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Home , Andreaeaceae, Blasiales, Fossombroniales, Grimmiales, Notothyladaceae, Pelliales

Phylogeny Poster Systematics and Characteristics of Nonvascular Land (, Liverworts, )

Bryophyte

Phylogeny anacrogynous; lvs in three rows (2 lateral, succubous, 1 dorsal lobule) Poster oil bodies scattered; mucilage on ventral surface CS parenchymatous, with glomerophycotean fungus di- or monoicous; single S per gynoecium; gemmae in axils of dorsal lobules Treubiales axis subterranean; lvs mostly isophyllous; rhizoids – Tracheophyte shoot calyptra + CS +, cells thin-walled, perforated Phylogeny di- or monoicous; gametangia lateral, bracts –; seta massive blepharoplast: lamellar strip and spline < 90 microtubules, aperture on left side; S several/gynoecium Poster CAP 4-valved; walls unistratose; elaterophore basal; elaters filamentous; asex. repro. – Haplomitriales Haplomitriaceae thalli winged ("leafy"), 2 ventral scale rows; air chambers –; gametangiophores – ventral "auricles" with Nostoc; dioicous; AN dorsal, solitary; AR dorsal, behind apex Angiosperm blepharoplast: marchantialean; CAP 4(-6)-valved elaters 2-helical; elaterophore basal, rudimentary Phylogeny gemmae receptacles flasked-shaped (unique in liverworts) Poster air chambers +; chlorophyllose filaments – rhizoids smooth ventral scales +, appendages – Marchantiidae archegoniophores branched Liverworts gemmae Neohodgsoniales Neohodgsoniaceae thalli rosettes or stems; axes: winged or lobes leaf-like air chambers –, mucilage cells –, pores – thallose or foliose AR and S in pear-shaped involucres (dorsal on thallus) rhizoids + rhizoids +, smooth oil bodies + seta very short; CAP cleistocarpous; elaters – Monocarpaceae Riellaceae perforated water-conducting cells M mycothallus with endophytic Glomeromycota thallus differentiated; air chambers +; air pores +/–; chlorophyllose filaments + a storage parenchyma +; ventral scales +; rhizoids +, pegged or smooth gametangia protective structures + di- or monoicous; antheridiophores +, archegoniophores +, AR involucre +/– Aytoniaceae Cleveaceae Conocephalaceae Corsiniaceae gametangial ontogeny without apical cells r pseudoperianths usu. –; seta short Cyathodiaceae Dumortieraceae Exormothecaceae Lunulariaceae blepharoplast: plastid and associated posterior elaters usu. 2-3-helical or –; gemmae, occ. tubers mitochondrion positioned at cell terminus c Marchantiaceae Monosoleniaceae Oxymitraceae Ricciaceae Targioniaceae zygote division transversal: : thallus; branching pseudodichotomous; di- or monoicus epi- and hypobasal cells h AN individual in covered chambers (dorsal); AR in distinct groups (dorsal); involucre +, short-tubular or flaplike CAP without columella CAP with 4 valves; elaterophores basal; asex. repro. very rare Noterocladaceae: thalli leafy; lvs succubous; branching lateroventral; AN in ostiolate chambers; AR clustered, involucre + elaters (unicellular) a CAP spheroidal; seta <10 cm; spores large (<100 µm), "multicellular", germination endosporic +/–; elaterophores basal Pelliaceae Noterocladaceae stomata – ca. 5,000 spp. n thallose, leaflike lobes succubous, obliquely inserted lunularic acid rhizoids purplish (rar. pale brown) t Pelliidae AN and AR in simple acropetal sequence i S protected by caulocalyx; CAP wall 2-6 stratose; spores sculptured Petalophyllaceae asex. repro. by fleshy stems (stolons), subterranean tubers, endogenous gemmae Allisoniaceae Calyculariaceae Makinoaceae o thallose or leafy, prostrate, erect or dendroid CS +, cells thick-walled, with pores p AN/AR clustered dorsally on thallus S protected by inner involucre or shoot calyptra Pallavicinaceae Hymenophytaceae h seta massive; CAP 2-14 valved, wall bistratose Pallaviciniales Moerckiaceae Phyllothalliaceae Sandeothallaceae y foliose; apical cell with 2 cutting faces (unique in leafy liverworts) lvs bilobed, smaller leaf lobe usu. a complex water sac; trigones very large t