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15The genus Oxymitra (Marchantiophyta) in Russia
A. D. Potemkin1, Yu. A. Rebriev2
'Komarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg, Russia 2Southern Scientific Center of the Russian Academy of Sciences, Rostov-on-Don, Russia Corresponding author. A. D. Potemkin, potemkin_alexey@binran.ru
Abstract. Differentiation, ecology and conservation and collecting issues are discussed for the genus and species Oxymitra incrassata obtained from the Rostov Region as new for Russia. Its description, variation, macro- and microphotographs and photograph of preferred habitat are provided. SEM study and images of spores display their characteristically smooth surface resulting in a shiny appearance.
Keywords: Oxymitra incrassata, conservation, differentiation, ecology, morphology, SEM, spores, European Russia, Rostov Region.
Род Oxymitra (Marchantiophyta) в России А. Д. Потемкин1, Ю. А. Ребриев2
'Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия 2Южный научный центр РАН, Ростов-на-Дону, Россия Автор для переписки: А. Д. Потемкин, potemkin_alexey@binran.ru
Резюме. Обсуждаются вопросы дифференциации, экологии, особенностей сбора и охраны недавно выявленного в Ростовской обл. нового для России рода и вида Oxymitra incrassata. Приведены описание, характеристика изменчивости, макро- и микрофотографии и фотографии среды обитания; выполненное СЭМ исследование строения спор выявило их сглаженную поверхность, обусловливающую их блеск.
Ключевые слова: Oxymitra incrassata, морфология, отличия, охрана, споры, СЭМ, экология, Европейская Россия, Ростовская область.
The genus Oxymitra Bisch. ex Lindenb. (Oxymitraceae Müll. Frib. ex Grolle, Ricciales Lindenb.) includes two species: the widespread O. incrassata and the South African O. cristata Garside. American plants of Oxymitra incrassata were distinguished by Howe (1914a) as a separate species, O. androgyna Howe, a status that was not confirmed by subsequent studies (Schuster, 1992). The species of Oxymitra are xero-thermophytes, occurring in habitats subjected to long periods of desiccation.
Oxymitra incrassata was first reported from adjacent to the Russia territory of Ukraine about 75 years ago (Zerov, 1964). At present it is known in Ukraine from 12 squares (Zerov, 1964; Maslovsky, 2017). Also it was recorded from a location from Mongolia adjacent to Russia (Schubert et al, 1977), which was cited by Abramov and Abramova (1983). On this basis, Oxymitra was listed as a provisional genus for
https://doi.org/10.31111/nsnr/2019.532.417
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the liverwort flora of Russia (Potemkin, Sofronova, 2009). With these nearby records, finding a Russian occurrence was viewed as highly probable. The genus Oxymitra was indeed recently recorded as new for Russia on the basis of a collection made by the second author in April, 2017 in the Rostov Region (Ellis et al, 2019). A subsequent targeted search of O. incrassata by both authors in the southern European Russia area was unsuccessful. The first record of Oxymitra incrassata in Russia, the heretofore lack of its description and illustrations in the Russian literature as well as the necessity to attract attention to its further search in Russia persuade us to discuss separately its morphology, ecology and distribution, provide photographs and appropriate references to facilitate further finds of O. incrassata in Russia.
Material and Methods
Material was collected at the end of April, 2017 by Rebriev and identified by Potemkin. Photomicrographs of pores were obtained with light microscope Lomo Mic-med2 equipped with camera Nicon D5100. The pore images were combined from several optical sections using the software package HeliconFocus 3.0. The other photomicrographs were made by camera Olympus Tough TG-3 using its macro- and stacking modes. For SEM study spores were placed on stubs without any pretreatment, and images were taken using a JEOL JSM-6390LA scanning electron microscope.
Description is based on specimens examined and extended on the basis of the treatments and illustrations by Müller (1951-1958), Zerov (1964), Schuster (1992), Potemkin, Sofronova (2009), Campos-Sandoval, Campos-Sandoval (2017), World Flora Online (2018). Data from the literature not confirmed by the specimen study are provided in square brackets in the species description.
