Studies on Victorian Bryophytes 6. Key to Thallose Liverworts and Hornworts

2006·1.6 MB·English

by  David Meagher

Bryophytespecialissue
overlapstheone fartherthe stemapex Trigone Triangular to cordate thickening
(orwould iftheywerecloseenough). atthepointwherethreecellsjoin.
Tooth Small, tapering projection on Underleaves Leaves ofa different size
margin ofleafor thallus, consisting (usually much smaller) and shape
ofone or a few cells, or formed by than the lateral leaves, and attached
an extension ofacell wall. ontheventral side ofthestem.
Transverse Having thejoin between the
leaves and stem running sideways Ventral Onthe underside ofthethallus or
acrossthestem,notangled. shoot,i.e. closesttothesubstratum.
Fa /
kr
transverse leaves
vY
Studies on Victorian bryophytes 6.
Key to thallose liverworts and hornworts
David Meagher
SchoolofBotany,TheUniversityofMelbourne,Victoria3010
Abstract
Anewkeytothegeneraandmanyspeciesofthalloseliverwortsandhornworts(exceptFossombronia
andRicciaspecies)inVictoriaisprovided.{TheVictorianNaturalist123(4),2006,247-254)
Introduction
This artificial key complements the key Tasmania, New Zealand or other parts of
to the genera of leafy liverworts in this the world might still be found in Victoria.
volume. It is based on the key to southern This key is not valid for other regions of
Australian liverworts in Scott (1985), but Australia.
is substantiallyupdated and revised to take Most ofthe thallose liverworts and horn-
into account taxonomic changes and addi- worts in Victoria are described and illus-
tions to the Victorian flora in the last 20 trated in Scott (1985) and Meagher and
years. Common mistakes are allowed for Fuhrer(2003).
inthe main keyand group keys. Namesoftaxa follow thecurrentnational
The key can be used to identify speci- checklist(McCarthy 2006).
mens to species level, except forspeciesof A basic glossaryofterms used inthis key
Fossombronia (which are very difficult to is included in the key to leafy liverworts
identify without detailed analysis) and (Studies 5 in this issue). For a complete
Riccia (which is under review in Australia and beautifully illustrated glossary ofbry-
and is likely to undergo substantial ological terms, see Malcolm and Malcolm
changes). Also keep in mind that species (2000).
and genera presently known only from
Vol. 123 (4) 2006 247
1
Bryophytespecialissue
Key tothallose liverworts and hornworts
Thallus leafy, orwith leaf-like lobes oneitherside ofacentral axis 2
Thallus lobcd orunlobcd, but not leafy 5
Rhizoidscrimson Fossombroniaceae (Group A)
Rhizoids notcrimson, orabsent, ornot seen 3
3 Lobes pinnateoralternate, arranged all alongthecentral thallus;
orplant leafy or lettuce-like 4
Lobespalmate, the lobes radiatingorbifurcating(Y-branching) from
a central point; orthallus arising from a narrow,± upright stalk 1
4 Thallusbrightgrass-green, thickand lleshy,ratherbrittle;
rare plant Treubiaceae (Treubia tasmanica)
Thallusgreen ornot; not thickand fleshy, not brittle;
common plants Fossombroniaceae (Group A)
5 Thallusone cell thick(except midrib) 6
Thallus mostlyseveral cellsthick in part 8
6 Thallus narrowthroughout(< 3 mm), lobespinnate, alternateorbifurcated 7
Thalluswide, at least in part(usually> 4 mm).
