BryophytesBryophytes

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BryophytesBryophytes UbiquitousUbiquitous

Moist temperate and Moist temperate and tropical locationstropical locations

Dominate Arctic TundraDominate Arctic Tundra AntarcticaAntarctica Above the treelineAbove the treeline Dry desertsDry deserts Some aquaticSome aquatic

Important colonizers Important colonizers with lichenswith lichens

Sensitive to air Sensitive to air pollutionpollution

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HypnumHypnum sp. - Moss on fallen log. GSMNP sp. - Moss on fallen log. GSMNP

HypnumMOSS

NowelliaLEAFY LIVERWORT

Lichen

Photo by Alan Heilman

Bartramia Bartramia pomiformipomiformi

(apple moss) + (apple moss) + PeltigeraPeltigera, a lichen., a lichen.

Photo by Alan Heilman

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Multicellular specialization (tissues)Cell walls made of cellulosePigments:

a. Chlorophyll ab. Chlorophyll b

Primary storage substance: starchOogamous -- Male gamete is motile & asymmetricalBreakdown of nuclear membraneZygotes retained

Features shared with Charophytes

Other algae

Xylem and phloem

Outer layer on sporesStomataAbility to distinguish D-methionine

from L-methionine

Zygote retained by gametophyte

Flavenoids

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BryophytesBryophytes Transitional between Charophytes and Transitional between Charophytes and

Vascular PlantsVascular Plants Synapomorphies for land plantsSynapomorphies for land plants

Antheridia and archegonia with protective Antheridia and archegonia with protective layerlayer

Retention of zygote & multicellular embryoRetention of zygote & multicellular embryo Presence of multicellular diploid Presence of multicellular diploid

sporophytesporophyte Multicellular sporangiaMulticellular sporangia Meiospores with sporopolleninMeiospores with sporopollenin Apical meristemApical meristem

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BryophytesBryophytes ““Autapomorphies”Autapomorphies”

Lack of xylem and phloemLack of xylem and phloem No ligninNo lignin Gametophyte generation dominant; Gametophyte generation dominant;

free-livingfree-living Sporophyte nutritionally dependantSporophyte nutritionally dependant Sporophyte unbranched with single Sporophyte unbranched with single

sporangiumsporangium

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10Ubc.edu

Bryophyte StructureBryophyte Structure Thalloid -- Hornworts & (some) liverworts Thalloid -- Hornworts & (some) liverworts Differentiated -- Mosses & (some) Differentiated -- Mosses & (some)

liverwortsliverworts Not Not truetrue leaves/stems leaves/stems

o Gametophyte generationGametophyte generationo No xylem & phloemNo xylem & phloem

RhizoidsRhizoids Mosses – multicellularMosses – multicellular Liverworts & hornworts – unicellularLiverworts & hornworts – unicellular Only anchor plantOnly anchor plant

PlasmodesmataPlasmodesmata

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Bryophyte StructureBryophyte Structure Hornworts – single plastidHornworts – single plastid Sperm only flagged cell – water Sperm only flagged cell – water

requiredrequired Gemmae cupsGemmae cups Gametophytes oftenGametophytes often

sexedsexed Sex chromosomes 1Sex chromosomes 1stst

in bryophytes!in bryophytes!

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Photo by Alan Heilman

Reproductive StructuresReproductive Structures AntheridiumAntheridium

Each cell Each cell single single spermsperm

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Reproductive StructuresReproductive Structures ArchegoniaArchegonia

Single eggSingle egg Zygote persistentZygote persistent Matrotrophy Matrotrophy Enlarged venter Enlarged venter

calyptracalyptra

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Nearly Mature Nearly Mature SporophyteSporophyte

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EmbryophytesEmbryophytes Matrotrophy + placenta Matrotrophy + placenta multicellular multicellular

sporophyte (produced by ……sporophyte (produced by …… More divisions between fertilization and More divisions between fertilization and

meiosis meiosis Larger sporophyteLarger sporophyte More sporesMore spores

Sporophytes of hornworts & mosses Sporophytes of hornworts & mosses stomatastomata

Sporopollenin – survival valueSporopollenin – survival value Spore generates Spore generates protonema protonema

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The three phyla are:

• Bryophyta – Mosses• ca. 10,000+ sp.