evidence of zoophagy; amphigastria – gametangiophores on short lateral-axillary branches, gynoecia partly sterile; perianth elongate seta + a CAP spherical to short-ovoid; germination endosporic; epiphytic (mostly) Pleuroziales Pleuroziaceae thalli linear, winged; branching dichotomous; midrib distinct (Metzgeriaceae) or thalli irregularly or Metzgeriidae pinnately branched (Aneuraceae); CS –; unicellular hairs +/–; oil bodies +/– or very small di- or autoicous; gametangia on short branches arising from midrib or reduced lateral branches thalloid or foliose shoot calyptra +; pseudoperianth –; CAP 4-valved, elaterophore apical lvs from 3 prim. initials elaters 1- or ehelical; asex. repro. – or gemmae/adventive thalli/caducous branches Aneuraceae Metzgeriaceae S branches lateral, exogenous e lvs incubous, unequally 2-/3-lobed, often conduplicate bilobed, lobules often inflated water sacs amphigastria +/–; S enclosed by a perianth and CY or shoot calyptra or stem perigynium t germination endosporic Jubulaceae Frullaniaceae Lejeuneaceae gemmae + (rare); fungal endosymbionts – Lepidolaenaceae Porellaceae Radulaceae a cauloid, 1-3-pinnate, rhizoids scarce lvs incubous, transversely inserted, bilobed (and further subdivided); margins ciliate p oil bodies 15-40/leaf cell; amphigastria +, bilobed, ciliate ** dioicous; gametangiophores apically on shoot; perianth bottle-shaped, perigynium – h CAP wall 4-7-stratose; germination exo-/endosporic Ptilidiales Ptilidiaceae Herzogianthaceae Neotrichocoleaceae Jungermanniidae y branches exo- or endogenous, ventral or lateral; lvs succubous, incubous, or transverse, undivided or variously lobed, occ. conduplicate bilobed, but then usu. with the smaller Acrobolbaceae Anastrophyllaceae Balantiopsaceae t foliose lobe(s), or lobules, dorsal (inflated water sacs rare); amphigastria +/– Calypogeiaceae Cephaloziaceae Cephaloziellaceae lvs from 2 prim. initials S enclosed by a perianth or stem perigynium; spore germination usu. exosporic a gemmae +/–; fungal endosymbionts + Geocalycaceae Gymnomitriaceae Jungermanniaceae Lepidoziaceae Lophocoleaceae Lophoziaceae protonema –; rhizoids –; complex shoot system of rhizomatous axes, erect leafy shoots perforated water-conducting cells Plagiochilaceae Scapaniaceae Trichocoleaceae dioicous; AN 1-2 in lvs axils of (3-)4-lobed lvs; AR solitary and scattered (1-4/shoot) S (only in T. ceratophylla, rar. encountered) terminal, single; stomata on S –; seta persistent CAP twisted, dehisce along single slit; CY +; asex. repro. by deciduous lvs or shoots Takakiales Takakiaceae cladocarpous; main stems with capitulum: central parenchyma, internal cylinder, cortex branches fascicled, rar. –; retort cells +/–; lvs with hyalocysts and chlorocysts dioicous occ. autoicous; AN single, subglobose, long-stalked in lvs axils AR terminal on short branches in capitulum; spore sac dome-shaped spores in tetrads, trilete mark +; CY +; common in mires, i.e., fens and bogs: peat mosses Sphagnales Sphagnaceae Flatbergiaceae Ambuchananiaceae Mosses CS –; lvs cells (rounded-)quadrate; costa +/– homogeneous or – autoicous, rar. syn- or dioicous; AN without specialized cap; seta – CAP elevated by a pseudopodium; valves 4-10, attached at apex spore sac dome-shaped; germination endosporic; CY small, usu. bistratose mostly cool-temp. and trop.-montane; saxicolous Andreaeales CS –; lvs cells (rounded-)quadrate; costa + dioicous; perichaetia developing after fertilization; AN without specialized cap B seta short, massive; CAP valves irregular 4-8, occ. separate at apex r spore sac dome-shaped germination endosporic; CY covering entire CAP Andreaeobryales Andreaeobryaceae y protonema thallose from filamentous chloronema; plants scattered or in lax, pale green turfs; CS –; axillary hairs club-shaped lvs obovate-spathulate to orbicular, margins at base long-ciliate; costa single, broad, homogeneous o aut- or synoicous; seta straight, succulent p CAP erect, symmetric; apophysis