Results and Discussion
Oxymitra incrassata (Brot.) Sergio et Sim-Sim, 1989, J. Bryol. 15: 662. = Riccia incrassata Brot., 1804, Fl. Lusit. 2: 428.
= Oxymitra paleacea Bisch., 1829, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 14(2, Suppl.).
= Tesselina pyramidata (Willd.) Dumort., 1822, Comment. Bot.: 78. = Riccia pyrami-data Willd., 1788. Bot. Mag. (Römer & Usteri) 2(4): 9.
(Plates I: 1; II; III; IV)
Thalli 2.5-5.5[7] mm wide and up to 10 mm long, dull green to yellowish green and cinnamon, dark brown or tinged purple, simple or 2-[3 times branched and then often forming hemi-rosettes], often convolute when dry and then covered by bleached scales, with remarkable deep median groove particularly in wet sterile thalli, which becomes indistinct in dry thalli, in cross section somewhat wider than high or higher than wide, with narrow tall unistratose air chambers. Epidermal cells thin-walled, pores ± stellate, bounded by 5-6[7] cells with ± thickened internal and radial walls. Ventral scales large and ± extending beyond thallus margins, most remarkable in female and convolute dry plants, bleached in distal part and brown or purplish in proximal part mostly, gradually narrowed in narrowly triangular ciliate apex, without ap-
pendages and oil bodies. Dioicious or monoicous. Antheridia and archegonia situated along median groove on separate thalli, [intermingled with each other or situated on separate branches]. Antheridia immersed in the thallus with ostioles projected above median groove. Archegonia on surface of median groove enclosed in becoming cinnamon [or purple] prominent pyriform involucres with air chambers and a short beak, [bounded by inconspicuous white filiform scales]. Mature capsules enclosed in strongly projecting involucres in 1-2(3) rows. Spores black opaque and shiny, 80-200 per capsule, (90)100-140 |im in diam. in European plants [and up to 180 |im in American plants]. Spore structure invisible in light microscope because of deep black pigmentation. Spore surfaces nearly smooth in SEM images, distal face with 4-5(6) shallow areoles across diam., (15)23-40(48) |im in diam., proximal surface smooth, triradiate mark distinct. [Spores dispersed in periods of rains due to the floating ability of the involucre with air chambers]. Elaters absent.
Note on spore surface structure. Provided by Sergio and Sim-Sim (1989: Fig. 3) SEM spore images from a specimen collected by F. Welwitch, which was selected as neotype of Oxymitra incrassata, were clarified by a solution of sodium hypochloride that resulted in a change of the spore surface structure which became tuberculate with acute ribs of meshes. Spores studied by us were mounted on a SEM stub without pretreatment and have a smooth surface structure and rounded ribs of meshes. As it was shown by our study spores of O. incrassata in collections over 100 years old are often destroyed or covered by fungal hyphae seen also in: Fig. 3, a, d by Sergio, Sim-Sim (1989). Such spores are unsuitable for proper investigation.
The spores, drawn and described by Schuster (1992: Fig. 982: 1) were redrawn from Hassel de Menendez (1962: Fig. 74, G), and also have acute ribs defining the areolae and show the surface with verrucae (0.2-0.5)2-5 |im high. Such a description may result also from some pretreatment of spores to facilitate observation with a light microscope. Schuster (1992) describes spores as shiny, which completely corresponds to our observations and can be observed at Plate III: 5. The shiny structure is correlates with the smooth spore surface shown in Plate IV.
Specimens examined: Russia, Rostov Region, Orlovsky District, Rostovsky State Nature Reserve, Starikovsky plot, second flood plain terrace of Manych River, ca. 46.538211°N, 42.890220°E, upper part of gentle south-faced valley steppe slope with dominance of Sti-pa lessingiana, Festuca valesiaca, Agropyron desertorum, Artemisia lerchiana and abundant Artemisia santonica, Galatella villosa, Iris pumila, Limonium gmelinii, in seasonally moist probably drying in summer stria, on chestnut somewhat saline soil, plants with androecia, 26 IV 2014, Yu. Rebriev, LE; Italien, Sondrio im ville Tellina (Lombardei)... auf sandiger Erde und Zwischen trockenen Felsen der Weinberge, 360-400 m, 17 IX 1901, E. Levier (V. Schiffner. Hepaticae europaeae exsiccatae 1168), LE; France, Frankreich, Rosquehaute (Dep. Herault), auf sandigem Diluvium und auf vulkanischem Boden, a) Nov. 1902, b) Dez. 1902, A. Crozals (V. Schiffner. Hepaticae europaeae exsiccatae 1167a, b), LE; Austria inferior: ad terram nudam ad "Rothenhof" prope Stein a. D. J. Baumgartner (Kryptoga-mae exsiccatae 181), LE; Niedorösterreich, Südhänge (Schiefer) oberhalb Rotenhohof bei
Plate I. Oxymitra incrassata (1) and its habitat (2) in the Rostov Region, by Yu. A. Rebriev.