• Hepatophyta – Liverworts

• ca 6000 sp.• Leafy liverworts• Thallose liverworts

• Anthophyta – Hornworts

• ca 100 sp.

THE PHYLA OF THE PHYLA OF BRYOPHYTESBRYOPHYTES

Liverworts: Phylum Liverworts: Phylum HepatophytaHepatophyta

Ca. 6000 speciesCa. 6000 species Mostly terrestrialMostly terrestrial Some have protonema-like structureSome have protonema-like structure Apical meristemApical meristem 3 major groups3 major groups

Complex thalloid liverwortsComplex thalloid liverworts Simple thalloid liverwortsSimple thalloid liverworts Leafy liverwortsLeafy liverworts

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Complex Thalloid Complex Thalloid LiverwortsLiverworts

Marchantia, Riccia, Marchantia, Riccia, RicciocarpusRicciocarpus

TerrestrialTerrestrial Thallus 10-30 cells thickThallus 10-30 cells thick

Upper layer -- air poresUpper layer -- air pores Middle layer -- chloroplast Middle layer -- chloroplast

richrich Lower layer – Lower layer –

carbohydrate storage & carbohydrate storage & rhizoidsrhizoids

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RicciaRiccia sporophytes sporophytes

Little more than a sporangiumLittle more than a sporangium

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MarchantiaMarchantia sporophytes sporophytes Gametangia -- stalked specialized Gametangia -- stalked specialized

structuresstructures Sporophyte – foot, seta, and capsuleSporophyte – foot, seta, and capsule

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Conocephalum Conocephalum conicumconicum - thallus + - thallus + archegoniophore, archegoniophore,

with dev. sporophytewith dev. sporophyte

archegoniophore (part of gametophyte, not a sporophyte)

Photo by Alan Heilman

MarchantiaMarchantia

Mature capsule contains elaters Mature capsule contains elaters (green strands)(green strands) Aid in spore dispersalAid in spore dispersal

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Asexual ReproductionAsexual Reproduction

Fragmentation Fragmentation Gemmae – splash cupsGemmae – splash cups

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MarchantiaMarchantia Life Cycle Life Cycle

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Marchantia polymorphaMarchantia polymorpha - antheridiophores - antheridiophores malemale

Photo by Alan Heilman

Marchantia Marchantia polymorphapolymorpha - -

archegoniophores archegoniophores femalefemale

Photo by Alan Heilman

Ricciocarpus natansRicciocarpus natans - floating aquatic - floating aquatic liverwortliverwort

Photo by Alan Heilman

Riccia fluitansRiccia fluitans - floating aquatic liverwort - floating aquatic liverwort

Photo by Alan Heilman

Conocephalum conicumConocephalum conicum - detail of upper - detail of uppersurface of thallus with poresurface of thallus with pore

Photo by Alan Heilman

Leafy LiverwortsLeafy Liverworts Very diverseVery diverse

> 4000 of the 6000 > 4000 of the 6000 speciesspecies

More tropicalMore tropical Diagnostic …Diagnostic …

Moss leaves Moss leaves o SpiralSpiralo MidribMidribo Entire Entire

Liverwort leaves Liverwort leaves o Distichous Distichous o Highly dissectedHighly dissected

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FrullaniaFrullania

Photo By Paul Davison

CephaloziaCephalozia

Photo By Paul Davison

Moss and Liverwort Moss and Liverwort LeavesLeaves

Moss Leafy Liverwort

Mid Vein

Margin teeth

Sporophyte of leafy Sporophyte of leafy liverwortliverwort

Capsule

Seta

Photo by Alan Heilman

Hornworts: Phylum Hornworts: Phylum AnthocerophytaAnthocerophyta

ca 100 speciesca 100 species Gametophytes similar Gametophytes similar

to liverwortsto liverworts Large single Large single

chloroplastchloroplast Phylogenetically most Phylogenetically most

closely related to closely related to vascular plantsvascular plants

NostocNostoc is symbiont -- is symbiont -- NN

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GametophytesGametophytes

Rosette-likeRosette-like Unisexual and Unisexual and

bisexualbisexual Clustered Clustered

antheridiaantheridia Single gametophyte Single gametophyte

– many sporophytes– many sporophytes

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wikipedia

SporophytesSporophytes

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SporophytesSporophytes

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Life cycleLife cycle

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Anthoceros laevisAnthoceros laevis - - hornwort, habitat hornwort, habitat