long-tapering, swollen, rugose; stomata many; operculum conic; annulus and PS –; CY cucullate asex. repro. by multicellular axillary gemmae Oedipodiales Oedipodiaceae h CS +/–; lvs cells parenchymatous; costa +, homogeneous y autoicous; CAP erect, symmetric, cylindric, stomata +/–; annulus – operculum +; PS of 4 teeth; CY small, mitrate, glabrous; gemmae + () t mostly northern-temp., on various substrates in moist sites Tetraphidales a protonema short-lived or persistent; subterranean root-like "rhizome"; CS of hydroids and leptoids acro- costa complex (Polytrichum-type), often broad, with adaxial lamellae carpous dioicous, rar. autoicous; perigonia often conspicuous (splash cups) PS CAP erect to horizontal, rounded or 2-4-angled; stomata +; PS of 32 or 64 teeth or as bunch of bristles; CAP with epiphragm protonema thallose x 6 (nemato- spores small (up to 60 10 /CAP); CY mitrate or cucullate, hairy to glabrous often covering CAP; largest terrestrial mosses Polytrichales foliose dontous) lvs cells parenchymatous hydroids protonema persistent; G reduced; ♂ of one leaf surrounding single AN rhizoids multicellular lvs ecostate; dioicous CAP usu. asymmetric, flattened on upper side; annulus +; operculum + mycorrhiza – protonema filamentous PS of -type; exostome + (short teeth in 1-4 rings); endostome + (membranaceous); parastome + gametangial ontogeny chloro- and Buxbaumiidae CY small, mitrate or cucullate Buxbaumiaceae with apical cells caulonema Buxbaumiales blepharoplast: plastid and associated protonema short-lived with funnel-shaped flaps; CS – posterior mitochondrion positioned lvs 2(-3)-stratose; costa +; dioicous; perichaetial lvs ciliate above, costa long-excurrent along inner nuclear surface seta very short; CAP asymmetric, stomata phaneropore; annulus +; operculum + occurrence of stray microtubules PS of Buxbaumia-type (exostome – or rudimentary, endostome +, parastome –) Diphysciidae Diphysciaceae CAP with PS and columella CY small, mitrate Diphysciales elaters – protonema short-lived; leafy stems short from subterranean axes; CS – stomata on S arthro- dontous cladocarpous; lvs cells usu. collenchymatous; costa + (in Gigaspermum –) ca. 13,000 spp. par- or synoicous CAP immersed in some; gymnostomous or cleistocarpous; stomata with 2 guard cells igaspermidae G spores large; CY small, mitrate, fugacious; gemmae + Gigaspermales Gigaspermaceae protonema short-lived (rar. persistent); CS +; lvs lamina cells rectangular to hexagonal; costa + (rar. –) aut- or paroicous, rar. syn- or polyoicous CAP immersed in some; symmetric or asymmetric, operculate or rar. cleistocarpous annulus+/–; PS of Funaria-type or – unariidae F CY large, cucullate or mitrate, typically lobed (mitrate-campanulate in encalypts) Funariales Disceliaceae Encalyptaceae Funariaceae CS +; lvs in 8 rows, base sheathing lvs cells in sheath linear, in limb quadrate to hexagonal, mamillose; costa + di- or autoicous; CAP inclined to horizontal, stomata phaneropore annulus +; operculum + immiidae T PS +, exostome of 16 large teeth, endostome of 64 filaments from basal membrane; CY cucullate, often persistent Timmiales Timmiaceae protonema short-lived CS usu. + lvs cells often incrassate, sinuous, glass hairs common; alar cells +/– costa + (Dicranum-type or homogeneous); PS usu. + saxicolous (most) Grimmiales D i protonema short-lived or rar. persistent c CS +/– Bruchiaceae Calymperaceae Catoscopiaceae r lvs cells quadrate, rectangular, or more rar. elongate a * alar cells often differentiated Dicranaceae Ditrichaceae Fissidentaceae n costa + (Dicranum-type, leucobryoid, or reduced); PS +/– Dicranales i d protonema short-lived or rar. persistent Leucobryaceae Rhabdoweisiaceae a CS usu. +; lvs cells often papillose, glass hairs often + e alar cells rar. differentiated costa + or reduced, Pottia-type; PS usu. + (pottioid) or – PS often xerophytes of "harsh environments" Pottiaceae Ephemeraceae Liverworts Pottiales setaphytes