CHAPTER 4 o secondary wall
Evolution and Diversiy -secreted outside of plasma membrane &
 Vascular Plants (Tracheophyta)
-monophyletic group of land plants
Apomorphies
a.) Independent, long-lived sporophyte
b.) branched sporophyte
c.) lignin in lignified secondary walls (pits)
d.) schlerenchyma (cells for structural support)
e.) tracheary elements (xylem; for water transport)
f.) sieve elements (phloem; for sugar transport)
g.) endodermis (transfer of compounds)
h.) roots (anchorage & absorption of water and
nutrients)
 independent long lived sporophyte
o haplodiplontic = “alternation of generation”
o sporophyte (2n)
-dominant, free-living, photosynthetic
-grows larger and independent w/ death of
gametophyte
o gametophyte (n)
-free-living, photosynthetic, short-lived, smaller
than sporophyte
 branched sporophyte
o sporophyte axis, or stems
o Polysporangiophytes
-bear multiple sporangia
-branched sporophyte
*these are apomorphies shared w/ fossil plants
o earliest vascular plants stems had dichotomous
branching
o pseudomonopodial
-modified growth pattern
-dichotomous
-one is dominant and the other is lateral
o sporophyte functions:
-support
-bears reproductive organs and leaves
-conductive organ via vascular tissue
 lignified secondary cell walls
o lignin
-chemical; complex polymer of phenolic
compounds in secondary wall
-secreted into space b/w cellulose microfibrils
forming interbinding cement
-for strength and rigidity of cell wall
primary cell wall (cellulosic)
-thicker than cell wall and contain cellulose
o pits
-holes in secondary wall; in pairs in primary cell
wall
-communication b/w cells via plasmodesmata
-water conducting cells
o primary pit field
-group of plasmodesmata in primary cell wall
o Plant cells w/ secondary walls are sclerenchyma
and tracheary elements
 schlerenchyma
o scleros – hard; enchyma – infusion
o for greater stem height
o nonconductive cells – have thick, lignified
secondary cell wall w/ pits (dead at maturity)
o 2 types of schlerenchyma
1.fibers – long narrow cells w/ sharply tapering
end; for mechanical support; in bundles;
may/maybe not be components of vascular
tissues
2.sclereids – isodiametric to irregular or
branched; for structural support; deter
herbivory
o collenchyma – consists of live cells w/ unevenly
thickened, pectic primary cell walls; for
structural support; not an apomorphy
 tracheal elements (xylem)
o major adaptive breakthrough in plant evolution
o elongate, dead at maturity; have lignified
secondary cell walls; oriented end-to-end
o functions in water and mineral conduction
o associated w/ parenchyma and schlerenchyma
o 2 types of tracheal elements
1. tracheids – imperforate: fluid flow b/w
adjacent cells at pit-pairs in primary cell walls;
2. vessel members – perforate; no intervening
cell walls b/w adjacent cells and fluid pass
through holes or perforations
-from preexisting tracheids independently
(ex. Equisetum, leptosporangiate ferns,
gnetales, angiosperms)
*perforation plate – contact area of 2 adjacent
vessel members (may be simple – single
opening or compound – several perforations)
o Tracheids are the primitive type
 sieve elements (phloem)
o phloe – bark
 o function in conduction of sugars
o Regions:
1. Sugar-rich “Source” – leaves (sugars are
synthesized during photosynthesis); mature
storage organs (sugars are released by starch
hydrolysis)
2. Sugar-poor “Sink” – dividing cells, developing
storage organs, reproductive organs (flowers or
fruits)
o associated w/ parenchyma and schlerenchyma
o elongate; semi-alive at maturity (lose nucleus
but retain mitochondria, ER and plastids)
o no lignified secondary cell wall
o primary cell wall
-has pores: aggregated in sieve areas
-callose: polysaccharide w/ beta-1-3 glucose
units; surrounds spores
o 2 types of sieve elements
1.sieve cells – sieve areas on both ends and side
walls; ancestral sugar-conducting cells; in non-
flowering plants
2.sieve tube members – sieve areas and sieve
plates; in flowering plants (angiosperms)
*both function to load and unload sugars in
their cavity
*sieve plates – consists sieve areas at the end
wall junction of two sieve tubes; pores – larger
than pores of sieve areas on their side walls
o Parenchyma cells
1.Albuminous cells – sieve cells; different
parent cells
2.Companion cells – sieve tube members; same
parent cells
o stele - organization of xylem and phloem in the
stem
o Stelar types
1. Protostele – central solid cylinder of phloem
and xylem; ancestral type (ex. Rhyniophytes)
2. Plectostele – modification of protostele;
phloem and xylem interdigitate
o cortex – large parenchyma tissue b/w epidermis
vascular tissues
 endodermis
o special cylinders of cells in roots or stems
o controls which plants are absorbed by the plant
