Chapter 21

1. The earliest fossils that definitely were vascular land plants belong to Cooksonia,
a genus of extinct plants. They branched equal dicothomous branching, both
branches being of equal size and vigor.
2. Draw a plant of Rhynia. Be certain to include the reproductive parts. Now assume
that it has a life cycle with an alternation of isomorphic generations and draw a
complete life cycle.

3. If Rhynia or its contemporaries were the ancestors to the ferns, how did the
gametophytes and sporophytes change during evolution?
At first, it was postulated that algae gave rise to true plants whose life cycle was
dominated by the gamenophyte and whose sporophyte was small and dependent,
hemiparasitic on the gametophyte. It was postulated that this evolved into the
vascular plants as the sporophyte become larger and more elaborate while the
gametophyte became reduced. But it is possible that hornworts evolved by the
reverse process, reading this series from left to right.
4. In the vascular bundles of flowering plants, protoxylem is closest to the center of
the stem, and metaxylem is farther out. Is this an endarch or an exarch
arrangement, but what about the early vascular plants. Were they endarch or
exarch, or did both types occur originally.
In an endarch protostele, protoxylem is located in the center and the metaxylem
differentiates on the outer edge of the xylem mass. Protoxylem is the xylem that
differentiates while the cells are small and narrow, and metaxylem differentiates

after the cells have expanded for a few more hours or days and are larger. The other
type of stele present in early vascular plantsis an exarch protostele, with metaxylem
located in the center of the xylem mass and protoxylem on the edges as several
groups next to the phloem.
5. The vascular bundles of flowering plants surround a pith, but the earliest vascular
plants had no pith. A vascular system with a solid mass of xylem with no pith is
called a protostele. A stele that does have a pith is called a siphonostele.
6. Rocks that contain the fossils of Rhynia also often contain fossils of Sciadophyton.
We know that Rhynia was a sporophyte and that Sciadophyton was a gametophyte.
Because they grew together, we suspect that they may be the two generations of a
single species. If so, did Rhynia have alternation of isomorphic or heteromorphic
generations? Also, if this is true, evolution has caused one generation to become
more complex, the other to become less complex. Which is which? Explain.
Phynia-type sporophytes occur together with Sciadophyton-type gametophytes,
they might have been alternate phases of the same species. If true, this these have
an alternation of isomorphic generations and later evolution into seed plants
involved reduction of the gametophyte to just a few cells and elaboration of the
sporophyte into a more complex plants.
7. There are two alternative hypotheses about the life cycle of the early vascular
plants, the transformation hypothesis and interpolation hypothesis. If Rhynia and
Sciadophyton are two generations of one species, which hypothesis would be
favored? Briefly describe the other hypothesis.
An alternative hypothesis, the transformation theory, postulats that after
dibiontic life cycle originated, both gametophyte and sporophyte became larger,
more complex, and vascularized, in a life cycle with an alternation of isomorphic
generations.
8. What are the zosterophyllophytes? How did they differ from rhyniophytes? Why
do we think they are related to lycophytes but not to ferns and seed plants?
They were small herbs without secondary growth. They differ from rhyniophyes
because their sporangia were lateral, not terminal; sporangia opened traansversaly
along the top edge, and their xylem was an exarch protostele, that is, protoxylem
on the outer margin and metaxylem in the center. Some of the simplest vascular
plants alive today, the lycophytes, have lateral sporangia, and they may represent a
line of evolution based on Zosterophyllum- like ancestors.
9. The reconstruction of Asteroxylon in Figure 21-10 shows thin, leaf-like flaps of
tissue on the plants surface. What are these called? Did they ever have stomata in
any zosterophyllophytes? Did they have vascular bundles? They did become large?