viewview

Phaeoceros sp.Phaeoceros sp.Photo by Ken McFarland

Mosses: Phylum Mosses: Phylum BryophytaBryophyta

Things that are Things that are NOTNOT mosses …. mosses …. Reindeer moss – lichenReindeer moss – lichen Scale moss – leafy liverwortsScale moss – leafy liverworts Club moss – Club moss – LycopodiumLycopodium (Fern ally) (Fern ally) Spanish moss – bromeliad (like Spanish moss – bromeliad (like

pineapple …..)pineapple …..) Spike moss – Spike moss – SelaginellaSelaginella (Fern ally) (Fern ally) Sea moss (or Irish moss) – red alga Sea moss (or Irish moss) – red alga

(Carrageenan!)(Carrageenan!)

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3 groups discussed in text3 groups discussed in text Sphagnidae – peat mossesSphagnidae – peat mosses Andreaeidae – granite mossesAndreaeidae – granite mosses Bryidae – true mossesBryidae – true mosses

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Mosses: Phylum Mosses: Phylum BryophytaBryophyta

Peat Mosses – Peat Mosses – Sphagnidae Sphagnidae

Sphagnum Sphagnum Earliest diverging mossEarliest diverging moss UnusualUnusual

Protonema a “plate” w/ marginal meristemProtonema a “plate” w/ marginal meristem Distinctive leaf morphologyDistinctive leaf morphology Distinctive sporophytesDistinctive sporophytes Gametophytes -- clusters of branchesGametophytes -- clusters of branches

Diapers and wound dressingsDiapers and wound dressings 20x dry weight in water (cotton only 4-6x)20x dry weight in water (cotton only 4-6x)

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SphagnumSphagnum Unique leaf Unique leaf

morphologymorphology Antheridia and Antheridia and

archegonia at tips archegonia at tips of specialized of specialized branchesbranches

Unique Unique sporophyte sporophyte morphologymorphology

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Granite Mosses -- Granite Mosses -- AndreaeidaeAndreaeidae

Mountainous or Arctic regionsMountainous or Arctic regions Protonema & rhizoids – 2 rows of Protonema & rhizoids – 2 rows of

cellscells Unusual sporangial dehiscenceUnusual sporangial dehiscence

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True Mosses -- BryidaeTrue Mosses -- Bryidae

Filamentous protonemaFilamentous protonema Multicellular rhizoidsMulticellular rhizoids Leaves one cell thickLeaves one cell thick Specialized tissuesSpecialized tissues

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Hadrom & LeptomHadrom & Leptom Found in setaFound in seta Water conducting tissue – hydroidsWater conducting tissue – hydroids

Similar to tracheids no protoplast Similar to tracheids no protoplast butbut no no ligninlignin

Food conducting tissue – leptoidsFood conducting tissue – leptoids Similar to sieve tubesSimilar to sieve tubes

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Sexual ReproductionSexual Reproduction Male and female gametangiaMale and female gametangia

Branch tipsBranch tips Lateral branchesLateral branches Unisexual & bisexualUnisexual & bisexual

o Antheridia in splash cupsAntheridia in splash cups Capsules – 6-18 months to mature!Capsules – 6-18 months to mature!

Maternally nourished sporophyteMaternally nourished sporophyte Specialized spore dispersalSpecialized spore dispersal

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Sexual ReproductionSexual Reproduction

Mnium

Archegonia and Antheridia

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paraphyses

Antheridial Splash CupsAntheridial Splash Cups

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Peristome TeethPeristome Teeth

Control release of Control release of

~ 50 million ~ 50 million sporesspores

Characteristic of Characteristic of BryidaeBryidae

Taxonomically Taxonomically importantimportant

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Eurhynchium serrulatumEurhynchium serrulatum

calyptra

operculum

peristome teeth

Photo by Alan Heilman

Dicranella Dicranella variavaria

Dehiscing operculum

Photo by Alan Heilman

Peristome teeth of Peristome teeth of Dicranum Dicranum flagellariaflagellaria

Photo by Alan Heilman

Dicranella Dicranella varia varia sporesspores

Photo by Alan Heilman

GrimmiaGrimmia gracilisgracilis

peristome peristome teeth openteeth open

Photo by Alan Heilman

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