o Casparian strip
-band or ring of lignin; water impermeable
-binds to plasma membrane of endodermis
*water & minerals pass through plasma
membrane (controls solute transfer)
o Suberin – similar to lignin; infiltrates the cell
wall, oriented tangenially (transverse walls) and
axially (radial walls)
 roots
o for anchorage and absorption of water and
animals
o lost roots secondarily or not found in Psilotales,
Salviniales
o if no roots, have uniseriate (one cell thick) and
filamentous rhizoids –for absorption
o apical meristem (in root tip)
-region of continuous mitotic divisions
-where roots & stems develop
-Selaginellaceae & Monilophytes: single, apical
cell
-Lycopodiaceae, Isoetaceae & seed plants:
complex, dividing cells
o Anatomical structures
1. rootcap
-aka calyptra; covers apical meristem;
provide lubrication; not in stems
2. root hairs
-hairlike extensions in the epidermal cells
away from the root tip
-greatly increase the surface area
-not in stems and species of Psilotopsida –
has rhizoids instead
3. central vascular cylinder
*cortex
-parenchymatous region b/w the
vasculature and epidermis
*pith
-center of vascular cylinder (if vascular
tissue is lacking)
4. endodermis w/ casparian strips
-surrounds the vascular cylinder
-selective absorption
*pericycle
-undifferentiated layer internal to the
endodermis
5. endogenous lateral roots
-origin of new lateral roots
-arise at pericycle or endodermis
-penetrate tissues of cortex
o mycorrhizae
-symbiotic interaction w/ fungi
-fungi aids plant in increasing surface area and
increase efficiency in selective absorption
-fungi benefits by obtaining photosynthates
(sugars) from the plant
 VASCULAR PLANT DIVERSITY -lycophylls originated from enations (small
o features that classify vascular plants: appendages) or from sterilization and planation
-sporophyte vegetative morphology (flattening) of sporangia
-life cycle -dorsiventral organs
-reproductive morphology
-gametophyte morphology  Lycopodiopsida
-spore features o club-mosses
o laesura (spore feature) o no ligules
-differentially thickened wall region on the o homospory – one type of spore
immature spores; divide through meiosis o sporangia develop laterally in sporophylls
3 basic spore types *sporophylls – axils of specialized leaves;
1. trilete – w/ a 3 branched laesura similar to vegetative leaves
2. monolete – linear and unbranched laesura o strobilus or cone – determinate reproductive
3. alete – lacking any evidence of a laesura shoot consisting of terminal aggregate of
sporophylls w/ sporangia; scale-like and
nonphotosynthetic
RHYNIOPHYTES
o paraphyletic F
Lycopodiaceae
o w/ dichotomous branched sporophyte axis that o lykos – wolf; podion – foot
bears terminal sporangia o terrestrial, epiphytic, pendulous, perennial,
o lack roots and leaf bearing system lycophyllous herbs
o stems are protostelic o roots – dichotomously branched, adventitious,
o protoxylem was “centrarch” endogenous
o Rhynia sp. o stems – dichotomously branched rhizoids or
corms, w/ protostelic vasculature;
pseudomonopodial (L. complanatum)
LYCOPODIOPHYTA - LYCOPHYTES o bulbils or gemmae – specialized short shoots;
o Zosterophyllophytes may detach, function as vegetative granules
-extinct, paraphyletic fossil group o leaves – simple, sessile, spiral/whorled, w/ a
-no leaves single midrib (microphyllous/lycophyllous), no
-w/ lateral sporangia ligule (eligulate)
-leads to lycophytes o sporangia – homosporous, reniform born on
sporophylls or strobili
Apomorphies of lycophytes o spores – globose/tetrahedral w/ trilete laesura
1. dichopodial roots o gametophytes – mycorrhizal, either
-root apical meristem branched into 2 roots photosynthetic or saprophytic
-no lateral roots o Genera – Huperzia, Lycopodiella, Lycopodium,
2. endarch protoxylem (roots) Pseudolycopodiella, Phylloglossum
-protoxylem: first tracheary cells w/ smaller and
thinner than metaxylem  Isoetopsida
-interior to the metaxylem toward the stem o Selaginellaceae and Isoetaceae – two extanct
center lycophyte families
3. exarch protoxylem (stems) o have ligules
-exterior to the metaxylem or away the stem o heterosporous
center -microspores and megaspores in sporangia
4. dorsiventral xporangia (produced in shoots)
-dehisce (splits) transversely o megaspore
5. sporophytic leaves -megaspores are fewer (4 per megasporangium
-lycophylls: single, unbranched vein; no gap in in Selaginella, more in Isoetes)
the stem vasculature and develops by -develops into female gametophyte: archegonia
intercalary meristem; develops from a shoot housing egg cell
apex
*shoot: stem + leaves
 o microspore
-develops into male gametophyte: antheridia,
the sperm manufacturing organs
o gametophyte – endosporic; develop within the