Scale bar: 5 mm.
Plate II. Oxymitra incrassata. 1-3 — pores; 4, 5 — ventral scales; 6 — wet thallus; 7— cross section of old part of male thallus with anteridial chamber and ostioles above. All from 26 IV 2014, Rebriev, LE, by A. D. Potemkin. Scale bar: 1-3 — 17 |m; 4, 5 — 375 |m; 6 — 1400 mm; 7 — 700 |im.
Plate III. Dry plants of Oxymitra incrassata. 1-2 — from the Rostov Region (26 IV 2014, Yu. Rebriev, LE); 3-4 — from Texas, USA (4 VIII 1991, G. Merrill 13338, LE); 5 — from Austria, thallus with detached involucre and spores in thallus cavity below involucre (Apr. 1903, J. Baumgartner, V. Schiffner (V. Schiffner. Hepaticae europaeae exsiccatae 1170), LE; 6 — from Crete Island, Greece, thallus with mature involucres and mature capsules inside (16 III 2007, Frahm K-175, LE), all by A. D. Potemkin. Scale bar: 1-4 —1.5 mm; 5 — 425 |im; 6 — 540 |im.
Plate IV. Spores of Oxymitra incassata. 1 — distal surface; 2 — proximal surface; 3— lateral aspect; 4 — fragment of distal surface. All from 16 III 2007, Frahm K-175, LE, by A. D. Potemkin and L. A. Kartseva.
Stein a. d. Donau — 350 m, Apr. 1903, J. Baumgartner, V. Schiffner (V. Schiffner. Hepaticae europaeae exsiccataeae 1170), LE (with mature capsules); Greece, Griechenland — Kreta (Crete Island), ... 35.30598°N, 23.93681°E, 1250-65 m, Zypressenwälder auf Kalk, 16 III 2007, Frahm K-175, LE (with mature capsules); USA, Texas, Burnet County, mesquite-oak savanna with granite outcrops on slope SE of lake, Inn Lake State Park, 30°44'N, 98°22'E, on thin soil on level of rock outcrop in full sun, 4 VIII 1991, G. Merrill 13338, R. Stotler, LE.
Illustrations. Drawings: Zodda, 1934, Fig. 62; Müller, 1951-1958, Fig. 94; Häs-sel de Ménendez, 1962, Figs. 74, 75; Zerov, 1964, Fig. 46; Augier, 1966, Fig. 24: 28; Schuster, 1992, Figs. 981-982; Casas et al., 2009, Fig. 6: 10, 11; Potemkin, Sofrono-va, 2009, Figs. 15, 43d, 45 at pages 160, 176, 178; photographs: Howe, 1914b, Fig. 1; Campos-Sandoval, Campos-Sandoval, 2017: Figs. 1, 2; Information., 2007; see also photographs on Internet.
Distribution. Oxymitra incrassata has a disjunct xerothermic Pangean distribution and is widely distributed in the Mediterranean region of Europe and North Africa, ranging northward to Central Europe, southward to the Canary Islands and eastward
to Jordan and Socotra, southern Arabian Peninsula, Central South America (Uruguay, Paraguay, Northern Argentina, Brazil), central México and south central USA (Frye, Kürchner, 1988; Campos-Sandoval, Campos-Sandoval, 2017; Gradstein, 2017; Ellis et al, 2019), Australia (Information., 2007).
