CHAPTER 4 o secondary wall

Evolution and Diversiy -secreted outside of plasma membrane &

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

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