spore wall
o wood was an apomorphy (ex. in Lepidodendron
and Isoetes)
F arise from endodermis for monilophytes or
Isoetaceae
o quillworts
o isos – ever; etas – green
o aquatic or terrestrial, perennial herbs
o roots – adventitious, endogenous, dichopodial
o stems – protostelic, vertically-oriented and
corm-like, rarely rhizomatous, w/ apical and
lateral meristem, base is lobed
o leaves – simple, spiral, basal rosette; blades are
sheathing, apical linear or circular, have ligules;
single midrib (lycophyllous)
o sporangia – heterosporous, located in
sporophylls (adaxial side)
o megasporangia – in outer leaves; 50-300 per
megasporangium; trilete, spore sculpturing for
species identification
o microsporangia – in inner leaves, monolete, 1M
o gametophytes – covered by membrane velum
and transversed by trabeculae (sterile strands);
endosporic
o CAM photosynthesis
o Air chambers in roots and leaves
Selaginellaceae
F o Trimerophytes
o spike-mosses
o selago – moss-like; ella – diminutive
o perennial herbs, rarely tree-like, xeric-adapted
“resurrection plants”
o roots – adventitious and dichopodial, some are
rhizophores (leafless stems)
o stems – dichotomously branching w/ erect
cespitose, prostate or climbing habit;
pseudomonopodial or sympodial; protostelic
o leaves – simple, sessile, spiral, single midrib
(microphyllous), ligulate, blades are small,
homomorphic (isophyllous) or dimorphic
(anisophyllous) in 4 rows
o sporangia – heterosporous
o ligulate sporophylls – microsporophylls and
megasporophylls; where microsporangia and
megasporangia occur
o gametophytes – endosporic
EUPHYLLOPHYTA – EUPHYLLOPHYTES
o sister group of lycophytes
Apomorphies of euphyllophytes
1. roots are monopodial
-do not dichotomously
-branch at the apical meristemslateral roots
pericycle for spermatophytes
2. roots have exarch protoxylem
-in outer part of metaxylem
3. sporangia
-terminal in position; dehisce longitudinally
4. molecular apomorphy
-30 kilobase inversion in chloroplast DNA
5. leaves
-euphylls: synonymous to megaphylls
o euphylls
-photosynthesis
-leaf gap: nonvascular parenchyma tissue
-have 1 vein per leaf; highly branched system of
veins
-mesophylls: chloroplast-containing tissue
-grow by marginal or apical meristems
-evolve via transformation of 3-dimensional non
planar lateral branched system into a leaf
-planation: flattening of the axes into a 2
dimensional plane
-webbing: development of thin tissues b/w axes
-“web” functioned a mesophyll
-extinct, paraphyletic group
-leads to the common ancestor of lycophytes
and euphyllophytes
-sporophytes w/ no leaves
-stems were photosynthetic and
pseudomonopodial
Leaf characteristics
o leaf primordium – result of growth
differentiation of apical meristem region;
matures into a leaf
o node – point of attachment of a leaf to the
stem
o internode – region between 2 nodes
o vascular strands – run b/w stem and leaf
providing connection for fluid transport
o vascular bundles – aka veins; for conduction to
mesophyll; contain vascular tissues
o mesophyll – upper columnar palisade cells;
lower irregular spongy cells
 o stomata – gas exchange; in the lower epidermis
of leaves
o axil – tissue at the upper region of stem and
upper leaf
o immature shoot system – axil develops into
bud
o monopodial – pattern of growth in which
lateral branches develop from axillary buds
MONILOPHYTA – MONILOPHYTES, FERNS
o 4 major lineages: Equisetopsida (horsetails),
Psilotopsida (whisk ferns & ophioglossoid
ferns), Marratiopsida (marratoid ferns),
Polypodiopsia or Leptosporangiatae
(leptosporangiate ferns)
o monilo – necklace or string of beads; phyt –
plant
o monophyletic group
o former name: moniliformopses
Apomorhies of Monilophytes
1. Siphonostele
-type of stem vasculature in which a ring of
xylem is surrounded by an outer layer of
phloem (ectoploich siphonostele) or inner layer
of phloem (amphiphloic siphonostele or
solenostele) and if dissected (dictyostele)
2. stem protoxylem is mesearch
-tracheary elements first mature near the
middle of a patch of xylem
-restricted to the lobes of the xylem
 Equisetopsida
o horsetails
o equisetophytes, sphenophytes, sphenopsids
o monophyletic
o Calamites – large woody tree
o Only species remaining – Equisetum
Apomorphies of Equisetophytes
1. ridged stems
-associated w/ hollow canals
2. reduced whorled leaves
-marginally fused
3. sporangiophores
-consists of a peltate axis bearing pendant
longitudinally dehiscent sporangia
4. photosynthetic spores w/ elaters
 Equisetaceae
o Horsetail/scouring-rush family
o Equus – horse; seta – bristle
o Perennial herbs; growing in wet (sometimes
xeric) habitat
o stems
-rhizomes; aerial stems are ridged and
photosynthetic w/ epidermis containing silica
inclusions
