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15Calypogeia azurea (Calypogeiaceae, Marchantiophyta) in the Northwestern European Russia
A. D. Potemkin1, E. A. Borovichev2,3, E. G. Ginzburg1
1 Komarov Botanical Institute, Professor Popov Str., 2, St. Petersburg, 197376, Russia; potemkin_alexey@binran.ru 2 Institute of Industrial Ecology Problems of the North of the Kola Science Centre of RAS, Fersmana Str., 14a, Apatity, Murmansk Region, 184209, Russia; borovichyok@mail.ru 3 Forest Research Institute of Karelian Research Centre of RAS, Pushkinskaya Str., 11, Petrozavodsk, Republic of Karelia, 185910, Russia
Abstract. Calypogeia azurea is reported for the first time in the Leningrad Region and for the second time for the Republic of Karelia. Description of C. azurea based on alive collections from the Leningrad Region and the Republic of Karelia, and their photographs are provided. The variability of the species in studied specimens is rather broad and corresponds to the plants from Great Britain and Nordic countries. C. azurea is known in Russia from alive collections from the Northwestern European Russia, Caucasus, South Urals, South Siberia and South Kuril Islands. Ecology and differentiation of C. azurea are discussed. The revision of all kept in LE specimens from Russia, Latvia, Finland and Georgia identified as C. trichomanis, nom. rej., has shown that most of them cannot be attributed to C. azurea or C. muelleriana, which usually correspond to this rejected name. Except two specimens from Finland and one from Georgia tentatively attributed to Calypogeia cf. azurea, they represent mostly materials of C. integristipula, partly of C. neesiana and exceptionally of C. suecica and C. fissa. LE older collections of C. trichomanis from Caucasus are C. fissa, from Arctic Siberia and Arctic Far East are C. muelleriana.
Keywords: Calypogeia azurea, Calypogeia trichomanis, blue oil bodies, distribution, taxonomy, variation, Leningrad Region, Republic of Karelia, Russia.
Calypogeia azurea (Calypogeiaceae, Marchantiophyta) на северо-западе европейской части России
А. Д. Потемкин, Е. А. Боровичев, Э. Г. Гинзбург
1 Ботанический институт им. В. Л. Комарова РАН, ул. Профессора Попова, д. 2,
197376, Санкт-Петербург, Россия; potemkin_alexey@binran.ru
2 Институт проблем промышленной экологии Севера КНЦ РАН, ул. Ферсмана,
д. 14а, г. Апатиты, Мурманская обл., 184209, Россия; borovichyok@mail.ru 3 Институт леса КарНЦ РАН, Пушкинская ул., д. 11, Петрозаводск, Республика Карелия, 185910, Россия
Резюме. Calypogeia azurea впервые достоверно приводится на основании прижизненных коллекций для Ленинградской области и во второй раз для Республики Карелия. Приведены описание и фотографии C. azurea, выявленной в
прижизненном состоянии в Ленинградской области и Республике Карелия. В образцах с северо-запада России вид характеризуется широким диапазоном изменчивости, сопоставимым с описанным у растений из Великобритании и Скандинавии. Обсуждены экология и отличия C. azurea. Вид достоверно известен в России с северо-запада европейской части России, Кавказа, Южного Урала, Южной Сибири и с Южных Курильских островов. Ревизия всех хранящихся в гербарии Ботанического института (LE) образцов, определенных как Calypogeia trichomanis, nom. rej., показала, что большинство из них не может быть отнесено к C. azurea или С. muelleriana, c которыми обычно соотносят это отвергнутое название. Большинство имеющихся образцов из Лениградской обл. и Латвии относятся к C. integristipula, часть к C. neesiana, единичные образцы к C. fissa и С. suecica. Все образцы с Кавказа относятся к C. fissa, из арктических Сибири и Дальнего Востока к C. muelleriana и лишь два образца из Финляндии и один из Грузии могут быть условно отнесены к C. azurea.
