Botanica Pacifica. A journal of plant science and conservation. (2012) 1, 31–95 Scapania and Macrodiplophyllum in the Russian Far East Seung Se CHOI1, Vadim A. BAKALIN2, 3* & Byung­Yun SUN1 ABSTR A C T Seung Se Choi The genera Scapania and Macrodiplophyllum have been studied in the Russian Far 1 Faculty of Life Sciences East. Mac ro diplophyllum is represented by three species that are known in this ge­ Chonbuk National University nus. Scapania includes 38 species that are distributed among 12 sections. Keys are Jeonju, Jeonbuk Province, South Korea provided for all taxa found in the Russian Far East. All species based on the Far E­mail: [email protected] Eastern material are illustrated. Some corrections have been made in respect to the infrageneric structure of Scapania. Spe cifically , section Brevicaules R. M. Schust. Byung­Yun Sun 1 Same institution was united with section Apiculatae H. Buch, S. sphaerifera H. Buch was transferred E­mail: [email protected] to section Aequilobae Müll. Frib., and accordingly, section Spheriferae Konstant. & Po temkin was synonymized with the latter. The understanding of the distributions Vadim A. Bakalin of some species has been improved. S. parvidens Steph. was reevaluated at the 2 Botanical Garden­Institute FEB RAS spe cies level. S. integerrima Steph., S. ligulata Steph. and S. ampliata Steph. are now Vladivostok, 690024, Russia excluded from the hepatic flora of Russia.Scapania magadanica S. S. Choi, Baka lin & B.­Y. Sun sp. nov. is described based on the combination of paroicous in flo­ 3 Institute of Biology and Soil Science res cence, green to colorless two­celled gemmae, a rounded apex of the dorsal FEB RAS lobe, and a non­decurrent ventral lobe, all of which are not present in other taxa Vladivostok, 690022, Russia Email: [email protected] of the genus. K e y w o r d s : Scapania, Macrodiplophyllum, Russian Far East, Scapaniaceae, taxonomy, Hepaticae * Corresponding author Чои С.С., Бакалин В.А., Сан Б.-Ю. Scapania и Macrodiplophyllum на российском Дальнем Востоке Manuscript received: 02.04.2012 Проведена обработка родов Scapania и Macrodiplophyllum для российского Review completed: 24.05.2012 Дальнего Востока. Macrodiplophyllum представлен 3 видами, охватывающи­ Accepted for publication: 15.07.2012 ми все разнообразие рода. Scapania включает 38 видов, объединенных в 12 секций. Составленные ключи охватывают все таксоны, представленные на российском Дальнем Востоке. Для видов, изученных по дальневосточ­ ным материалам, составлены оригинальные иллюстрации. Внутриродо вая структура рода Scapania подверглась некоторой переработке. Секция Bre­ vicaules R. M Schust. объединена с секцией Apiculatae H. Buch, S. sphaerifera H. Buch перенесена в секцию Aequilobae Müll. Frib. и, таким образом, секция Spheriferae Konstant. & Potemkin была сведена в синонимы с последней. Рас­ пространение большинства таксонов было существенно уточнено. Восста­ новлен видовой ранг Scapania parvidens Steph. S. integerrima Steph., S. ligulata Steph. и S. ampliata Steph. исключены из флоры печеночников российско­ го Дальнего Востока. Scapania magadanica S. S. Choi, Bakalin & B.­Y. Sun sp. nov. опи­­сана на основе пареции, зеленых до бесцветных выводковых почек, закруг ленной верхушки дорзальной лопасти и не сбегающей вентральной лопасти, ­ комбинации, ранее неизвестной внутри рода Scapania. Ключевые слова: Scapania, Macrodiplophyllum, российский Дальний Восток, Scapaniaceae, таксономия, Hepaticae INTRODUCT I O N and Macrodiplophyllum, which are taxonomically diverse in The Russian Far East covers more than 3 millions northern latitudes, including Arctic and Subarctic Asia km2, lying between 42ºN and 71°N, and is characterized and North America, with noticeable diversity in East Asia by no ti ceable diversity in floristic composition, orography, and the Himalayas, 2) and Diplophyllum, which is diverse in and climate. Scapania and Macrodiplophyllum, as well as the the tropics, descending into the Antipodes. Scapania and more remotely related Diplophyllum, are called "scapanioid" Macrodiplophyllum include 41 species (38 and 3, respectively) genera, which are taxonomically diverse in the Russian Far in Far Eastern Russia, and represent approximately 12% East. This group was previously circumscribed as the oli ­ of the total known hepatic taxonomic diversity in this area. gogeneric family Scapaniaceae s. str. (e.g., Schuster 1974, Only five species of Diplophyllum are found in the Russian Schljakov 1981), which has since been considerably enlarged Far East, and with the exception of D. taxifolium (Wahlenb.) when many former members of the Lophoziaceae were Dumort., which is common throughout in mountainous included (e.g. Vilnet et al. 2010). It is represented by two areas, are sporadically distributed. We