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ПОРЯДОК BARTRAMIALES M. Menzel Клювовидной

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ПОРЯДОК BARTRAMIALES M. Menzel Клювовидной Bartramiaceae: общая часть 13 ПОРЯДОК BARTRAMIALES M. Menzel клювовидной. Колечко не отпадающее. Перистом прикреплен ниже устья, двойной, простой, реже М.С. Игнатов, Е.А. Игнатова очень ломкий, рано обламывающийся и выглядя- Растения от мелких до крупных, в рыхлых или щий отсутствующим; зубцы экзостома согнуты и густых дерновинках, от сизовато-зеленых и зеле- закрывают устье коробочки в сухом состоянии, в ных до желто- и буро-зеленых. Стебель часто крас- основании сетчато-папиллозные или гладкие; эндо- ный, прямостоячий, простертый только у мелких, стом с высокой или низкой базальной мембраной угнетенных растений (Philonotis); ветвление или редуцирован; сегменты эндостома хотя морфо- подверхушечное, с одним или несколькими побе- логически и очередны зубцам экзостома, но выгля- гами; густо 3-, 5- или многорядно облиственный, дят супротивными из-за того, что они обычно рас- часто с б. м. выраженным спиральным располо- щеплены по килю сегмента, и половинки сегмен- жением листьев, внизу у большинства видов б. м. тов срастаются с половинками соседних сегментов войлочный; с центральным пучком или без него, (Рис. 2). Споры чаще крупные (15–)20–35 (–50)m. с гиалодермисом или без него; ризоиды в нижней Колпачок клобуковидный. части стебля, располагаются по линии, продолжа- Порядок включает одно семейство. ющей нижний угол низбегания листа. Листья пря- мо отстоящие до отстоящих, реже отогнутые или СЕМ. BARTRAMIACEAE Schwägr. — извилистые до курчавых, из б. м. яйцевидного, БАРТРАМИЕВЫЕ иногда плотно прилегающего к стеблю основания Семейство включает 10 родов, с центром раз- узко или широко ланцетные, постепенно длинно нообразия в высокогорьях Южной Америки. заостренные; край вверху или по всей длине пиль- 1. Сизые растения с плотно прижатыми в сухом чатый, плоский, отогнутый или отвороченный; состоянии, жесткими листьями, расположен- жилка мощная, оканчивается в верхушке листа ными в пяти четко выраженных прямых рядах; или выбегает, на поперечном срезе б. ч. слабо диф- крышечка с клювиком; перистом простой, ференцированная или ближе к основанию с выра- зубцы гладкие, попарно сближенные, на вер- женным рядом указателей, окруженными одним хушке б. м. плотно соединенные в виде конуса; или двумя пучками стереид или субстереид и аркто-альпийский вид .......... 4. Conostomum верхним и нижним эпидермисом, образованными — Буро-, желто- или сизо-зеленые растения, с более мелкими клетками; клетки в верхней части жесткими или мягкими, отстоящими, реже листа квадратные до удлиненно прямоугольных, с б. м. прижатыми листьями, не расположенны- мамиллами или папиллами в углах или над про- ми рядами или же в рядах, образующих спи- светом клетки, или с кутикулярными папиллами, раль вдоль стебля; крышечка плоская или низ- заметными в виде продольной штриховатости, или ко выпуклая; перистом двойной, зубцы папил- гладкие; в основании листа клетки прямоугольные лозные или с сетчатой орнаментацией, не до продолговатых или линейных, гладкие или с бывают попарно сближенными, на верхушке папиллами в нижних углах, тонко- или умеренно свободные; многие виды широко распростра- толстостенные, клетки углов основания не диффе- нены как в горах, так и на равнине ............ 2 ренцированы. Вегетативное размножение пазуш- ными выводковыми почками (Philonotis) или легко 2. Стебель трехгранный, без центрального пучка; обламывающимися листьями (Bartramia). Одно- клетки в верхней части листа без папилл и домные, обоеполые, двудомные. Андроцеи и гине- мамилл, с мелкими кутикулярными папил- цеи верхушечные. Перигониальные листья у дву- лами, заметными б. ч. в виде продольной домных видов звездчато распростертые вокруг дис- штриховатости ........................... 1. Plagiopus ковидных перигониев. Перихециальные листья сла- — Стебель б. м. округлый или пятигранный, с бо дифференцированные. Спорофиты по 1(–2) из центральным пучком; клетки в верхней части одного перихеция. Коробочка на короткой или длин- листа папиллозные, мамиллозные или почти ной ножке, слабо или умеренно наклоненная, до гладкие, без кутикулярных папилл ............. 3 начала рассеивания спор, когда она закрыта 3. Листья сухие прямые, жесткие, прямо отсто- крышечкой – шаровидная, с высокой спинкой и ко- ящие, из беловато-блестящего, прилегающего сым устьем, сухая округло-овальная, глубоко про- к стеблю основания резко суженные в длинную дольно бороздчатая. Крышечка значительно уже шиловидную верхушку; пластинка вверху 2– диаметра урночки, низко коническая до широко 3-слойная ............................. 