often palmately divided Pallavicinaceae (Group C)
7 Growing on wet or dried soil, commonly mud (either saline or fresh);
lobepattern usually not obvious Sphaerocarpales (Group D)
Growingon trees orrocks, neveron mud; lobes
bifurcating(Y-branching) Metzgeriaceae (Group E)
8 Chloroplastsusually l or2 percell; capsuleerect, needle-like,
splittinggraduallydown from tip; large cavitiescontainingdark
cyanobacteria often evident in thallus Anthocerophyta (Group B)
Chloroplasts several tomanypercell; capsuleovoidto globose,
notneedle-like; cavities in thallus (ifpresent) not containingdark cyanobacteria . 9
9 Thallus half-buried in soil,firmlyanchoredbycopiousrhizoids;
capsule formed in marsupium buried in soil Enigmellathallina
Thallusonorabovesoil, ornot on soil, anchored ornot byrhizoids;
capsule not formed inmarsupiumburiedin soil 10
10 Thallussurface without pores, uppersurfacehomogeneous, withoutpores;
rhizoidsall smooth 11
Plants surfacewith pores, often openingtothe uppersurfacebyairpores;
rhizoidsnormallyoftwo sorts: onesmooth and theotherwith internal
peg-likethickenings 15
11 Midrib conspicuous; lamina 1 cell thick inouterparts 12
Nomidrib; laminanever 1 cell thick, except sometimes attheveryedge 13
12 Plants on bark orrock, neveron soil; paleyellow-green, neverrose-tinted;
prostrate; linehairspresent on ventral surfaceand usuallyalso on
thallus margins Metzgeriaceae (Group E)
Plantsonsoil orrotting logs,alsoepiphytic in humid habitats;
mid todarkgreen, often rose-tinted, often arising from anarrow,
±uprightstalk; ventral surface andmargins lackinghairs
(but may be toothed) Hymenophytaceae/ Pallavicinaceae (Group C)
248 The Victorian Naturalist
Bryophytespecialissue
13 Plantswith regular, few-celled lobes intheposition ofleavesand
underleaves miskevedZoopsisoreroded leafy liverwort
Plants usually irregularly lobed; lobes many-celled 14
14 Sporophyte needle-like; chloroplasts usually 1 or2 percell;cavities in
thallus containing dark cyanobacteria often present ... Anthocerophvta (Group B)
Sporophytenot needle-like; chloroplasts usually several to many percell;
cavities in thallus containingdark cyanobacteria
never present Aneuraceae (Group F)
15 Gemmacupscircularorcrescent-shaped,obviouson uppersurface
ofthallus * Marchantiaceae(Group G)
Gemma cups lacking 16
16 Uppersurface ofthallus spongy, often whitish 17
Uppersurface ofthallus firm, usually green 18
17 Plants usually formingcompleteorpartial rosettes
on theground, orelse free-floating; not in salt pans Ricciaceae (Group H)
Plantsnotformingrosettes; in saltpansoron
compacted soil Sphaerocarpales (Group D)
18 Uppersurfaceofthallusflat, notfurrowed; sporophytes
carried outsidethe thallus * 19
Uppersurface furrowed, V-shaped at leastatapex; sporophytes
embedded in thallus Ricciaceae (Group H)
19 Thallus usually> 7 mm wide; many long, free rhizoidson
ventral surface Marchantiaceae (Group G)
Thallus usually< 6 mm wide;neverwith rhizoids
as above Aytoniaceae and Targioniaceae (Group I)
Group A
Fossombroniaceae
1 Plantsaquatmicmorsemi-aquatic; thallus erect,
upto 30 tall Austrofossombronia australis
Plants notaquaticorsemiaquatic(butmaybeon dryingmud);
