Botanica Pacifica. A journal of plant science and conservation. 2019. 8(1): 85–103 DOI: 10.17581/bp.2019.08109 Liverworts of the Russian Far East: the taxa with ciliate leaves Vadim­A.­Bakalin Vadim­A.­Bakalin ABSTRACT e­mail:­[email protected] The liverwort taxa with ciliate leaves occurring in the Russian Far East are reviewed. In total, 9 taxa belonging to four families (Jubulaceae, Ptilidiaceae, Neotrichocolea­ Botanical Garden­Institute FEB RAS ceae and Trichocoleaceae) are recorded, 6 species occur in Russia in the Far East Vladivostok,­Russia on­ly. ­The­identification­keys,­morphological­descriptions­and­figures­based­on­ma­ te rials from the Russian Far East are provided. Keywords: Jubulaceae, Ptilidiaceae, Neotrichocoleaceae, Trichocoleaceae, Hepaticae, the Russian Far East, East Asia * corresponding author РЕЗЮМЕ Manuscript­received:­23.10.2018 Бакалин В.А. Печеночники российского Дальнего Востока: таксоны с Review­completed:­28.02.2019 рес нитчатыми листьями. Ревизованы­таксоны­печеночников­с­реснитча­ Accepted­for­publication:­03.03.2019 ты­ми ­листьями,­встречающиеся­на­российском­Дальнем­Востоке:­9­видов­ Published­online:­05.03.2019 из ­4­семейств­(Jubulaceae,­Ptilidiaceae,­Neotrichocoleaceae­and­Trichocoleaceae).­ Шесть­видов­встречаются­в­России­только­на­Дальнем­Востоке.­Приводятся­ ключи­для­определения,­морфологические­описания­и­иллюстрации,­со­ став­ленные­на­основе­изучения­материала­с­российского­Дальнего­Востока. Ключевые слова: Jubulaceae,­Ptilidiaceae,­Neotrichocoleaceae,­Trichocoleaceae,­пе­че­ ноч­ники,­российский­Дальний­Восток,­Восточная­Азия The liverworts with ciliate leaves occurring in the Rus­ 2. ­Leaves­conduplicate­or­ventral­lobe­pyxidate,­without­wa­ si an Far East do not form the monophyletic group; these ter sac in the ventral base of the leaf ..................................... 4 are representatives of various families belonging to even dif­ 3.­Cells­along­leaf­margin­with­large­papilla­like­pro­tu­be­ran­ fe­rent ­orders:­Porellales­(Jubulaceae),­Ptilidiales­(Pti­li­dia­ceae,­ ces by one per each cell, water sacs present in the base of Neotrichocoleaceae) and Jungermanniales (Tricho co lea ceae). vent ral lobes and underleaves ...... Ascidiota blepharophylla Plants are also varying in the size, from tiny in Neo hat toria [Arctic­Alpine­broadly­mega­Beringian­highly­disjunctively­dist­ri­ bu­ted ­taxon,­not­known­in­the­Russian­Far­East,­but­may­be­found,­ herzogii to large in Trichocolea tomentella and Pti li di um ciliare. The treated­in­Bakalin­&­Klimova­2019] attempt to unite all of them in the same treat ment is based on ­practical­reason­only: the­majority­of ­taxa­could­not­be­ 3.­No­papilla­like­protuberances­along­leaf­margin,­un­der­ leaves without water sacs, water sac of the leaf is completely keyed­out­using­keys­existed­in­Russia.­More­over,­the­keys­ trans formed ventral lobe .............. Trichocoleopsis sacculata existed­ in­ adjacent­ countries­ do­ not­ covers­ all­ re­cog­nized­ Russian taxa of this group. Therefore the main goal of the 4. ­Leaves­with­pyxidate­ventral­lobe­......................................­5 present account was to accumulate all available infor mation 4. ­Leaves­with­ventral­lobe­appressed­to­the­dorsal­lobe,­near­ on ­ liverworts­ with­ ciliate­ leaves­ known­ in­ the­ Rus­sian­ Far­ ly­plane­or­slightly­concave­to­convex,­with­1–4­teeth­along­ East ­and­provide­descriptions,­figures,­iden­ti­fi­cation­keys­and­ upper half .................................................... Nipponolejeunea data on ecology and distribution within one paper. The paper treats taxa whose leaf lamina is more or less developed, and 5. ­Underleaves­only­slightly­wider­than­stem­(1.2–1.7­of ­stem­ width), bilobed, neither ciliate nor dentate, ventral lobes does not include such genera as Blea ph ro stoma where leaves easi ly caducous, leaves dentate (sometimes prominently so) divided nearly to the base and no true lamina exists. to ­shortly­ciliate,­plants­300–500­µm­wide­............................... ................................................................. Neohattoria herzogii TAXONOMIC TREATMENT 5. ­Underleaves­2–4­times­wider­than­the­stem­diameter,­bi­ The­key­to­the­genera­with­ciliate­leaves­and­developed­ tetra lobed, commonly with cilia along margin, leaves ciliate, leaf­lamina­occurring­in­the­Russian­Far­East: plants­commonly­wider­1.0­mm­wide­.........­Jubula japonica 1. ­Leaves­conduplicate­or­with­pyxidate­ventral­lobe­or­bear­ 6. ­Leaf ­lamina­reduced,­the­80­%­of ­the­leaf ­are­cilia,­plants­ wa ter sac in the base of the leaf ............................................. 2 whi tish greenish to brightly so ......... Trichocolea tomentella 1. ­Leaves­not­conduplicate,­ventral­lobe­not­pyxidate,­no­ 6.