-have a hollow pith and 2 rings of canals
(lacunae); inner ring canals (carinal)
o leaves
-small, simple, whorled microphyllous (1
veined), nonphotosynthetic at maturity and
laterally connate in a sheath w/ tooth-like
apices
o sporangia
-homosporous, born in terminal strobili, peltate
sporangiophores
o spores
-lack an attachment scar
-spherical, green, 4 spatulate hygroscopic
elaters
o gametophytes
-photosynthetic and cushion-like
o subgenus: Equisetum (horsetails; whorled
lateral branches and stomates flush w/ the
epidermis), Hippachaete (scouring rushes; lack
lateral branches w/ sunken stomates)
 Psilotopsida
OphioglossalesO
o Ophioglossoid ferns
o Each leaf consist of sterile segment (contains
the photosynthetic blade or lamina) and a
fertile fragment (bears sporangia)
o Rhizomes give rise to unbranched roots that
lack root hairs
o Sporangia – eusporangia
-relatively large
-has a sporangial wall w/ more than 1 cell layer
-ancestral condition of the land plants
F
Ophioglossaceae
o Adder’s Tongue
o ophis – snake; glossa – tongue
o terrestrial, perennial herbs
o roots – fleshy, mycorrhizal, lacking root hairs,
bears adventitious buds (grows into a plantlet)
o stems – subterranean, erect; protostelic
vasculature or ectophloic siphonostele
o leaf – solitary, lacking circinate vernation, blade
simple and unlobed or compound to divided (1-
2 pinnatifid), venation is open-dichotomous or
reticulate
o sori – lacking
o sporangia – homosporous; eusporangia – born
on a stalked fertile segment, either spike-like
(Ophioglossum) or branched and panicle-like
sporangial dehiscence is transverse
o spores – tetrahedral and trilete
o gametophytes – nonphotosynthetic and
mycorrhizal or mycotrophic
PsilotalesO
o whisk ferns
o 2 genera: Psilotum &Tmesipteris
o sporophyte consists of a horizontal rhizome
that gives rise to aerial, photosynthetic,
dichotomous branching stems
o epiphytic w/ rhizomes having mychorrhizal
associations; only absorptive rhizoids arise from
rhizomes
o lack true roots (an apomorphy)
o leaves
-reduced and peg-like
-lack a vascular strand
-enations:outgrowths
-w/ midrib
o sporangia
-2 to 3 lobed
-synangium: fusion product of 2 or 3 sporangia;
yellowish at maturity and are subtended by a
forked appendage (apomorphy)
o gametophytes
-nonphotosynthetic and may contain
mycorrhizal fungi
o Psilotum nudum – whisk broom; cultivated in
greenhouses and naturalized in warm climates
F o large, terrestrial ferns; tissues with
Psilotaceae
o whisk ferns
o psilos – naked
o consists of terrestrial/epiphytic, perennial herbs
o roots are absent
o stems
-underground: rhizoid-bearing with endophytic
mycorrhizae
-aerial: photosynthetic, terete, ridged, or
flattened; dichotomously
branched/unbranched; protostele or
solenostele vasculature
o leaves
-simple, spiral, or distichous
-blades small & subulate to scale-like
-Tmesipteris: larger blades & vertically oriented,
either with single midrib (microphyllous) or
lacking vasculature
-enation/prophyll: vasculature runs to base
o sporangia
-eusporangiate; homosporous, arising from
short lateral branches;subtended by a bifid
appendage
-synagiate: the product of fusion of 3 (Psilotum)
or 2 (Tmesipteris) sporangia
o spores
-reniform & monolete
o gametophytes
-nonphotosynthetic; cylindrical; saprophytic;
mycorrhizal; sperm is multiflagellate
MARATIOPSIDA - MARATTOID FERNS
o contains the single order Marattiales and
family Marattiaceae
o general form,
-large pinnate or bipinnate leaves with circinate
vernation; very similar to the Polypodiopsida or
leptosporangiate ferns
o sporangia
-located on the abaxial surface of leaflet blades
-eusporangia; synangium: sporangia fused in a
common structure (in some taxa)
o gametophyte
-photosynthetic
o Sporangia : eusporangiate
Apomorphy of Marattoid ferns
o “polycyclic siphonostele”
-appears as concentric rings of siphonosteles in
cross-section
MarattiaceaeF
mucilage canals
o stems – rhizomatous or erect, stout,
and trunk-like, the anatomy a polycyclic
dictyostele
o leaves – develop by circinate vernation;
mature leaves large; stipulate: stipules
large, persistent on stems; simple or 1–
4-pinnate
o petioles and rachillae — with swollen
pulvini and prominent pneumatodes
(lenticels)
o sori – abaxial, intramarginal, exin-
dusiate
o sporangia – eusporangiate;
homosporous; distinct; vertically
dehiscent: plane of dehiscence
perpendicular to blade surface;
 arranged in a narrow ring or fused into
a raised or sunken synangium that
dehisces tangentially into two valves
o spores – trilete, monolete, or alete
o gametophytes – large, thalloid, and
photosynthetic
o Economic importance: cultivated ornamentals
and food (from edible stems), perfume oil, and
an alcoholic drink (from stem starch).