Ecology. Xerothermophyte. On fine-grained compacted soil, preferably over limestone exposures, subject to long periods of desiccation in Europe (DierBen, 2001), apparently absent from calcareous soils in western North America (Schuster, 1992). In the Rostov Region in upper part of gentle south-faced valley steppe slope (Plate I: 2) in seasonally moist probably drying in summer stria, on chestnut somewhat saline soil, shallow siliceous soil over a wet ledge in quartzite west faced rocky slope in Spain (Campos-Sandoval, Campos-Sandoval, 2017). See labels of specimens examined for details.
Collecting notice. Habit of Oxymitra incrassata depends greatly on the state of thalli. Extremely dry sterile thalli become convolute with bleached ventral scales covering the thallus (Plate III: 3-4), female plants may be less convolute (Plate III: 5-6). Dry plants of Oxymitra may be hardly discernible on dry soil that causes difficulties for their collection. In wet conditions, thalli of O. incrassata become ± plane and their scales may be hardly discernible, in sterile thalli particularly (Plates I: 1; III: 1-2). Due to the tender structure of thalli they may absorb water ± quickly (Plate II: 6) and apparently use early morning dew as a source of water. Therefore, early morning time for collecting is preferable.
Variation and differentiation. Plants from the Rostov Region were easily distinguished due to characteristic deep median groove and distinctive ventral scales gradually narrowed in bleached narrowly triangular apices ± extending beyond thal-lus margins.
When sterile, O. incrassata may be mistaken for a species of Riccia L. from which it differs by the distinctive deep median groove, the presence of ventral scales and stellate pores. In the latter two characters Oxymitra incrassata recalls species of Athalamia Falc., which have thalli without median groove, considerably broader than high, and broader often bistratose air chambers.
Distinctions of European and American plants of O. incrassata in spore size and ecological requirements are not constant and overlap that support recognizing them as one species (Schuster, 1992: 407).
Conservation. Despite the broad range, O. incrassata is quite rare and had been considered a candidate for a new European Red List (Schumacker, Vana, 2005; Maslovsky, 2017). Even in detailed bryophyte studies in Europe it is found in a few locations as it was shown by Campos-Sandoval, Campos-Sandoval (2017). However, it is listed in "European Red List of mosses, liverworts and hornworts" (Hodgetts et al., 2019) with status LC (Least concern). Nevertheless its state in Russia corresponds to IUCN criteria B2a and D1 (IUCN..., 2012). Taking into account these facts and the establishment of O. incrassata at its eastern limit of distribution in Europe it is recommended to include it in forthcoming Red Data books of Russia and the Rostov Region.
Acknowledgments
John J. Engel and James R. Shevock are thanked for careful reading of the manuscript and correcting the English. The valuable advices of Evgeny A. Borovichev and help of Mrs. Lyudmila A. Kartseva in SEM study are gratefully appreciated. The study by A. D. Potemkin was carried out in the framework of the institutional research project of Komarov Botanical Institute of the Russian Academy of Sciences "Flora and systematics of lichens and bryophytes of Russia and phytogeographical-ly important regions" (AAAA-A19-119020690077-4) and the work of Yu. A. Rebriev was supported by government assignments for South Science Center RAS (project AAAA-A19-119011190176-7 "Structural and functional organization and dynamics of plane landscape biocoenoses of the south part of Russia in the conditions of climate change and anthropogenic impact"). The research was done using equipment of The Core Facility Center "Cell and Molecular Technologies in Plant Science" at the Komarov Botanical Institute RAS (St. Petersburg, Russia).
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Литература
[Abramov, Abramova] Абрамов И. И.. Абрамова А. Л. 1983. Конспект флоры мхов Монгольской Народной Республики. Л.: 222 с.
Augier J. 1966. Flore des Bryophytes. Morphologie, Anatomie, Biologie, Ecologie, Distribution, Géographie. Paris: 702 p.
Campos-Sandoval J. Á., Campos-Sandoval A. 2017. First recorded locality for the liverwort Oxymi-tra incrassata (Brot.) Sérgio & Sim-Sim (Marchantiales: Oxymitraceae) in the province of Málaga (Spain). BV news Publicaciones Científicas 6(69): 5-9. https://www.biodiversidadvirtu-al.org/taxofoto/sites/default/files/bv_news_publicaciones_cientificas_volumen_6.pdf (Дата обращения: 9 VII 2019).