Ключевые слова: Calypogeia azurea, Calypogeia trichomanis, изменчивость, распространение, синие масляные тела, таксономия, изменчивость, Ленинградская область, Республика Карелия, Россия.
The name Calypogeia trichomanis (L.) Raddi, nom. rej., since its treatment by K. Müller (1947) and until recently was used for plants with blue oil bodies. Oil bodies in this species are fugacious and need to be studied when plants alive that often makes identification of this species in collections without oil bodies untenable. Before Müller (1947) this name was applied to various species of the genus that resulted from misleading description of Mnium trichomanis L., nom. rej., as «Mnium foliis distichis integerrimis» (Linne, 1753: 1114) referring to not informative illustrations of Dillenius (1741: 236, Plate XXXI) (Fig.). The broad circumscription of C. trichomanis in Synopsis Hepaticarum by Gottsche et al. (1844: 198) with inclusion of infraspecific taxa with various shape of leaves and underleaves worsen the situation.
Despite Stotler and Crotz (1983) have shown that majority of older reports of C. trichomanis in both Europe and eastern North America belong to C. muelleriana the data on distribution of C. azurea in Russia until present remain uncertain because most of them are not annotated by data on oil bodies or not published. Following Stotler and Crotz (1983), Kon-stantoinova et al. (1992) and Potemkin, Sofronova (2009) noted that old records of C. azurea (as C. trichomanis) based on identification of materials without oil bodies might be erroneous and based on specimens of C. muelleriana. Konstantinova, Bakalin et al. (2009: 36) mentioned also that C. azurea is restricted to subalpine belt of mountains and the middle and south taiga, and do not occur in the Far North. Revision of LE collections labelled as C. trichomanis has shown their various identifications (see Results).
Stotler and Crotz (1983) have shown that C. trichomanis sensu K. Müller and modern authors cannot be correlated with any previous legitimate name and needs to be replaced by name C. azurea Stotler et Crotz which has limited occurrence in Europe and North America.
Buczkowska and coauthors (Buczkowska, 2004; Buczkowska et al., 2004) made biometrical analysis of morphological and anatomical characters of the alive plants of European species of the genus Calypogeia Raddi with oil bodies in Poland. The study was also combined with the isoenzyme analysis and demonstrated a distinct segregation of C. azurea from the other European species at the species level and from American C. peruviana Nees et Mont. Those results are supported by recent molecular studies (Buczkowska, pers. com. 19.04.2017).
The facts above and recent finds of C. azurea in the Leningrad Region and the Republic of Karelia persuade us to provide its description on the basis of collections identified with oil bodies, to analyze and discuss its distinctive characters and differentiation, and generalize all known information on its distribution and ecology in Russia.
Material and Methods
All materials of Calypogeia trichomanis kept in Herbarium of Koma-rov Botanical Institute, St. Petersburg (LE) from different regions of Russia and some adjacent territories, Latvia, Finland and Georgia mostly (34 specimens cited below) have been revised.
Data on C. azurea from the Northwestern European Russia are based on identified with oil bodies collections from the Leningrad Region and the Republic of Karelia, which certify correct identification of this still poorly known in Russia species. Three specimens with oil bodies were found and identified in the area. Specimens of C. azurea in the Leningrad Region were collected in minigrip (= ziplock) plastic bags to prevent desiccation.
Photographs of alive plants from the Leningrad Region have been made by Olympus Tough TG-3 and Nikon Coolpix 7900.
Data on specimens deposited in KPABG and INEP are available in CRIS (Cryptogamic Russian Information System, http://kpabg.ru/ cris/?q=node/16).