o 6 genera: Angiopteris, Christensenia (2 spp.),
Danaea (30 spp.), Eupodium (neotropics),
Marattia s.s., and Ptisana (paleotropics)
o distinctive in being large, terrestrial ferns with
mucilage canals; the stems with a polycyclic dic-
tyostele; the leaves generally large, simple to
several-pinnate, with abaxial, intramarginal
eusporangia, sometimes fused into synangia
POLYPODIOPSIDA - LEPTOSPORANGIATE FERNS
o also known as Leptosporangiatae or Filiopsida or
Filicales
o commonly known as Leptosporangiate ferns
o sporophytes – perennial herbs or trees (almost all
Leptosporangiate ferns)
o (exception) aquatic ferns: annuals
o Stem
-horizontally oriented
-rhizome: may grow under or upon the ground
(terrestrial), on or in cracks of rocks (epipetric), on
or in water (aquatic), or upon another plant
(epiphytic).
o arborescent: with a tall, erect, aerial stem, which in
the tree ferns (some species)
o a few ferns are vines with weak stems or with
elongate, vine-like leaves that sprawl on the ground
or upon another plan
o stem anatomy – ectophloic or amphiphloic
siphonostele/solenostele dictyostele or protostele
o leaf development
- circinate vernation: both major and minor axes or
leaf divisions are coiled early in development and
uncoil at maturity
o coiled heads: fiddleheads or croziers
o leaf: frond; petiole: stipe
o pinnae/pinna
-first discrete leaflets or blade division of ferns
-if there are 1 or more divisions: 1° pinna, 2° pinna,
and so forth are be used
o pinnules or segments: ultimate leaflets or blade
divisions
o monomorphic: leaves of ferns are similar to
another
o dimorphic: fertile leaves differ from sterile,
vegetative leaves
o Leaf venation
-pinnate: with a central vein giving rise to veinlets
on either side; overalls leaf venation of ferns
-palmate: less common; with more than one main
vein arising from the base
-ultimate vein pattern: pinnules (smallest segments
in the leaf)
*2 types of ultimate venation
1. Open/Free: veins arising from the midvein or
base of a pinnule do not join back together
-free veins maybe simple (not branching)
-forked / bifurcate (more common) ; each
vein gives rise to pairs of veins
a. dichotomous (equal forks)
1. Reticulate/Anastomosing: veins appear to join
back together forming a net-like reticulum
enclosing an area called areole
o Trichomes: hairlike structures
o Scales: flattened, minute, leaf-like structures on
the stem, shoot apex, petiole or blade
1. clathrate- scales with cell walls of
adjacent cells (anticlinal walls) that are
thick
2. non-clathrate- thin anticlinal walls
a. fibrillose
b. denticulate
c. marginate
o Primary apomorphy: leptosporangium
-developing from a single cell
-having a single layer of cells
-smaller spore # that eusporangia
o Leptosporangia
-have a proximal stalk and distal sporangial
body
-annulus - single row of specialized cell; cell
walls are differentially thickened on the inner
cell face and on the faces between adjacent
annular cells
o Sori/sorus- clusters of leptosporangia
o Receptacle- common region of attachment of
sorus
o Paraphyses- sterile, hair-like structures ; arising
from the receptacle
o Indusium
-flap of tissues arising from the blade
surface
-function to protect young leptosporangia
-control the dispersal of spores
 Examples: Aspleniaceae are indusiate;
Polypodiaceae are exidusiate
b. Shapes :
a. reniform - kidney-shaped
b. orbicular - circular
c. linear - narrowly elongate
c. Types of attachments :
a. Peltate - with a central stalk
b. Lateral - attached to the side
o Acrostichoid - sori are scattered on the leaf
surface
o False indusium - possessed by those who lack
indusium ; extension of the blade margin which
overlaps the sorus
o Development of Leptosporangia within the
sorus can be :
a. Gradate (sequential) - the sporangia of a
sorus mature in succession from the
base toward the apex (acropetalous ) or
from the apex toward the base
(basipetalous)
b. Simultaneous (simple) - sporangia or a
sours mature at the same time
c. Mixed - combination of gradate and
simultaneous
d. Intermingled - no consistent
developmental pattern
o Haploid gametophyte phase is separate from
the “dominant” sporophyte phase
o Fern gametophytes
a. small, thin flat sheet of green,
photosynthetic cell, cordate in shape
b. typically surficial (grow upon the
ground) and bear rhizoids (attachment
and absorption)
c. bear sperm-producing antheridia and
egg producing archegonia
d. sperm cells are coiled and
multiflagellate ; attached to a
cytoplasmic vesicle
o Sperm fertilizes an egg in the cell of
archegonium
o Zygote divides and differentiates into a new
embryo ; remains attached to the gametophyte
o Soon sporophyte attains independence of the
gametophyte (dies) and the sporophyte
becomes persistent (dominant phase)
o Fern sporophytes can develop from
gametophytes asexually termed apogamy /
apogamous
