary
Seed plants appeared about one million years ago, during the Carboniferous period. Two major innovations—
seed and pollen—allowed seed plants to reproduce in the absence of water. The gametophytes of seed plants
shrank, while the sporophytes became prominent structures and the diploid stage became the longest phase of
the lifecycle. Gymnosperms became the dominant group during the Triassic. In these, pollen grains and seeds
protect against desiccation. The seed, unlike a spore, is a diploid embryo surrounded by storage tissue and
protective layers. It is equipped to delay germination until growth conditions are optimal. Angiosperms bear both
flowers and fruit. The structures protect the gametes and the embryo during its development. Angiosperms
appeared during the Mesozoic era and have become the dominant plant life in terrestrial habitats.

GYMNOSPERMS
Gymnosperms, meaning “naked seeds,” are a diverse group of seed plants and are paraphyletic. Paraphyletic
groups are those in which not all members are descendants of a single common ancestor. Their characteristics
include naked seeds, separate female and male gametes, pollination by wind, and tracheids (which transport
water and solutes in the vascular system).
Gymnosperm seeds are not enclosed in an ovary; rather, they are exposed on cones or modified leaves.
Sporophylls are specialized leaves that produce sporangia. The term strobilus (plural = strobili) describes a tight
arrangement of sporophylls around a central stalk, as seen in cones. Some seeds are enveloped by sporophyte
tissues upon maturation. The layer of sporophyte tissue that surrounds the megasporangium, and later, the
embryo, is called the integument.
Gymnosperms were the dominant phylum in Mesozoic era. They are adapted to live where fresh water is scarce
during part of the year, or in the nitrogen-poor soil of a bog. Therefore, they are still the prominent phylum in the
coniferous biome or taiga, where the evergreen conifers have a selective advantage in cold and dry weather.
Evergreen conifers continue low levels of photosynthesis during the cold months, and are ready to take advantage
of the first sunny days of spring. One disadvantage is that conifers are more susceptible than deciduous trees to
infestations because conifers do not lose their leaves all at once. They cannot, therefore, shed parasites and
restart with a fresh supply of leaves in spring.

The life cycle of a gymnosperm involves alternation of generations, with a dominant sporophyte in which the
female gametophyte resides, and reduced gametophytes. All gymnosperms are heterosporous. The male and
female reproductive organs can form in cones or strobili. Male and female sporangia are produced either on the
same plant, described as monoecious (“one home” or bisexual), or on separate plants, referred to
as dioecious (“two homes” or unisexual) plants. The life cycle of a conifer will serve as our example of reproduction
in gymnosperms.

Life Cycle of a Conifer

Pine trees are conifers (cone bearing) and carry both male and female sporophylls on the same mature
sporophyte. Therefore, they are monoecious plants. Like all gymnosperms, pines are heterosporous and generate
two different types of spores: male microspores and female megaspores. In the male cones, or staminate cones,
the microsporocytes give rise to pollen grains by meiosis. In the spring, large amounts of yellow pollen are
released and carried by the wind. Some gametophytes will land on a female cone. Pollination is defined as the
initiation of pollen tube growth. The pollen tube develops slowly, and the generative cell in the pollen grain divides
into two haploid sperm cells by mitosis. At fertilization, one of the sperm cells will finally unite its haploid nucleus
with the haploid nucleus of a haploid egg cell.
Female cones, or ovulate cones, contain two ovules per scale. One megaspore mother cell, or megasporocyte,
undergoes meiosis in each ovule. Three of the four cells break down; only a single surviving cell will develop into
a female multicellular gametophyte, which encloses archegonia (an archegonium is a reproductive organ that
contains a single large egg). Upon fertilization, the diploid egg will
give rise to the embryo, which is enclosed in a seed coat of tissue
from the parent plant. Fertilization and seed development is a long
process in pine trees: it may take up to two years after pollination.
The seed that is formed contains three generations of tissues: the
seed coat that originates from the sporophyte tissue, the
gametophyte that will provide nutrients, and the embryo itself.
Figure illustrates the life cycle of a conifer. The sporophyte (2n)
phase is the longest phase in the life of a gymnosperm. The
gametophytes (1n)—microspores and megaspores—are reduced
in size. It may take more than year between pollination and
fertilization while the pollen tube grows towards the
megasporocyte (2n), which undergoes meiosis into megaspores.
The megaspores will mature into eggs (1n).
This image shows the life cycle of a conifer. Pollen from male cones
blows up into upper branches, where it fertilizes female cones.