Specimens examined
Calypogeia azurea (3 specimens identified with oil bodies). Russia. Republic of Karelia, Kondopoga District, Kivach State Nature Reserve, W part of the reserve, 2 km NW of the administration place, 62°16'58"N, 33°58'14"E, 95 m alt., spruce forest near Suna River, peaty bank of the river, with Fuscocephaloziopsis lunulifolia
(Dumort.) Vana et L. Söderstr., 26.06.2015, E. A. Borovichev BE19-45-15, INEP; Petrozavodsk City, Sulazhgora Lake, spruce forest with Oxalis acetosella L., on the hummock, 21.08.1998, V. A. Bakalin, KPABG. Leningrad Region, Vsevolozhsk District, 60.1814456°N, 30.4622110°E, 500 m W of Kavgolovskoe Lake, bilberry-sphagnum spruce forest, on needle litter and soil under spruce, 11.09.2016, E. G. Ginzburg 1109-3, det. A. D. Potemkin, LE.
Calypogeia cf. azurea (2 specimens identified without oil bodies as C. trichomanis, all in LE). Finland. Finlandia australis, Lojo, 22.06.1878, S. O. Lindberg (mod. parvifolia); Nylandia, Porvoo, 17.07.1937, H. Buch. Georgia. Lagodekhi Nature Reserve... 26.06.1948, A. Dolukhanov (trans. ad C. muelleriana).
Calypogeia fissa (8 specimens identified as C. trichomanis or Kantia trichomanis (L.) Gray, all in LE). Russia. Caucasus, Krasnodar Territory, Adler District. 16.07.1948, Raspopov; Sakhalin Region, Iturup Island. 23.09.1974, Blagodatskikh (C. fissa s. l.). Latvia. Livland, Kreis Riga. 21.06 (= 04.07) 1909, J. Mikotowicz, Bryotheca Baltica Nr. 2f (mod. subintegrifolia, admixture to C. neesiana). Georgia. Adzharia Republic, Batumi Botanical Garden. 03.07.1961, A. Abramo-va, I. Abramov; ibid., 17.06.1961, A. Abramova, I. Abramov; ibid., 03.07.1961, A. Abramova, I. Abramov; ibid., 18.06.1961, A. Abramova, I. Abramov (mod. densifo-lia with rather large underleaves); ibid., 20.06.1961, A. Abramova, I. Abramov.
Calypogeia integristipula (15 specimens identified as Kantia trichomanis or Calypogeia trichomanis, all in LE). Russia. Leningrad Region, on rotten stump in forest near Dno Railway Station, 07.06.1940, I. I. Abramov. Baltic countries or Russia. Musci hepatic. et frondosi Rus. Balt. Exs. I, Leif. 3, N 132. G. Girgensohn. Latvia or Estonia. Musci hepatic. et frondosi Rus. Balt. Exs. II, Abt. I, 1862-63, die Laub. Un Lebermoose Liv.-Est. und Kurlands. G. Girgensohn 56. Latvia. Kurland, Kreis Doblen. 29.06 (= 12.07) 1908, J. Mikotowicz, Bryotheca Baltica Nr. 2a; Livland, Kreis Riga. 18.05 (= 31.05) 1909, J. Mikotowicz, Bryotheca Baltica Nr. 2b; Kurland, Kreis Doblen. 29.06 (= 12.07) 1908, J. Mikotowicz, Bryotheca Baltica Nr. 2a; Livland, Kreis Riga. 18.05 (= 31.05) 1909, J. Mikotowicz, Bryotheca Baltica Nr. 2b; Kurland, Kreis Doblen. 25.06 (= 08.07) 1906, J. Mikotowicz, Bryotheca Baltica Nr. 2; Livland, Kreis Pernau. 23.07 u. 25.07 (= 05.08 u. 07.08) 1909, J. Mikotowicz, Bryotheca Baltica Nr. 2c; Livland, Kreis Pernau. 23.07 u. 25.07 (= 05.08 u. 07.08) 1909, J. Mikotowicz, Bryotheca Baltica Nr. 2c (trans. ad C. neesiana); Kurland, Kreis Doblen. 29.06 (= 12.07) 1908, J. Mikotowicz, Bryotheca Baltica Nr. 2a; Livland, Kreis Riga. 28.09 (= 11.10) 1908, J. Mikotowicz, Bryotheca Baltica Nr. 2d; Livland, Kreis Riga. 28.09 (= 11.10) 1908, J. Mikotowicz, Bryotheca Baltica Nr. 2d; Kurland, Kreis Doblen. 25.06 (= 08.07) 1906, J. Mikotowicz, Bryotheca Baltica Nr. 2 (trans. ad C. neesiana: no marginal border in apical area); Livland, Kreis Pernau. 23.07 u. 25.07 (= 05.08 u. 07.08) 1909, J. Mikotowicz, Bryotheca Baltica Nr. 2c; Livland, Kreis Riga. 18.05 (= 31.05) 1909, J. Mikotowicz, Bryotheca Baltica Nr. 2b.