o Economic importance : cultivated ornamentals
o Some ferns (croziers) are edible and used as
medicines, fibers and matting or flavoring
o Leptosporangiate fern classification : 7 orders
and 33 families
OSMUNDALES - OSMUNDACEOUS FERNS
o Cinnamon Fern family
o 3 genera, 20 species
o Terrestrial plants
o Distinctive in having erect stems (sometimes
arborescent), an ectophloic siphonostele with
separate xylem strands, dimorphic leaves or leaf
segments with sori and indusia absent, and
short-stalked, large bodied sporangia with an
apical slit and a lateral annulus
o Stems : erect with an ectophloic siphonostele
having a ring of discrete xylem strands
o Leaves : 1-2 pinnate or pinnatifid, stipulate,
with stipules at the base of petioles, and
dimorphic
o Sori and indusia : absent
o Sporangia ocurring on the abaxial surface of
leaves or leaf segments
o Sporangia : large bodies and shirt stalks,
dehiscing by apical slit
o Annulus : lateral
o Spores : green, subglobose, and trilete, 128-512
per sporangium
o Gametophytes : relatively large, green, cordate,
surficial
o Chromosome # : 22
o Distribution : tropical and temperate region
o Economic importance : cultivated ornamentals ;
local uses for fiber and food
o Osumunda regalis : used in brewing Celtic ale
HYMENOPHYLLALES - FILMY FERNS
o Greek hymen - “membrane” + phyllum - “leaf”
o 9 genera , 600 species
o contain both epiphytic and terrestrial plants
o distinctive in having scale-less, usually slender,
creeping, protostelic rhizomes, thin leaves
usually 1-cell thick, lacking stomata, and
marginal sori with conical, tubular, or 2-lobes
indusia and elongate receptacle
o Stems : rhizomatous, rhizomes withput scales,
slender and creeping or stout and erect,
protostelic
o Leaves : 1-cell thick, stomata absent, cuticle
absent or reduced
o Blade scales : absent
o Trichomes : sometimes present
o Venation : open
o Soria : marginal
o Receptacle : elongate and continuous with vein
tips
o Indusia : present and conical, tubular or 2-lobed
(bivalvate)
o Sporangia : basipetalous
o Annulus : oblique
o Spores : green, globose, and trilete
o Gametophytes : filamentous or ribbon-like,
reproducing by fragmentation or gemmae
o Chromosome # : 11, 12, 18, 28, 32, 33, 34, 36
o Distribution : humid regions of pantropical and
south-temperate regions
o Contains 2 sister groups : “trichomanoid” clade
and “hymenophylloid” clade
o Family members are found in humid regions
o Economic importance : few cultivated
ornamentals
GLEICHENIALES - GLEICHENOID FERNS
o contains 3 families
o Forking fern family
o terrestrial, often growing in dense open
thickets, dichotomously branched, with a
“vitalized” protostele or rarely a solenostele
o distinctive in often forming dense, open
thickets, the leaves usually long, indeterminate,
pseudodichotomously branched, with round
exindusiate sori, leptosporangia with a
transverse-oblique annulus
o Leaves : indeterminate with the rachises
pseudodichotomously branched, 1-2 pinnate
o Veins : free
o Sori : round, abaxial, not marginal, exindusiate
o Sporangia: round to pear-shaped, 5-15
sporangia per sorus developing simultaneously
o Annulus : transverse-oblique
o Spores : globose-tetrahedral or bilateral, trilete
or monolete, 128-800 per sporangium
o Gametophytes : large, green, surficial, with
club-shaped hairs, developing endotrophic
mycorrhizae
o Chromosome # : 22, 34, 39, 43, 56
SCHIZAELES - SCHIZAEOID FERNS
o contain3 families : Anemiaceae, Lygodiaceae,
and Schizaeaceae
o common morphological features : dimorphic
leaves, lack of well-defined sori, and sporangia
having a transverse, subapical, continuous
annulus
o Lygodiaceae
-climbing fern family
-terrestrial, climbing plants
o distinctive in having indeterminate leaves, with
twinning/climbing rachises, alternately bearing
pseudodichotomously-branching pinnae, leaf
segments dimorphic, the sori at the tips of ultimate
segments, each with only one sporangium covered
by indusium-like flap, sporangia with a transeverse,
subapical, continuous annulus
o Stems : rhizomatous, rhizomes slender, creeping,
bearing hairs, and protostelic
o Leaves : mostly indeterminate with and elongate,
twinning and climbing rachis that bears pinnae
alternately
o Pinnae : pseudodichotomously branching
o Veins : free or anastomosing
o Sori : abaxial, on lobes of ultimate leaf segments, an
o Indusium : indusium-like flange covering the
sporangium
o Sporangia : 1 per sorus
o Annulus : transverse, subapical, and continuous
o Spores : tetrahedral, trilete, 128-256 pero
sporangium
o Gametophytes : green, cordate, and surficial
o Chromosome # : 29, 30
o Distribution : pantropical
o Economic importance : use of the twinning leaf
rachis as a fiber / mat material
SALVINIALES - AQUATIC / HETROSPOROUS FERNS
o 2 families : Marsileaceae and Salvinaceae
o unusually being aquatic
o unique among leptosporangiate ferns in being