Diversity of Gymnosperms
Modern gymnosperms are classified into four phyla. Coniferophyta, Cycadophyta, and Ginkgophyta are similar in
their production of secondary cambium (cells that generate the vascular system of the trunk or stem and are
partially specialized for water transportation) and their pattern of seed development. However, the three phyla
are not closely related phylogenetically to each other. Gnetophyta are considered the closest group to
angiosperms because they produce true xylem tissue.

• Conifers (Coniferophyta)
Conifers are the dominant phylum of gymnosperms, with the most variety of species (Figure). Most are typically
tall trees that usually bear scale-like or needle-like leaves. Water evaporation from leaves is reduced by their thin
shape and the thick cuticle. Snow slides easily off needle-shaped leaves, keeping the load light and decreasing
breaking of branches. Adaptations to cold and dry weather explain the predominance of conifers at high altitudes
and in cold climates. Conifers include familiar evergreen trees such as pines, spruces, firs, cedars, sequoias, and
 yews. A few species are deciduous and lose their leaves in fall. The European
larch and the tamarack are examples of deciduous conifers (Figurec). Many
coniferous trees are harvested for paper pulp and timber. The wood of conifers
is more primitive than the wood of angiosperms; it contains tracheids, but no
vessel elements, and is therefore referred to as “soft wood.”
Conifers are the dominant form of vegetation in cold or arid environments and
at high altitudes. Shown here are the (a) evergreen spruce Picea sp., (b)
juniper Juniperus sp., (c) sequoia Sequoia Semervirens, which is a deciduous
gymnosperm, and (d) the tamarack Larix larcinia. Notice the yellow leaves of
the tamarack.

• Cycads
Cycads thrive in mild climates, and are often mistaken for palms because of the shape of their large, compound
leaves. Cycads bear large cones (Figure), and may be pollinated by beetles rather than wind: unusual for a
gymnosperm. They dominated the landscape during the age of dinosaurs in the
Mesozoic, but only a hundred or so species persisted to modern times. They face possible
extinction, and several species are protected through international conventions. Because
of their attractive shape, they are often used as ornamental plants in gardens in the
tropics and subtropics.
This Encephalartos ferox cycad has large cones and broad, fern-like leaves.

• Gingkophytes
The single surviving species of the gingkophytes group is the Gingko biloba (Figure). Its fan-
shaped leaves—unique among seed plants because they feature a dichotomous venation
pattern—turn yellow in autumn and fall from the tree. For centuries, G. biloba was cultivated
by Chinese Buddhist monks in monasteries, which ensured its preservation. It is planted in
public spaces because it is unusually resistant to pollution. Male and female organs are
produced on separate plants. Typically, gardeners plant only male trees because the seeds
produced by the female plant have an off-putting smell of rancid butter. This plate from the
1870 book Flora Japonica, Sectio Prima (Tafelband) depicts the leaves and fruit of Gingko biloba,
as drawn by Philipp Franz von Siebold and Joseph Gerhard Zuccarini.
• Gnetophytes
Gnetophytes are the closest relative to modern angiosperms, and include three dissimilar genera of
plants: Ephedra, Gnetum, and Welwitschia (Figure). Like angiosperms, they have broad leaves. In tropical and
subtropical zones, gnetophytes are vines or small shrubs. Ephedra occurs in dry areas of the West Coast of the
United States and Mexico. Ephedra’s small, scale-like leaves are the source of the compound ephedrine, which is
used in medicine as a potent decongestant. Because ephedrine is similar to amphetamines, both in chemical
structure and neurological effects, its use is restricted to prescription drugs. Like angiosperms, but unlike other
gymnosperms, all gnetophytes possess vessel elements in their xylem.
(a) Ephedra viridis, known by the common
name Mormon tea, grows on the West Coast of the
United States and Mexico. (b) Gnetum
gnemon grows in Malaysia. (c) The
large Welwitschia mirabilis can be found in the
Namibian desert.