thallus prostrate, < 10 mmtall 2
2 Rhizoidshyalineorbrown, nevercrimson; Thallus± as longaswide, ruffledand
lamellateon dorsal surface Petalophyllumpreissii
Rhizoids usuallycrimson; thallususuallymuch longerthan wide,
not lamellate Fossombronia
Group B
Anthocerophyta
1 Chloroplasts2 ormorepercell; capsulewith spirallythickened
and unsegmented elaters 2
Chloroplasts usually percell; capsule with irregularsegmented pseudo-elaters 3
1
2 Thallus < 25 mm long Megacerosgracilis
Thallus35-50 mm long Megacerospellucidus
3 Thalluswith aroughand cavernoussurface, usuallypalegreen
with crisped margins; spores blackish Anthocerospunctatus
Thallus with smooth uppersurface,usuallydarkgreen
with marginsrarelycrisped; sporesyellowish 4
4 Plants dioecious Phaeoceros laevis
Plantsmonoecious Anthocerosbrotheri
Vol. 123 (4) 2006 249
Bryophytespecialissue
Group C
Hymenophytaceaeand Pallavicinaceae
1 Plants with sex organs 2
Sexorgans lackingornot visible.. 8
2 Sex organs on specialised shortbranches atbaseoron undersideoffrond 3
Sex organs on uppersideoffrond, noton specialised branches 4
3 Sexual branchesatbaseoffrond; thallussimple orsparselybranched.
not palmate Podomitriumphyllanthus
Sexual brancheson underside offrond; thalluspalmately
divided above Hymenophytonflabellatum
4 Sporophytebase encased in a thick, fleshytubebearingarchegontanearapex;
maleplants with scales overlapping midrib dorsally 5
Sporophytebase surroundedbya longtubularpseudoperianth,not fleshy;
maleplantswith scales in 2 rowsdown eachsideofmidrib 6
5 Thallusbranched, margins coarselytoothed Symphyogynapodophylla
Thallusunbranched, consistingofbroadandnarrow sections,
margins entire Symphyogyna interrupta
6 Marginsstrongly toothedwith conspicuousteeth,
several cells long Pallaviciniaxiphoides
Marginsgenerally entireorwithteeth ofonly 1 or2 cells 7
7 Thallus± flat, the marginsrarely ifeverflexed upwards* Pallavicinialyelli
Thallus commonly concave, the margins flexed upwards Pallavicinia ruhristipa
8 Frond margins toothed, at least nearapex 9
Frond margins entireornearly so 10
9 Marginswithteeth ofonly or 2 cells** Pallavicinia lyelli
1
Margins with teeth several cells long P. lyelli/S. interrupta (inseparable)
10 Thallus, commonly concave, the margins flexed upwards Pallavicinia ruhristipa
Thallus± flat, the marginsrarely ifeverflexed upwards 11
11 Fronds borne onerect stalks; plantcompletelygreen Hymenophytonflabellatum
Fronds prostrate; plantmay havearose-pinktinge 12
12 Stalk rose-pink, at leastnearbase Pallavicinia lyelli
Stalk completely green Podomitriumphyllanthus
* Schuster (1991) gave the name Pallavicinia pseudo/yellii to Australasian material of
lyellir and gave a Latin diagnosis, but did notvalidate the name by nominatinga
type.
** Jenseniaconnivens, discounted from the Australian flora by Schaumann etal. (2004),
would keytocouplet9; ithas frondsborneon erectstalksbut istinged roseredbelow.
250 The Victorian Naturalist
Bryophytespecialissue
Group D
Sphaerocarpales
31 Thallus±circular,usuallyalmostcoveredbyinflated,bottle-like involucres 2
Thalluselongated, consistingofa stem with awingalongoneside,
spore-bearing involucres atthe edge and tip 3
2 Thallusbubble-like,with a single poreontop;
in saline habitats Monocarpussphaerocarpus
Thallusflat, withseveral involucrestogether;
not in saline habitats Sphaerocarpostexanus
Monoicous; on freshwatermud; spineson spores 12 pm long Riellaspiculata
Dioicous; on salinemud; spineson spores4-5 pm long Riellahalophila
6
Group E
Metzgeriaceae
1 Thalluswith hairson both dorsal andventral surfaces* Metzgeria sp. A
Thalluswithouthairson dorsal surface 2
2 Thallus lobed and saccate Metzgeriasaccata
Thallus flat, notlobedorsaccate 3
3 Hairsweaklytodistinctly falcate,mostlypaired;midribcovered
by2-3 cellson dorsal side Metzgerialeptoneura
Hairs notfalcate, pairedand/orsingle; midrib coveredby2-4cellson dorsal side ... 4
4 Thallustapered to anarrow apexon mostlobes Metzgeriaconsanguinea
Thallusrarely ifatalltapered, most lobeswith an obtuse, rounded apex 5
5 Midrib coveredby3(-4)cells on dorsal side,4-6 on ventral side Metzgeria rigida
Midrib coveredby2(-3) cells on dorsal side, 2-4 onventral side 6
Midribcovered by2 cells onventral side** Metzgeriadecipiens
Midrib covered by (3-)4cells on ventral side Metzgeriafurcata
Notes:
Cells covering the costa should be counted about halfway between one thallus branch
andthenext.
* Metzgeria sp. A from Carlisle State Park seems closest to M. follicola of
Melanesia.
** So (2002) followed Grolle (2002) in reducing M. decipiens to a synonym of
M. furcata based on the variability in the number of cells covering the midrib
,
on the dorsal side of the thallus. However, the number of cells on the ventral
side seems to distinguish the two clearly. Until a full assessment of the two
taxa ismade, 1 prefertomaintainthem asseparateentities.