2. Bartramia p.p. 14 BARTRAMIACEAE A B C D E F G H Рис. 2. Bartramia pomiformis Hedw. (A, C, D), B. ithyphylla Brid. (B), Plagiopus oederianus (E–H): A, B, E – перистомы, 59, 120 и 78; C, F, G, H – фрагменты перистомов, показывающие расположение зубцов экзостома очередно сегментам эндостома (и супротивно ресничкам), но кажущееся супротивным сегментам, из-за того, что последние разделены по килю почти до основания, 112, 685, 196,358; D – орнаментация зубца экзостома в средней части снаружи, 1490. Plagiopus 15 A B Рис. 3. Conostomum tetragonum (A–B): A – общий вид перистома, заметно попарно сближенное расположение его зубцов, 108; B – внутрь согнутые зубцы экзостома,250. — Листья сухие серповидно согнутые, прямые — Stems round to pentagonal, central strand или извилистые до кудрявых; переход от при- present; upper leaf cells smooth, mamillose or легающего основания к пластинке листа б. м. papillose, papillae never appearing as longitudi- постепенный; пластинка вверху однослойная nal striolations ............................................... 3 или только у края 2–3-слойная ................... 4 3. Leaves rigidly straight when dry; leaf bases whit- 4. Листья сухие б. ч. извилистые до курчавых, ish, sheathing, abruptly tapered into subulate, 2– реже вверх направленные и в сухом состоянии 3-stratose upper leaf limbs with unistratose mar- торчащие; край листа 2–3-слойный; клетки gins ...................................... 2. Bartramia p.p. верхней части листа очень коротко прямо- — Leaves falcate-secund, erect (rarely straight), угольные, с пологой центральной мамиллой; flexuose or crisped when dry; leaf bases yellow- однодомный; мезофит ........ 2. Bartramia p.p. green, not sheathing, moderately to abruptly ta- — Листья сухие слабо односторонне согнутые pered into narrow, unistratose upper limbs or или прямые; край листа однослойный; клетки upper limbs with 2–3-stratose margins ......... 4 в верхней части листа удлиненные, с папиллой 4. Leaves flexuose, crisped or erect when dry; leaf в нижнем, реже в верхнем углу; двудомные, margins 2–3-stratose; upper leaf cells short-rect- очень редко однодомные; гигрофиты ............ angular or subquadrate; plants monoicous; mes- .................................................... 3. Philonotis ophytes ................................. 2. Bartramia p.p. — Leaves usually slightly falcate-secund, rarely 1. Plants glaucous; leaves rigid, tightly appressed straight when dry; leaf margins unistratose; up- when dry, distinctly arranged in five straight per leaf cells rectangular to elongate-rectangu- rows; opercula rostrate; peristome single, exos- lar; plants dioicous or very rarely monoicous; tome teeth grouped in eight pairs, united at tips, hygrophytes................................. 3. Philonotis smooth on outer (dorsal) surfaces; Arctic and Род 1. Plagiopus Brid. — Плагиопус mountain tundras .....................4. Conostomum — Plants green to yellowish, brownish or glaucous- М.С. Игнатов, Е.А. Игнатова green; leaves soft or rigid, spreading, falcate, Растения средних размеров, в более или менее flexuose or incurved, loosely appressed when dry, густых дерновинках, темно- или буровато-зеленые. arranged in multiple rows or indistinctly in three Стебель с 1–2 подверушечными побегами, уме- or five spiral rows; opercula convex to low-con- ренно густо трехрядно облиственный, без цент- ic; peristome double, single, or absent, exostome рального пучка, с гиалодермисом. Листья сухие teeth not paired, free at tips, papillose to reticu- извилистые до скрученных, влажные дуговидно late on outer (dorsal) surfaces, widespread in ter- отогнутые, из несколько расширенного, но нерез- restrial and saxicolous habitats ..................... 2 ко отграниченного основания узко ланцетные, по- 2. Stems triquetrous, central strand absent; upper степенно длинно заостренные, вверху килеватые; leaf cells papillose with cuticular papillae appear- край до середины или выше отвороченный, ввер- ing as longitudinal stiolations .... 1. Plagiopus ху двуслойный и пильчатый, зубцы двойные, ниже 16 BARTRAMIACEAE S t c H s S t f S t f C s F H h C b Рис. 4. Plagiopus oederianus: Hs, h 6; F 25; Stc 285; Stf 317; Cs, m, b 317. C m простые, ближе к основанию край цельный; жил- стом с высокой базальной мембраной, широкими ка оканчивается немного ниже верхушки листа, сегментами, реснички короткие или отсутствуют. вверху на нижней стороне пильчатая, на попе- Тип рода – Plagiopus serratus Brid. (= P. oede- речном срезе слабо дифференцированная, иногда rianus). Род включает три вида, имеющих сильно с двумя указателсями, выделяющимися более круп- дизъюнктивное распространение. Название от ными размерами; клетки в верхней части листа pl£gioj – коcой, poÚj – нога (греч.), относится к коротко прямоугольные и квадратные, толстостен- коробочке, косо сидящей на ножке. ные, с мелкими продолговатыми кутикулярными Plagiopus collections often have sporophytes, and папиллами, имеющими вид продольной исчер- their capsules have a distinctive, diagnostic shape that ченности. Специализированные органы вегетатив- immediately points
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