Section Summary
Gymnosperms are heterosporous seed plants that produce naked seeds. They appeared in the Paleozoic period
and were the dominant plant life during the Mesozoic. Modern-day gymnosperms belong to four phyla. The
largest phylum, Coniferophyta, is represented by conifers, the predominant plants at high altitude and latitude.
Cycads (phylum Cycadophyta) resemble palm trees and grow in tropical climates. Gingko biloba is the only
representative of the phylum Gingkophyta. The last phylum, Gnetophyta, is a diverse group of shrubs that produce
vessel elements in their wood.
ANGIOSPERMS
From their humble and still obscure beginning during the early Jurassic period, the angiosperms—or flowering
plants—have evolved to dominate most terrestrial ecosystems (Figure). With more than 250,000 species, the
angiosperm phylum (Anthophyta) is second only to insects in terms of diversification.
These flowers grow in a botanical garden border in Bellevue, WA. Flowering plants
dominate terrestrial landscapes. The vivid colors of flowers are an adaptation to
pollination by animals such as insects and birds.
The success of angiosperms is due to two novel reproductive structures: flowers and
fruit. The function of the flower is to ensure pollination. Flowers also provide
protection for the ovule and developing embryo inside a receptacle. The function of
the fruit is seed dispersal. They also protect the developing seed. Different fruit
structures or tissues on fruit—such as sweet flesh, wings, parachutes, or spines that grab—reflect the dispersal
strategies that help spread seeds.
• Flowers
Flowers are modified leaves, or sporophylls, organized around a central stalk. Although they vary greatly in
appearance, all flowers contain the same structures: sepals, petals, carpels, and stamens. The peduncle attaches
the flower to the plant. A whorl of sepals (collectively called the calyx) is located at the base of the peduncle and
encloses the unopened floral bud. Sepals are usually photosynthetic organs, although there are some exceptions.
For example, the corolla in lilies and tulips consists of three sepals and three petals that look virtually
identical. Petals, collectively the corolla, are located inside the whorl of sepals and often display vivid colors to
attract pollinators. Flowers pollinated by wind are usually small, feathery, and visually inconspicuous. Sepals and
petals together form the perianth. The sexual organs (carpels and stamens) are located at the center of the flower.
As illustrated in Figure, styles, stigmas, and ovules constitute
the female organ: the gynoecium or carpel. Flower structure
is very diverse, and carpels may be singular, multiple, or
fused. Multiple fused carpels comprise a pistil. The
megaspores and the female gametophytes are produced
and protected by the thick tissues of the carpel. A long, thin
structure called a style leads from the sticky stigma, where
pollen is deposited, to the ovary, enclosed in the carpel. The
ovary houses one or more ovules, each of which will develop
into a seed upon fertilization. The male reproductive organs,
the stamens (collectively called the androecium), surround
the central carpel. Stamens are composed of a thin stalk
called a filament and a sac-like structure called the anther.
The filament supports the anther, where the microspores
are produced by meiosis and develop into pollen grains.
This image depicts the structure of a perfect flower. Perfect
flowers produce both male and female floral organs. The flower
shown has only one carpel, but some flowers have a cluster of
carpels. Together, all the carpels make up the gynoecium.

• Fruit
As the seed develops, the walls of the ovary thicken and form the fruit. The seed forms in an ovary, which also
enlarges as the seeds grow. In botany, a fertilized and fully grown, ripened ovary is a fruit. Many foods commonly
called vegetables are actually fruit. Eggplants, zucchini, string beans, and bell peppers are all technically fruit
because they contain seeds and are derived from the thick ovary tissue. Acorns are nuts, and winged maple
whirligigs (whose botanical name is samara) are also fruit. Botanists classify fruit into more than two dozen
different categories, only a few of which are actually fleshy and sweet.

Mature fruit can be fleshy or dry. Fleshy fruit include the familiar berries, peaches, apples, grapes, and tomatoes.
Rice, wheat, and nuts are examples of dry fruit. Another distinction is that not all fruits are derived from the ovary.
For instance, strawberries are derived from the receptacle and apples from the pericarp, or hypanthium. Some
fruits are derived from separate ovaries in a single flower, such as the raspberry. Other fruits, such as the
pineapple, form from clusters of flowers. Additionally, some fruits, like watermelon and orange, have rinds.
Regardless of how they are formed, fruits are an agent of seed dispersal. The variety of shapes and characteristics
reflect the mode of dispersal. Wind carries the light dry fruit of trees and dandelions. Water transports floating
coconuts. Some fruits attract herbivores with color or perfume, or as food. Once eaten, tough, undigested seeds
are dispersed through the herbivore’s feces. Other fruits have burs and hooks to cling to fur and hitch rides on
animals.