Vol. 123 (4) 2006 251
Bryophytespecialissue
Group F
Aneuraceae
1 Thallus U-shaped in cross-section, at leastnearlobetips 2
Thallus flatorslightlycurved in cross-section, neverU-shaped 4
2 Thallus marginsplane; lobeapicesspoon-shaped, oftenyellowish
and bearinggemmae Riccardia cochleata
Thallus lobes flexuose tocrispate; lobe apices not asabove 3
3 Thallus>7 mm wide; marginsstronglycrisped; aquaticplantofalpine
or subalpine streams Aneura sp. A
Thallus< 6 mm wide;margins flexuosetoslightlycrisped;
not aquatic Aneura rodwayi
.
4 Apex ofthallus not dissected 5
Apexofthallusdissected 8
5 Plantdendroid orsemi-dendroid; thallus differentiated intoa central
stem and branches; lateral brancheswithevidentcentral strands;
cuticlepapillose Riccardiaeriocaula
Thallus withoutan erect stem; lateral brancheswithoutacentral strand;
cuticle smooth 6
6 Thallus brancheswith awing I cell thick; mucilage papillae lateral
andventral only, persisting; shoot calyptra smooth Riccardiaminima
Thallus branches notwinged; mucilagepapillaedorsal aswell as lateral
andventral, not persisting; shoot calyptracrownedwith hyaline hairs 7
7 Thallus lens-shaped incross-section Riccardiaaequicellutaris
Thallus circular in cross-section Riccardia alcicornis
8 Cuticle striolateorpapillose 9
Cuticle smooth 10
9 Cuticle striolate Riccardia crassa
Cuticlepapillose Riccardiacolensoi
10 Thallusmean width>2 mm (usually3-6 mm);
apex deeply dissected Aneuraalterniloba
Thallusmeanwidth<2 mm; apex shallowly divided 11
11 Thallusmean width< 1 mm; oftenwith prostratemainbranches
and erect, pinnately branched(almost palmate)secondarybranches;
monoecious Riccardia wattsiana
Thallus mean width > mm; branches prostrate; dioecious 12
1
12 Branchingoftenappearingpalmate; mucilage papillaenotpersisting;
stolonspresent Riccardiarupicola
Branchingalwaysclearlypinnate; mucilagepapillae persisting;
stolons absent Riccardiabipinnatifida
Note:
Aneura sp. A is an undescribed species known from the Bogong High Plains, Baw Baw
Plateauand KosciuszkoNational Park. Itisprobably alsopresentinNewZealand.
252 TheVictorian Naturalist
Bryophytespecialissue
Group G
Marchantiaceae
1 Gemmacups(ifpresent)crescent-shaped;poreson uppersurface
ofthallusnotsurrounded by polygonal shapes Lunulariacruciata
Gemmacups(ifpresent) circular;poreson uppersurface
surrounded bypolygonal shapes 2
2 Ventral scalesforminganarrowcrimson stripedownthecentreofthe
undersideofthethallus; archegoniophorelobes flat,
rectangular Marchantiafoliacea
Ventral scalescolourless,coveringundersideofthallus;
archegoniophore lobes rod-like 3
3 Colourofuppersurfaceofthallusevenly green; surfacewith aglossysheen;
marginal scalesnotprojectingbeyondthallus edge Marchantiaberteroana
Colourofuppersurfaceofthallus uneven, with adarkzone
downthemiddle; surfacewithoutaglossysheen; marginal scales
projecting slightlybeyond thallus edge Marchantiapolymorphavar.aquatica
Group H
Ricciaceae
Plants free-floating 2
1
Plants on soil ormud 3
2 Ventral scales conspicuous, purplish Ricciocarposnatans
Ventral scalesnot evident, notcoloured Ricciaduplexvar. duplex
3 Thallusheart-shaped; ondryingmud;ventral scalespurplish,
in bunches Ricciocarposnatans
Thallusheart-shapedornot; on varioussubstrates;
ventral scalesvariously coloured butnotin bunches 4
4 Dorsal surfaceofthalluswith compacttissuesforming
narrowveiticalairchambers, withoutspecialisedpores;
epidermal cellshyaline RicciasubgenusRiccia
Dorsal surfaceofthalluswith loosely arranged(often spongy)
tissuesformingpolyhedral orlargeand irregularairchambers
withwell-definedpores; epidermal cellschlorophyllose
exceptaround pores RicciasubgenusRicciella
Vol. 123 (4) 2006 253
Bryophytespecialissue
Group
I
AvtoniaceaeandTargioniaceae
1 Side branchesoriginating fromundersideofthallus;capsules formed
in blackspherical pouches beneath apicesofthallus Targioniahypophylla
Side branchesoriginatingfrom marginoruppersurfaceofthallus;
capsules formed in umbrella-like streutures (archegoniophores) ... (Aytoniaceae) 2
2 Sexorgansalwayspresent,in2 ormorereceptaclesdown the
midline ofthe thallus Plagiochasma rupestre
Sexorgansabsent, orscattered receptacles usuallyon themargin
orapex ofthethallus 3
3 Epidermal poressurrounded by4 ormoreringsofcells; perianth (involucre)
hemispherical, with one slit beneath Reboiiliaqueenslandica
Epidermal poressurroundedby 1-3 ringsofcells; perianthconical,
openingwithnumerousvertical slits Asterella4
4 Thalluscrimson underneath, generally 5-7 mm wide;
perianth with 12-14 slits* Asterelladrummondii
mm
Thallusgreen underneath,generally3-4 wide;
perianth with about8 slits Asterellatenera
*Asterella conocephala, A. tasmanica and A. whiteteggeana are almost certainly
conspccificwithA. drummondiiorA. tenera.