heterosporous, producing 2 types of spores and
sporangia : megaspores, produced within
megasporangia and microspores forming within
microsporangia ; both developed and formed in
sporocarp
o Sporocarp : rounded, seed-like structure with a
hard outer layer ; for protection and dormancy
o Megaspores : large and produced in few #s
-single, haploid nucleus or megaspores gives -
rise to female gametophyte via mitosis and
differentiation
-female gamtetophyte bears 1 or more
archegonia
o Microscpores : small and produced in large #s
-develop into a male gametophyte, which bears
1 or more spem-producing antheridia
o Reproductive features:
-heterospory - reduction of meagspore # per
megasporangium
-endospory - in the evolution of seeds
MarsileaceaF
o clover fern family
o 3 genera and 75 species
o consist of rooted, aquatic, herbs with emergent
leaves, the blade sometimes floating
o Stems : elongate, slender and creeping
rhizomes often bearing hairs with aerenchyme
and solenostelic anatomy
o Leaves : circinate, simple or palmate with 2 or 4
sessile leaflets
o Veins : dichotomous often fusing apically
o Sporocarps : reniform with a stlak arising from
the petiole base of leaf axil, each sporocarp
bearing 2 halves, each of these with a several
rows or internal sori
o Sori : consist of column of megasporangia and
microsporangia that lacks an annulus and are
enveloped by a hood-like indusium
o Sporophore : gelatinous structure where sori
are attached
o Megasporangium : bears a single, trilete
meagspore
 after imbibing water megaspore
releases a gelatinous mass -
acrolamellae
 Acrolamellae - with apical longitudinal
folds and basal horizontal folds,
contains a central liquid, filled region
(spearm lake)
 Megasporangial wall breaks away ,
female gametophyte forms a single
archegonium at the megaspore apex,
rupturing the apex of the spore wall
and protruding in the sperm lake
o Microsporangia - produce several trilete
microspores, each microspore forming an
endosporic, male gametophyte that bears and
relase numerous coiled, multiflagellate sperm cells,
some of which enter the opening of the
acrolamellae into the sperm lake region, which
leads to archegonium
o Chromosome # : 10 (pilularia ), 20 (marsilea)
o Distribution subcosmopolitan
o distinctive in being rhizomatous, aquatic ferns, the
leaves lacking blade tissue or palmate with 2 or 4,
sessile leaflet, sori developing within seed-like,
dessication-resistant sporocarps, which, upon,
imbibing water, each release an elongate,
gelatinous sporophore bearing the sori and
sporangia, the spores heterospores
SalviniaceaeF
o floating fern family
o 2 genera , 16 species
o consist of floating, aquatic herbs
o Roots : absent in Salvinia present in Azolla
o Stems : dichotomously branched rhizomes,
protostelic, aerenchymous
o Leaves : simple, dimorphic, aerenchymatous, at
maturity distichous
o Sporocaprs : globose, heterosporous, each bearing
either one megasporangium or microsporangium
 Megasporangium - one functional megaspore
surrounded by massulae (gelatinous mass from
multinucleate plasmodium)
 Each megaspore forming an endosporic,
female gametophyte with several, protruding,
apical archegonia
 Microspornagium - each bearing several
microspores surrounded by massulae
o Chromosome # : 9(Salvinia), 22 (Azolla)
o Distribution : subcosmopolitan
o distinctive in being floating, aquatic herbs, the
leaves simple, either in whorls of 3 (2 floating, 1
rootlike) bearing water repellant trichomes
(Salvinia) or 2-ranked and 2-lobed, the upper lobes
housing cyanobacteria (Azolla), sori modified as
sporocarps, each bearing either one megaspore or
several microspore surrounded by gelatinous
massulae
CYATHEALES - TREE FERNS
o 600 species, 8 families
o Stems : arboresecnet or rhizomatous bearing hairs
or prominent scales
o Sori : marginal or abaxial, indusiate or exindusiate
o Spores : trilete
o Gametophytes : green and cordate
CyatheaceaeF
o scaly tree fern family
o 4 genera
o mostly terrestrial, some epiphytic
o Stems : mostly arborescent (tree ferns); anatomy is
polycyclic dictyostele
o Trunks : often with marcescent (persisting) leaves of
leaf bases
o Shoot apices and petiole bases are covered with
large scales or hairs
o Leaves: up to 5m long
o Blades: 1-3 pinnate
o Petioles: obvious, usually discontinuous
pneumathodes (tissue with air spaces) in 2 ines
o Blade veins: free, simple to forekd rarely
anastomosing
o Sori : abaxial, round, superficial or terminal on veins
and marginal or submarginal
o Receptacle: raised
 o Paraphyses : present
o Exindusiate or indusiate
o Indusium : present; saucer-like, cup-like,
bivalvate, or globose, and completely
surronding sporangia
o Sporangia : mature gradately
o Annulus : oblique
o Spores : tetrahedral, trilete, variously
ornamented
o Gametophytes : green and cordate
o Chromosome # : 69
o Distribution : pantropical, especially in montane