The Life Cycle of an Angiosperm

The adult, or sporophyte, phase is the main phase of an
angiosperm’s life cycle (Figure). Like gymnosperms,
angiosperms are heterosporous. Therefore, they
generate microspores, which will generate pollen grains
as the male gametophytes, and megaspores, which will
form an ovule that contains female gametophytes.
Inside the anthers’ microsporangia, male gametophytes
divide by meiosis to generate haploid microspores,
which, in turn, undergo mitosis and give rise to pollen
grains. Each pollen grain contains two cells: one
generative cell that will divide into two sperm and a
second cell that will become the pollen tube cell.
The life cycle of an angiosperm is shown. Anthers and
carpels are structures that shelter the actual
gametophytes: the pollen grain and embryo sac. Double
fertilization is a process unique to angiosperms.

If a flower lacked a megasporangium, what type of

gamete would not form? If the flower lacked a
microsporangium, what type of gamete would not form?

The ovule, sheltered within the ovary of the carpel,

contains the megasporangium protected by two layers
of integuments and the ovary wall. Within each megasporangium, a megasporocyte undergoes meiosis,
generating four megaspores—three small and one large. Only the large megaspore survives; it produces the
female gametophyte, referred to as the embryo sac. The megaspore divides three times to form an eight-cell
stage. Four of these cells migrate to each pole of the embryo sac; two come to the equator, and will eventually
fuse to form a 2n polar nucleus; the three cells away from the egg form antipodals, and the two cells closest to
the egg become the synergids.
The mature embryo sac contains one egg cell, two synergids or “helper” cells, three antipodal cells, and two polar
nuclei in a central cell. When a pollen grain reaches the stigma, a pollen tube extends from the grain, grows down
the style, and enters through the micropyle: an opening in the integuments of the ovule. The two sperm cells are
deposited in the embryo sac.

A double fertilization event then occurs. One sperm and the egg combine, forming a diploid zygote—the future
embryo. The other sperm fuses with the 2n polar nuclei, forming a triploid cell that will develop into the
endosperm, which is tissue that serves as a food reserve. The zygote develops into an embryo with a radicle, or
small root, and one (monocot) or two (dicot) leaf-like organs called cotyledons. This difference in the number of
embryonic leaves is the basis for the two major groups of angiosperms: the monocots and the eudicots. Seed
food reserves are stored outside the embryo, in the form of complex carbohydrates, lipids or proteins. The
cotyledons serve as conduits to transmit the broken-down food reserves from their storage site inside the seed
to the developing embryo. The seed consists of a toughened layer of integuments forming the coat, the
endosperm with food reserves, and at the center, the well-protected embryo.
Most flowers are monoecious or bisexual, which means that they carry both stamens and carpels; only a few
species self-pollinate. Monoecious flowers are also known as “perfect” flowers because they contain both types of
sex organs (Figure). Both anatomical and environmental barriers promote cross-pollination mediated by a
physical agent (wind or water), or an animal, such as an insect or bird. Cross-pollination increases genetic diversity
in a species.
Diversity of Angiosperms
Angiosperms are classified in a single phylum: the Anthophyta. Modern angiosperms appear to be a monophyletic
group, which means that they originate from a single ancestor. Flowering plants are divided into two major groups,
according to the structure of the cotyledons, pollen grains, and other structures. Monocots include grasses and
lilies, and eudicots or dicots form a polyphyletic group. Basal angiosperms are a group of plants that are
believed to have branched off before the separation into monocots and eudicots because they exhibit traits from
both groups. They are categorized separately in many classification schemes. The Magnoliidae (magnolia trees,
laurels, and water lilies) and the Piperaceae (peppers) belong to the basal angiosperm group.

• Basal Angiosperms
The Magnoliidae are represented by the magnolias: tall trees bearing large, fragrant flowers that have many parts
and are considered archaic (Figured). Laurel trees produce fragrant leaves and small, inconspicuous flowers.
The Laurales grow mostly in warmer climates and are small trees and shrubs. Familiar plants in this group include
the bay laurel, cinnamon, spice bush (Figurea), and avocado tree. The Nymphaeales are comprised of the water
lilies, lotus (Figurec), and similar plants; all species thrive in freshwater biomes, and have leaves that float on the
water surface or grow underwater. Water lilies are particularly prized by
gardeners, and have graced ponds and pools for thousands of years.
The Piperales are a group of herbs, shrubs, and small trees that grow in the
tropical climates. They have small flowers without petals that are tightly
arranged in long spikes. Many species are the source of prized fragrance or
spices, for example the berries of Piper nigrum (Figureb) are the familiar
black peppercorns that are used to flavor many dishes.
The (a) common spicebush belongs to the Laurales, the same family as
cinnamon and bay laurel. The fruit of (b) the Piper nigrum plant is black pepper,
the main product that was traded along spice routes. Notice the small,
unobtrusive, clustered flowers. (c) Lotus flowers, Nelumbo nucifera, have been
cultivated since ancient times for their ornamental value; the root of the lotus
flower is eaten as a vegetable. The red seeds of (d) a magnolia tree, characteristic
of the final stage, are just starting to appear.