haplo- lepideous CS + Mosses lvs cells usu. rhombic; costa +; di- or autoicous some CAP neck often differentiated into broad and colored hypophysis Hornworts clado- carpous PS usu. + coprophytes (mostly), spores fly-dispersed Meesiaceae Splachnaceae Splachnales protonema globular or filamentous (incl. horsetails) CS –; lvs cells often papillose, glass hairs often + Tracheophytes B PPP +; costa usu. –; CAP immersed or exserted ycads C r annulus –; PS usu. –; spore germination exo- or endosporic Ginkgo Hedwigiaceae Helicophyllaceae Rhacocarpaceae y CY cucullate, smooth, glabrous Hedwigiales Ephedra i Welwitschia d stems tomentose; CS +; lvs often sheathed, limb narrow Gnetum lvs cells mamillose or papillose; costa +; di-, syn-, par-, or autoicous Seed a Plants e perigonia occ. splash cups; CAP globose ("apple mosses") ANA grade PS of Bryum-type +/–; annulus –; operculum + Bartramiaceae PS CY usu. minute, cucullate; asex. repro. by deciduous branchlet and bulbils Bartramiales Angiosperms Magnoliids diplo- lepideous- PPP usu. –

Monocots alternate lvs cells rhombic-hexagonal; costa + CAP often pyriform, pendulous ("pear mosses") – Fabids PS +/ (Bryum-type) asex. repro. by bulbils, filiform gemmae and rhizoid tubers Bryales Bryaceae Mniaceae Plagiomniaceae Malvids Rosids some pleurocarpous; CS +

Lamiids lvs cells rounded-hexagonal; costa +; di-, aut-, or synoicous Asterids perichaetia basally in tomentum; seta short or long Calomniaceae Cyrtopodaceae Mitteniaceae CAP widest at orifice; PS +/–; CY cucullate Campanulids predom. trop. mostly on forest floor Rhizogoniales Rhizogoniaceae Spiridentaceae CS –; upper lvs cells rounded occ. elongate, thick-walled, usu. papillose costa +; gonio- or cladautoicous, dioicous, or phyllodioicous CAP immersed or exserted, smooth or 8- (rar. 16-)ribbed, stomata phaneropore or cryptopore PS + (Orthotrichum-type) or reduced; annulus – or rudimentary; operculum +; CY +, often hairy gemmae +/–; saxicolous, corticulous Orthotrichales Orthotrichaceae pleurocarpids CS +; costa + aut-, par-, or dioicous, rar. heteroicous H CAP erect to horizontal, stomata phaneropore y annulus –; operculum +; PS ± reduced p CY cucullate; axillary propagules or rhizoidal gemmae Orthodontiaceae n Orthodontiales Theodor C. H. Cole, Dipl. Biol. Bernard Goffinet,Ph.D. a plants tomentose n CS +; lvs cells rounded-hexagonal, smooth or papillose Prof. Dr. Hartmut H. Hilger Dept. of Ecology and a costa +; di- or autoicous; perichaetia lateral Institute of Biology – Evolutionary Biology e CAP often striate to sulcate; annulus +; operculum +; CY cucullate gemmae-bearing pseudopodia (Aulacomnium); predom. temp. Aulacomniales Aulacomniaceae Freie Universität Berlin University of Connecticut 75 North Eagleville Road plants often stipitate ("dendroids") Altensteinstr. 6 CS +/–; PPP +, foliose D-14195 Berlin, Germany Storrs CT, 06269-3043, USA costa +/–; operculum +; PS +/– Braithwaiteaceae predom. trop. Hypnodendrales Hypnodendraceae Pterobryellaceae Racopilaceae COLE TCH, HILGER HH, GOFFINET B (2021) Bryophyte Phylogeny Poster CS –; PP +/–; lvs plicate/condupl., lamina cells ± incrassate • hypothetical tree based on molecular phylogenetic data (2021) costa +, double or – • branch lengths deliberate, not expressing actual time scale dioicous or phyllodioicous; CAP stomata +/–; annulus +/– occ. usu. Garovagliaceae • position of many characters on tree unclear spore germination precocious; CY cucullate (or mitrate) Ptychomniaceae Rhabdodontiaceae • some minor orders/families omitted gemmae +; trop. to south-temp.; usu. epiphytic Ptychomniales Orthorrhynchiaceae • characters do not necessarily apply to all members of the clades core Gondwanian dendroids • phylogenies: Cox et al. 2010, Dong et al. 2021, Flores et al. 2018, Knoop 2010, Liu et al. 2019, Long et al. 2016, modules: prostrate stolons, erect foliate stems, pleurocarps simple or branched, pinnate, flabellate, or dendroid Puttick et al. 2018, Shaw