forests
o distinctive in being mostly arborescent, the
shoots generally covered with trichomes or
scales, leaves very large, usually 1-3 pinnate
leaves, the sori exindusiate or indusiate,
indusia, when present, saucer-like, cup-like,
bivalvate or globose and completely
surrounding sporangia, the sporangium annulus
oblique
POLYPODIALES - POLYPOD FERNS
o 7,100 species, 15 families
o indusiate or exindusiate
o sporangia are distinctive in having a thin (1-3
cellled), generally long stalk, a lateral stomium and
an annulus that is vertically oriented and
interrupted by the stalk and stomium
o Gametophytes : green, cordate and surficial
o Aspleniaceae - Spleenwort family
 Stems : rhizomatous, rhizomes climbing, creeping,
ascending or suberect, bearing clathrate scales at
shoot apices and petiole bases
 Leaves : monomorpihc, simple to multipinnate,
often with small clavate hairs, 2 back to back C-
shaped vascular strands at petiole bases fusing
distally into an X-shape
 Venation : pinnate or forking, usualy free, without
veinlets
 Sori and indusia : elongate (linear) along veins
 Sporangia ; mixed, sporangial stalks of 1 row long
 Spores : reniform and
monolete, with a winged
perine
 Chromosome # : 36, rarely 38,
39
 Distribution :
subcosmopolitan
 Economic importance :
medicine and cultivars such as
Asplenium bulbiferum
 distinctive in having shoot
apices with clathrate scales and
leaves with elongate, linear sori
and indusial
DRYOPTERIDACEAE - DRYOPTEROID FAMILY
o Greek drys - “oak” + pteris - “fern”
o 40-45 genera, 1700 species
o terrestrial, epipetric or epiphytic perennials
o Stems : rhizomatous, the rhizomes dictyostelic,
creeping, ascending, erect, or scandent-
climbing, shoot apices with non-clathrate scales
o Leaves : monomorphic, rarely dimorphic, simple
or pinnate to decompound
o Veins : pinnate or forking, free to anastomosing,
with or without veinlets
o Sori : usually round, indusiate or exindusiate,
acrostichoid in some taxa
o Indusia : round-renform or peltate
o Sporangia : mixed, sporangial stlaks in 3 rows,
short to long
o Spores : reniform and monolete
o Chromosome # : x = 41 (40)
o Distribution : pantropical to temperate regions
o Economic imp : cultivated ornamentals
o distinctive in being rhizomatous, creeping to
climbing plants, the shoot apcies with non-
clathrate scales, sori exindusiate or indusiate
with indusia round-reniform or peltate
o POLYPODIACEAE
o Polygram / Polypody family
o Greek Polys - “many” + pous -“feet”
o 56 genera, 1200 species
o epiphytic (usually), epipetric or terrestrial
perennials
o Stems : rhizomatous, rhizomes dictyostelic, long
to short-creeping, bearing scales
o Leaves : simple to 1-pinnate, monomorphic or
dimorphic
o Blades : glabrous or with hairs or scales
o Veins : often anastomosing or reticulate
sometimes with veins or free
o Sori : abaxial, round, oblong, or elliptic, rarely
elongate or acrostichoid
o Receptacle : often with paraphyses, exindusiate
o Sporangia : mixed, sporangial stalks n 1-3 rows,
often long
o Spores : hyaline to yellowish, reniform and
monolete or greenish and globose-tetrahedral
and trilete
o Chromosome # : 35, 36, 37, (25, etc)
o Distribution : pantropical to temperate
o Economic imp : edible, medicinal, or flavoring
plants, # of ornamental cultivars inclding
species of Aglaomorpha, Drynaria and
Platycerium
o include the so-called grammitid ferns about 20
gener and 600 species of mostly small, tropical
epiphytes with simple leaves ; nested within
Polypodiaceae
o distinctive in being exindusiate, mostly
epiphytic ferns, usually round, oblong, or
elliptic, rarely elongate or acrostichoid

PTERIDACEAE PTEROID FERN FAMILY

o Greek pteris - “fern”
o 50 genera, 950 species
o terrestrial, epipetric, or epiphytic plants, rarely
aquatic
o Stems : rhizomatous, the rhizomes creeping to
erect, bearing scales or hairs
o Leaves : simple, pinnate, pedate or
decompound
o Veins : free or anastomosing
o Sori : exindusiate, either marginal with a false
indusium formed by a reflexed marginal flap or
intramarginal in lines along veins
o Receptacle : not raised
o Sporangia : mixed, sporangial stalks 1-3 cell
thick, often long
o Spores : globos or tetrahedral, trilete and
ornamented
o Chromosome # : 29, 30
o Distribution : subcosmopolitan, mostly tropical
and arid regions
o Economic imp : cultivated ornamentals, such as
Acrostichum, Adiantum, Cheilanthes,
Cryptogramma etc ; Pteris vittata used to
remove arsenic from toxic landfills
o 5 monophyletic groups : Parkerioideae,
Adiantodeae, Cryptogramms, Cheilantoideae,
Pteridoideae
o distinctive in having exindusiate sori, either
marginal with false indusia, or intramarginal in
lines along veins.

***DISCLAIMER: This reviewer is book-based and

does not contain information/notes from the
powerpoints given by the professor. Also, for
diagrams and images, please refer to your book.