• Monocots
Plants in the monocot group are primarily identified as such by the presence of a single cotyledon in the seedling.
Other anatomical features shared by monocots include veins that run parallel to the length of the leaves, and
flower parts that are arranged in a three- or six-fold symmetry. True woody tissue is rarely found in monocots. In
palm trees, vascular and parenchyma tissues produced by the primary and secondary thickening meristems form
the trunk. The pollen from the first angiosperms was monosulcate, containing a single furrow or pore through the
outer layer. This feature is still seen in the modern monocots.
Vascular tissue of the stem is not arranged in any particular
pattern. The root system is mostly adventitious and unusually
positioned, with no major tap root. The monocots include
familiar plants such as the true lilies (which are at the origin of
their alternate name of Liliopsida), orchids, grasses, and palms.
Many important crops are monocots, such as rice and other
cereals, corn, sugar cane, and tropical fruits like bananas and
pineapples (Figure). The world’s major crops are flowering plants.
(a) Rice, (b) wheat, and (c) bananas are monocots, while (d) cabbage,
(e) beans, and (f) peaches are dicots.

• Eudicots
Eudicots, or true dicots, are characterized by the presence of two cotyledons in the developing shoot. Veins form
a network in leaves, and flower parts come in four, five, or many whorls. Vascular tissue forms a ring in the stem;
in monocots, vascular tissue is scattered in the stem. Eudicots can be herbaceous (like grasses), or produce woody
tissues. Most eudicots produce pollen that is trisulcate or triporate, with three furrows or pores. The root system
is usually anchored by one main root developed from the embryonic radicle. Eudicots comprise two-thirds of all
flowering plants. The major differences between monocots and eudicots are summarized in Table. Many species
exhibit characteristics that belong to either group; as such, the classification of a plant as a monocot or a eudicot
is not always clearly evident.
 Comparison of Structural Characteristics of Monocots and Eudicots
Characteristic Monocot Eudicot
Cotyledon One Two
Veins in Leaves Parallel Network (branched)
Stem Vascular Tissue Scattered Arranged in ring pattern
Roots Network of adventitious roots Tap root with many lateral roots
Pollen Monosulcate Trisulcate
Flower Parts Three or multiple of three Four, five, multiple of four or five and whorls

Section Summary
Angiosperms are the dominant form of plant life in most terrestrial ecosystems, comprising about 90 percent of
all plant species. Most crops and ornamental plants are angiosperms. Their success comes from two innovative
structures that protect reproduction from variability in the environment: the flower and the fruit. Flowers were
derived from modified leaves. The main parts of a flower are the sepals and petals, which protect the reproductive
parts: the stamens and the carpels. The stamens produce the male gametes in pollen grains. The carpels contain
the female gametes (the eggs inside the ovules), which are within the ovary of a carpel. The walls of the ovary
thicken after fertilization, ripening into fruit that ensures dispersal by wind, water, or animals.
The angiosperm life cycle is dominated by the sporophyte stage. Double fertilization is an event unique to
angiosperms. One sperm in the pollen fertilizes the egg, forming a diploid zygote, while the other combines with
the two polar nuclei, forming a triploid cell that develops into a food storage tissue called the endosperm.
Flowering plants are divided into two main groups, the monocots and eudicots, according to the number of
cotyledons in the seedlings. Basal angiosperms belong to an older lineage than monocots and eudicots.

DIG DEEPER
All plants are related. They are made of the same basic body plan (an aboveground part that functions for
photosynthesis and reproduction, and a belowground part that functions for attachment to the substrate and
nutrient acquisition). Almost all plants also have the same basic needs to survive - sunlight, air, water, and
nutrients. Yet how come we ended up with some 390,000 species of plants (not including the bryophytes)?
Hundreds of millions of years of evolution allowed plants to have variations in how they cope with the different
environments on land, obtain nutrients from the air and soil, defend themselves against herbivores and
competitors, and reproduce. Among the flowering plants, their distinct reproduction and co-evolution with their
animal pollinators allowed them to occupy a wider variety of habitats and diversify more successfully than the
other plant life forms that appeared on land.