Asteroxylon, they contained stomata and a small trace of vascular tissue that ran
from the stele through the cortex to the base of the enation. In figure 21-10, it
shows its surface covered with enations, small flaps of photosynthetic tissue. In
species without enations, stems were round and oriented vertically.
10. What are microphylls? Are they related to the enations of Asteroxylon?
Leaf is an ambiguous term, and enations in the division Lycophyta are called
microphylls for clarity.
11. The lycophytes once contained many species of large trees that formed
extensive forests. Briefly describe plants of Lepidodendron and Sigillaria. Also
describe their wood.
Many extinct lycophytes Lepidodedron, Sigillaria, and Stigmaria had a vascular
cambium and secondary growth. Their wood looked remarkably like the secondary
xylem of pines and other living conifers, having a pith, rays, and elongate tracheids.
12. Name two genera of living lycophytes. What are the common names? About
how big do they get? Are they leafy or do they have naked stems? In a plant
identification book, they would probably be listed with ferns.
-Lycopodium cernuum is a common species that has extensive rhizomes, vertical
chlorophyllous shoots, and sporangia clustered into cones. These shoots are leaning
against abd being supporte by surrounding grasses and shrubs.
-Lycopodium lucidulum is one of several lycopod species in which sporangia are
distributed among the leaves rather than a strobili.
13. Trimerophytes were plants that evolved from rhyniophytes but with more
derived features. In one feature, certain stems grow longer than the others, and
thus, rather than having dichotomous branching, they have pseudomonopodial
branching (displayed especially by Pertica). Simultaneously, the positioning of
branching became more regular and controlled.
14. Describe the evolution of megaphylls. What are telomes?
Megaphylls, leaves that evolved from branch systems and are present in all seed
plants, ferns, and equisetophytes.
The ultimate twigs, those of the last dichotomy, are known as telomes.
15. Describe the trimerophytes. From what group did they produce? Even through
all rhyniophytes and trimerophytes are now extinct, would you consider them
unsuccessful?

Trimerophtyes became distinct from rhyniophytes during the Lower Devonian and
existed until the Upper DSevonian Period and then came to an end not by being
extinct but rather by evolving into the ancestor if ferns and the seed plants.
16. What are the two common names of equisetums? What is the appearance of the
plants? Their approximate height? Equisetums have strobili and canals. Briefly
describe the canals and strobili.
Equisitum, with 15 extant species known as horsetails and scouring rushes. The
living plants are all herbs without any secondary growth, and although ceratin
species may attain a height of up to 10 m, they are usually less than 1 m.
17. Ferns first appeared in the Devonian period. Look at the inside and the back
cover. How long was the Devonian period? Unlike all the groups mentioned so far in
this chapter, most of this group is still extant, not extinct. About how many species
of modern ferns are there?
The Devonian is a geologic period and system of the Paleozoic Era spanning from
the end of the Silurian Period, about 419.2 3.2 Mya (million years ago), to the
beginning of the Carboniferous Period, about 358.9 0.4.
The Devonian period experienced the first significant adaptive radiation of
terrestrial life. Since large vertebrate terrestrial herbivores had not yet
appeared, free-sporing vascular plants began to spread across dry land, forming
extensive forests which covered the continents. By the middle of the Devonian,
several groups of plants had evolved leaves and true roots, and by the end of
the period the first seed-bearing plants appeared.
18. Name two genera of ferns that are found in deserts. Name two that grow
floating on water. One that lives underwater. These ferns are called tree ferns. Do
tree ferns have woody trunks with secondary xylem?
All ferns are perennial and herbaceous; none is woody, but some do achieve the
size of small trees- the tree ferns. Although called tree ferns, they never have
secondary xylem.
19. A very important feature of ferns is shown in Figure 21-29: Their shoot xylem is
not solid as it was in the rhyniophytes. In the evolution of ferns and seed plants,
there was the evolution of pith. A stele with a pith is called a siphonostele. Another
significant feature of ferns is that the phloem occurs both to the outer side as well
as the inner side of the xylem. Where does it occur in the vascular bundles of
flowering plants?
Vascular tissue may be scattered in ground tissue or regularly arranged
forming a ring. In the latter arrangement, ground tissue is diff erentiated
into cortex (outer to vascular tissue) and pith (inner to vascular tissue).

20. You may have noticed that the undersides (but never uppersides) of fern leaves
have brown dots or brown streaks or brown patches. The brown dots are called sori,
clusters of sporangia where meiosis occurs. If a leaf carried these structures, the
leaf is called heterosporous.
21. When fern spores germinate, they grow into small, simple heart-shaped or
ribbon-shaped photosynthetic gametophytes with unicellular rhizoids on the lower
surface but with no vascular tissue and no epidermis.
22. Briefly describe eusporangia and leptosporangia. Which is the type that occurs
in all other vascular plants? Which is the type that occurs in most ferns?

In a leptosporangium, found only in ferns, development involves a single

initial cell that becomes the stalk, wall, and spores within the sporangium. There
are around 64 spores in a leptosporangium.

In a eusporangium, characteristic of all other vascular plants and some

primitive ferns, the initials are in a layer (i.e., more than one). A eusporangium is
larger (hence contain more spores), and its wall is multi-layered. Although the
wall may be stretched and damaged, resulting in only one cell-layer remaining.