et al. 2011, Söderström et al. 2016, Sousa et al. 2018, Villarreal et al. 2010, 2012, 2016 only pleurocarpous CS usu. +; PPP +/–; costa +, single • characters: Crandall-Stotler et al. 2009, Frey et al. 2009, Goffinet/Shaw 2009, Ligrone et al. 2012 lvs cells usu. prosenchymatous usu. dioicous; seta straight or curved, smooth or mamillose PS Bryum-type CAP exserted, subglobose to oblong-ovoid; stomata + Hypopterygiaceae • abbreviations: G gametophyte, S sporophyte, AR archegonia, AN antheridia, CAP capsule CY mitrate or cucullate Hypopterygiales CS central strand, CY calyptra, PP paraphyllia, PPP pseudoparaphyllia, PS peristome, lvs leaves occ. occasionally, rar. rarely, usu. usually, + present, – absent, +/– some with/some without, ± more or less homocostate CS +/–; PP (–); PPP rare pleurocarps lvs cells often large, lax * the early diversification of the gave rise to other lineages, such as families in the informal protohaplolepidae, or the pseudoditrichales, Scouleriales and Bryoxiphiales: these lineages are not included here, for their relationships remain uncertain! costa +, single/double, rar. – Daltoniaceae Hookeriaceae Leucomiaceae ** Ptilidiales sister to Porelliales based on Dong et al. 2021 operculum + ° acrocarpous used s.l. here for terminal sporophyte (varying ontogeny) CY usu. mitrate asex. repro. by caducous lvs Hookeriales Pilotrichaceae Saulomataceae Schimperobryaceae Bernard Goffinet is supported by NSF grant # DEB-1753811 or branch tips, CS +/– Special thanks to Harald Kürschner, Dietmar Quandt, Juan Carlos Villarreal, and Misha Ignatov for valuable advice and consultation or filiform gemmae PPP usu. + alar cells often + Amblystegiaceae Anomodontaceae Brachytheciaceae Calliergonaceae CY cucullate > 4200 spp. (ca. 1/3 of all mosses) Hypnales Cryphaeaceae Hypnaceae Hylocomiaceae Lembophyllaceae A Leskeaceae Meteoriaceae Miyabeaceae Neckeraceae n Plagiotheciaceae Pterobryaceae Pylaisiaceae Pylaisiadelphaceae t Sematophyllaceae Thuidiaceae Trachylomataceae AN jacket of 4 cell tiers h Nostoc in longitudinal canals; pyrenoid – spores yellow, smooth, monolete mark + o pseudoelaters long, usu. unicellular Leiosporocerotales Leiosporocerotaceae (Leiosporoceros)

c AN jacket of 4 cell tiers e Hornworts thallus cavernous with mucilage-filled schizogenous cavities spores dark-brown/blackish, trilete mark +, spinose r pseudoelaters helical thickenings partly present Anthocerotales Anthocerotaceae (Anthoceros s.l., incl. Sphaerosporoceros and Folioceros)

o thallus orbicular or strap-like, often rosettes AN 2-8 per chamber Nostoc in schizogenous slime cavities S with/without stomata t (mostly ventral via mucilage clefts) spores yellow-blackish, trilete mark +, equatorial girdle + chloroplast usu. 1/cell with pyrenoid pseudoelaters occ. – Notothyladaceae (, Mesoceros, , Paraphymatoceros) o oil droplets + Notothyladales water-conducting cells – ) p AN 1–many, of endogenous origin AN 1-3 per chamber AR single, embedded on dorsal thallus surface AN thalli narrow, lingulate; pyrenoid + or – jacket cells h blepharoplast: spline of 12 microtubules (inaperturate), dioicous; spores yellow, later darkening CC-BY lamellar strip rhomboidal, basal bodies 2 of equal size, side by side pseudoelaters without helical thickenings Phymatocerotaceae (Phymatoceros) ( irregularly Phymatocerotales y zygote division longitudinal, three-tiered embryo arranged seta – t S chlorophyllous, mostly horn-like, AN one per chamber growing from basal foot by indeterminate, intercalary meristematic activity S with or without stomata spores transparent or yellow a columella well or poorly defined pseudoelaters with helical thickenings Dendrocerotaceae (Dendroceros, Megaceros, Nothoceros, Phaeomegaceros) stomata on S Dendrocerotales pseudoelaters (mostly multicellular) spore production continuous lignans +, flavonoids – ca. 200 spp.

Vascular Plants see Tracheophyte and Angiosperm Phylogeny Posters © COLE, HILGER, GOFFINET 2021