Acknowledgements flora ofAustralia Supplementary Series No. 20.
wMhaonypotihanntkesdaoruetdeurerotrostiwnotahneonmyanmuosucsrirpetferaeneds Sc(hAuBstReSranRdMFN(G1V9:91C)anDbierarganoasndesBloafcknbuerwn)taxa of
madesomevaluablecommentsandsuggestions. HBeoptaainiiccaael.LIa.bJournagtconrnyanHn)i:id14a3c.J15o0u.rnaloftheHatton
References Scott GAM (1985) Southern Australian Liverworts.
Grolle R (2002)Typificationsofthree old names of CAuasntbrearlriaa)n Flora and FaunaSeriesNo. 2. (AGPS:
Metzgeriaspecies(Hepaiicae):Jungermanniafurca- SchaumannF,PfeifferTandFreyW(2004)Molecular
ta L. 1753. ./. linearis Sw. 1788 andJ. puhescens divergencepatternswithin theGondwanan liverwort
MaBSlrccyhoorlpahmnyktBe1s7a:9n2dA.nMCarIlylcplotulosmtgr.aamtNiede(GB2lr0oy0so0sl)aorMgyio.css(2e3Ms:ica1r1no9-d-O1Op2tt1hi.cesr gBbrerynyouopsphhyJytctane)ss.enS2it5au.d(iJPeoasulrlinunvaAilucsmotifraactlchaetee,HmapHetertpaoattneierBoaopitnbayntfioircneaas,lt
McLPCrievasersr:twhNoeyrltssoPnaM.nNdZ(H2)o0r0n6w)nrCthse.ckVleirsstionof6AAupsrtilra2l0i0a6n SoaLnaMdboLtrhae(t2oP0ra0yc2i)9f.6ic:,M2eN3t1ezgwe2r4Zi4e.aal(aHnedpaJtoiucranea)lionfABuosttarnalyas4i0:a
(www.anbg.gov.au/abrs).(ABRS:Canberra) 603-627.
Meagher Dand FuhrerB (2003)A FieldGuidetothe
Mosses andAlliedPlants ofSouthern Australia. Received13April2006;acceptedIJune2006
One hundred and nineteen years ago
MOSSES OFVICTORIA, WITH BRIEFNOTES
BY D. SULLIVAN
cWheretolookforthem. Onand in the crevicesofrocks,on logs,aboutthebasesand rootsof
trees, on banks ofwatercourses, lakes, lagoons, and waterholes, on the ground, from the low
landstothesummitsofourhighestmountains-both inwetanddrylocalities,butmoreespecial-
ly in the former. ... I would recommend Melbourne collectors to search well about the Yarra,
Dandcnong, You Yangs, Mount Macedon, Riddle’s Creek, Lancefield (Deep Creek), Sunbury,
Gisborne,etc. September,October,andNovemberarethebest months forthedry localities,and
December, January, and February for the higher mountains and moist forest country. Mosses
mayhefoundincertainlocalitiesthroughouttheyear,butinwinter,exceptinrarecases,theyare
not in a fit state fordetailed examination, having lost both the calyptras and operculums parts,
whicharesometimesofgreatvalueindecidingspecificdistinctions.’
From The VictorianNaturalistIV(1887-8),pp. 109-110
254 The Victorian Naturalist