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Dierßen K. 2001. Distribution, ecological amplitude and phytosociological characterization of European bryophytes. Bryophytorum Bibliotheca 56: 1-289.
Ellis L. T., Amélio L. A., Peralta D. F., Backor M., Baisheva E. Z., Bednarek-Ochy-ra H., Burghardt M., Czernyadjeva I. V., Kholod S. S., Potemkin A. D., Erdag A., Kirmaci M., Fe-dosov V. E., Ignatov M. S., Koltysheva D. E., Flores J. R., Fuertes E., Goga M., Guo S.-L., Hofbauer W. K., Kurzthaler M., Kürschner H., Kuznetsova O. I., Lebouvier M., Long D. G., Mamon-tov Yu. S., Manjula K. M., Manju C. N., Mufeed B., Müller F., Nair M. C., Nobis M., Norhaz-rina N., Aisyah M., Lee G. E., Philippe M., Philippov D. A., Plásek V., Komínková Z., Por-ley R. D., Rebriev Yu. A., Sabovljevic M. S., de Souza A. M., Valente E. B., Spitale D., Sri-vastava P., Sahu V., Asthana A. K., §tefänut S., Suárez G. M., Vilnet A. A., Yao K.-Y., Zhao J.-Ch. 2019. New national and regional bryophyte records, 59. Journal of Bryology 41(2): 177-194. https://doi.org/10.1080/03736687.2019.1613112
Frey W., Kürschner H. 1988. Bryophytes of the Arabian Peninsula and Socotra. Floristics, phytoge-ography and definition of the Xerothermic Pangaean element. Studies in Arabian bryophytes 12. Nova Hedwigia 46: 37-120.
Gradstein S. R. Amphitropical disjunctive species in the complex thalloid liverworts (Marchantii-dae). Journal of Bryology 39(1): 66-78. https://doi.org/10.1080/03736687.2016.1189662
Hassel de Menéndez G. G. 1962. Estudio de las Anthocerothales y Marchantiales de la Argentina. Opera Lilloana 7: 1-297.
Hodgetts N., Calix M., Englefield E., Fettes N., Garcia Criado M., Patin L., Nieto A., Bergamini A., Bisang I., Baisheva E., Campisi P., Cogoni A., Hallingback T., Konstantinova N., Lockhart N., Sabovljevic M., Schnyder N., Schrock C., Sergio C., Sim-Sim M., Vrba J., Ferreira C. C., Afonina O., Blockeel T., Blom H., Caspari S., Gabriel R., Garcia C., Garilleti R., Gonzalez Mancebo J., Goldberg I., Hedenas L., Holyoak D., Hugonnot V., Huttunen S., Ignatov M., Ignatova E., Infante M., Juutinen R., Kiebacher T., Kockinger H., Kucera J., Lon-nell N., Lüth M., Martins A., Maslovsky O., Papp B., Porley R., Rothero G., Soder-strom L., §tefänut S., Syrjanen K., Untereiner A., Vána J. 1, Vanderpoorten A., Vellak K., Alef-fi M., Bates J., Bell N., Brugues M., Cronberg N., Denyer J., Duckett J., During H. J., Enroth J., Fedosov V., Flatberg K.-I., Ganeva A., Gorski P., Gunnarsson U., Hassel K., Hespan-hol H., Hill M., Hodd R., Hylander K., Ingerpuu N., Laaka-Lindberg S., Lara F., Mazimpa-ka V., Mezaka A., Müller F., Orgaz J. D., Patino J., Pilkington S., Puche F., Ros R. M., Rumsey F., Se-garra-Moragues J. G., Seneca A., Stebel A., Virtanen R., Weibull H., Wilbraham J., Zarnowiec J. 2019. A miniature world in decline: European Red List of Mosses, Liverworts and Hornworts. Brussels: 88 p.
Howe M. 1914a. Further observations on the Texan Oxymitra (Tesselina). The Bryologist 17(6): 92-94. https://doi.org/10.2307/3238893
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