­Leaf ­lamina­not­reduced,­distinct­and­relatively­wide,­trian­ wa ter sac in the leaf base ......................................................... 6 gu lar lobes divided into several ‘sublobes’, which terminate by cilia (that may be poorly developed) .................... Ptilidium 2. ­Leaves­with­water­sac­in­the­ventral­base­..........................­3 ©Botanical Garden-Institute FEB RAS. 2019 85 Bakalin Jubulaceae­H.­Klinggr con vex, with apex distinctly curved to ventral side, densely Jubulaceae is the small family including three genera (all den­tate­ciliate,­600–800­×­280–600­µm,­smaller­leaves­with­ are ­known­in­the­Russian­Far­East),­distributed­in­temperate­ dor sal lobes not ciliate, with only attenuate­pilose apex; to tropical (in the latter absent in the low elevations, but vent ral lobe pyxidate, slightly longer than wide, lobe axis at com mon in mountains) amphioceanic areas, southward to 15–60(–80)°­with­stem­axis,­ca.­180–200­×­150­µm,­almost­ nor thern Australia. They are characterized by green to brow­ ina v riable in size across the shoot despite considerable va­ nish color of plants without red pigmentation, branching ria tions in the dorsal lobe size, stilus virtually absent to of Lejeunea and Frullania type, rhizoids originating from un­ reduced­to­slime­papilla.­Underleaves­contiguous­to­slight­ der leaf bases, incubously inserted leaves with distinct dif­ ly overlapping above situated underleaves, sinuously in ser­ fen re tiation into dorsal and ventral lobes and dorsal lobe ted, with hardly auriculate base, appressed to the stem or apex acute to piliferous, more or less thin­walled leaf cells, nar­rowly­obliquely­spreading,­small­underleaves­nearly­rec­ bi lobed (sometimes tetralobed) underleaves, absence of tan­gular,­ca.­350­×­200­µm,­with­sinus­descending­to­3/5­ pe ri gynium and reduced foot of seta, seta short, capsule of the length, margin entire or with additional teeth in one bis­tra­tose­ spherical­ (Frey­ &­ Stech­ 2009).­ The­ status­ and­ or both lateral sides, some small underleaves are si mi lar to tao x nomical composition of the family remained unclear larger­underleaves­that­are­widely­ovate­to­800­×­700­µm­ un­til ­recently,­when­Larraín­et­al.­(2015)­clearly­defined­the­ (excluding cilia), densely ciliate­dentate throughout, di vi ded structure of the family and its relationships with mor pho­ by­V­­to­U­shaped­sinus­descending­to­1/2­of ­the­length­into­ lo­gi­cally­ similar­ Frullaniaceae­ and­ Lejeuneaceae,­ especially­ 2 ­prominently­apiculate­lobes.­Cells­in­dorsal­lobe­middle­ ar guing Neohattoria as the member of Jubulaceae, but not subisodiametric­to­oblong,­20–35­×­12–25­µm,­thin­walled,­ mer ging with Frullania as it was commonly estimated be fore. with small concave trigones, cuticle smooth, inter mediate Jubulaceae in the Russian Far East are restricted to sou­ thickenings­absent­or­obscure­and­present­in­lower­half ­of ­ thern­most ­flank­of ­the­land­and­occur­in­Primorsky­Ter­ri­ the lobe; oil bodies ellipsoidal to shortly fusiform, nearly to­ry, ­southern­flank­of ­Khabarovsk­Territory,­Sakhalin­Is­ filling­cell­lumen,­distinctly­biconcentric.­Autoicous.­And­ land­and­southern­Kurils.­The­genera­were­keyed­out­in­the­ roecia on short lateral branch, bracts cupped, imbricate, bi­ general­key. lo­bed­for­1/3–1/2­of ­the­length,­margin­entire­or­remotely­ dentate, bracteole ovate, shortly incised. Perianth terminal Jubula­Dumort.,­Commentat.­Bot.­(Dumortier):­112,­1822 on­leading­axis,­distinctly­plicate­in­upper­half ­(2­ventral­and­ Jubula­is­the­small­genus­of ­Hepaticae­that­counts­5­spe­ 2­lateral­folds),­obscurely­beaked,­exerted­for­1/2–3/4­of ­ cies­(Söderström­et­al.­2016),­although­with­many­subspecies­ the ­length,­obconical­obovate,­ca.­1.7­×­0.6­mm;­bracteole­ within J. hutchinsiae­(Hook.)­Dumort.­Noticeable­all­taxa­that­ narw ro ly ellipsoidal, shortly incised, bracts bilobed, lobes were ­tested­(Larraín­et­al.­2015)­were­placed­into­sub­spe­cies­ lan­ceo­late,­unequal,­with­margin­dentate,­but­not­ciliate.­ of the last mentioned taxon, whereas other (although from Comment. This species is easily to recognize due to pi­ the ­morphological­point­of ­view­this­may­look­strange­e.g.­ xy date ventral lobe and ciliate leaves and underleaves. Pro­ for J. hattorii­r Uda ­et­V.Nath)­treated­in­the­last­checklist­as­ bab­lyy ­it­ma ­be­mistaken­with­Frullania and Neohattoria in dis crete species. Moreover, many taxa described by Stephani re­gio­nal­flora­due­to­pyxidate­ventral­lobe,­however­it­easi­ly­ were merged to J. hutchinsiae ssp. javanica (e.g. J. rostrata differs from regional Frullania in ciliate leaves and under ­ Steph., J. sikkimensis Steph., J. tonkiensis Steph.) following to leaves and in larger size of plants and relatively large under ­ Guerke­ (1978).­