Vascular plants evolved several key adaptations including:
1. An independent sporophyte generation that is long-lived and photosynthetic, while the gametophyte generation is short-lived.
2. A branched sporophyte axis that bears multiple sporangia and reproductive organs. This allowed for greater height and structural support.
3. Lignified secondary cell walls in sclerenchyma, tracheary elements, and sieve elements which provided strength, rigidity and conduction of water, minerals and sugars throughout the plant.
Vascular plants evolved several key adaptations including:
1. An independent sporophyte generation that is long-lived and photosynthetic, while the gametophyte generation is short-lived.
2. A branched sporophyte axis that bears multiple sporangia and reproductive organs. This allowed for greater height and structural support.
3. Lignified secondary cell walls in sclerenchyma, tracheary elements, and sieve elements which provided strength, rigidity and conduction of water, minerals and sugars throughout the plant.
Vascular plants evolved several key adaptations including:
1. An independent sporophyte generation that is long-lived and photosynthetic, while the gametophyte generation is short-lived.
2. A branched sporophyte axis that bears multiple sporangia and reproductive organs. This allowed for greater height and structural support.
3. Lignified secondary cell walls in sclerenchyma, tracheary elements, and sieve elements which provided strength, rigidity and conduction of water, minerals and sugars throughout the plant.
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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