restricted the present strongly phytogeographically detached clades: 1) Scapania study to examining Macrodiplophyllum (including all three of the globally recognized species) and Scapania. ©Botanical Garden-Institute FEB RAS. 2012 31 Choi, Bakalin & Sun Taxonomy of the genus Scapania is controversial. For Key to scapanioid genera of the Russian Far East example, some authors (Potemkin 1999, 2002) include Mac­ 1. Perianth usually strongly flattened, truncate, lacking plicae; gem­ ro diplophyllum in this genus, whereas others (Buch 1928) mae usually elliptical to spherical, smooth; keel usually sharp and divide Scapania s. str. into two genera and treat the subgenus acute throughout .................................................................... Scapania. Scapaniella as a separate genus. Currently, Scapania and Mac­ 1. Perianth usually weakly flattened, contracted, plicate; gemmae usually angular to stellate, rarely elliptical (Scapania sect. Kaalaasia); ro diplophyllum are accepted by the majority of bryologists keel usually obtuse or leaves are canaliculate; leaf base commonly (Kon­­stantinova et al. 2009, Crandall­Stotler et al. 2009, sheathing .............................................................................................. 2. Heinrichs et al. 2012) at the generic level. Thirty eight spe­ 2. Leaves evidently spreading from the base; gemmae elliptical ....... cies from our study area were placed into 12 sections. ........................................................................... Scapania sect. Kaalaasia. In total, 67 species were reported from eastern extra tro­ 2. Leaves with sheathing base; gemmae usually angular, stellate ... pical Asia due to their appearances in floristic papers (Piippo .......................................................................................................... 3. 1990, Hong 2003, Yamada & Iwatsuki 2006, Yamada & Choi 3. Plants larger 2.5–4.5 mm wide, robust; gemmae 2–4 celled; 2007, Zuo & Cao 2007a, Bakalin 2010), and approximately trigones usually convex ............................................ Macrodiplophyllum. 30 of these were described from this area (Stephani 1910, 3. Plants smaller 1.0–3.5 mm wide, not robust; gemmae 1–2 celled; trigones usually concave ................... Diplophyllum [not treated here]. Horikawa 1934, Amakawa & Hattori 1953, 1954, 1955, Po temkin 2000, 2001, Zuo et al. 2007b, 2007c). For this variable and diverse territory, some regional treatments, SCAPANIA (Dumort.) Dumort. Recueil Observ. such as the treatment of Japanese Scapaniaceae (Amakawa Jungerm. 14, 1835 nom. cons. & Hattori 1953, 1954, 1955), which is somewhat outdated, Martinellius Bennet & Gray, ex S. F. Gray, Nat. Arr. Brit. as well as some attempts to describe the Chinese Scapania Pl. I: 691, 1821. (Po temkin 2000, 2001, Zuo & Cao 2007a, Zuo et al. 2007b, Plants 6.0–150.0 × 0.5–35.01 mm, 2007c) were published. Scapania and Macrodiplophyllum were Description: pro­­­strate to ascen ding and erect, from yellowish green never examined specifically for the Russian Far East, but to brown and purplish to red­purple. sparsely books such as those by Schljakov (1981) and Buch (1928) Stems branched, dorsal surface yello w ish green to greenish pale include very valuable information on the distribution and brown, ventral surface not differs­­ from dorsal one to deep taxonomy of these genera in the Russian Far East, as reddish brown and fuscous brown, in cross section round­ these authors examined approximately 85% of the present ed to transversely elliptic, with thick­walled cortex cells species diversity within the area. Bakalin (2004) has recently in 1–3(–5) layers. sparse to numerous, isolate reviewed Macrodiplophyllum for Russia and published the Rhizoids to fasciculate and spreading at right angle from the stem, most recent information on the distribution of Scapania colorless to pale brown. unequally bilobed, usually and Macrodiplophyllum in this area (Bakalin 2009, 2010). He Leaves conduplicate, contiguous to im bri cate, rarely canaliculate; has reported 43 species from the Russian Far East (three keel (0.1–)0.2–0.7(–0.8) of vent ral lobe length2; dorsal lobe of which were excluded and one was added in the present transversely inserted, not or distinctly decurrent, obliquely paper), a number representing more than half of the East ovate to lanceolate or rounded­quadrate, 0.5–1.0 of ventral Asian diversity of that group, and including both Arctic lobe size; ventra l lobe transversely inser ted, insertion line taxa (e.g., S. simmonsii, S. spitsbergensis) and East Asian ones obscurely to dis tinctly arched, not or distinctly decurrent, (sect. Stephania). obliquely oblong to obovate or rounded, margin in both The present account is largely based on a critical re­