Calypogeia muelleriana (2 specimens identified as Calypogeia trichomanis, all in LE). Russia. [Central Siberia, Krasnoyarsk Territory] Putorana Plateau. 10.07.1984, A. Zhukova; Chukotka Autonomous Area, Anadyr District. 07.08.1981, O. Afonina.
Calypogeia neesiana (4 specimens identified as Kantia trichomanis or Calypogeia trichomanis, all in LE). Latvia. Livland, Kreis Riga. 21.06 (= 04.07) 1909, J. Mikotowicz, 2f; Kurland, Kreis Doblen. 25.06 (= 08.07) 1906, J Mikotowicz,
Bryotheca Baltica Nr. 2; Livland, Kreis Riga... 18.05 (= 31.05) 1909, J. Mikotowicz, Bryotheca Baltica Nr. 2b; Livland, Kreis Riga. 21.06 (= 04.07) 1909, J. Mikotowicz, Bryotheca Baltica Nr. 2f (with admixture C. fissa mod. subintegrifolia).
Calypogeia suecica (3 specimens identified as Kantia trichomanis, all in LE). Russia. Lomonosov District, Bolshaya Izhora settlement, on rotten stump, 01.07.1984, A. D. Potemkin (Calypogeia cf. suecica). Estonia. Livland, Kreis Per-nau... 08.08 (= 21.08) 1909, J. Mikotowicz, Bryotheca Baltica Nr. 2e (Calypogeia cf. suecica); Livland, Kreis Pernau... 08.08 (= 21.08) 1909, J. Mikotowicz, Bryotheca Baltica Nr. 2e.
Results and Discussion
The revision of all kept in LE collections of C. trichomanis from Russia, Latvia, Georgia and Finland has shown their different identifications. They represent mostly materials of C. integristipula, partly of C. neesiana and exceptionally of C. suecica and C. fissa. All kept in LE older collections of C. trichomanis from Caucasus are C. fissa, from Arctic Siberia and Arctic Far East C. muelleriana and none of them can be certainly attributed to C. azurea (see lists of examined specimens above). Despite there are no reliable records of C. azurea from Finland based on alive collections until present (S. Laaka-Lindberg, pers. com. 27.01.2017) there are two old collections in LE, which correspond in morphology to this species.
C. azurea was collected alive for the first time for the Leningrad Region by Ginzburg in the Vsevolozhsk District and by Borovichev in the Republic of Karelia from Kivach State Nature Reserve. Earlier in Karelia it was collected alive by Bakalin (1999: 22) in the outskirts of Petrozavodsk City. In the field, the species was recognized based on greyish green or bluish color and deep blue oil bodies discernible with high magnification lens x 20 as blue or dark minute spots.
Morphological description (based on collections from the Leningrad Region and the Republic of Karelia)
Calypogeia azurea Stotler et Crotz, 1983, Taxon 32(1): 74. — Plates I, II.
Plants greyish green or bluish when alive, yellowish green when dried, from 1.6-1.8(2) mm wide x 7-16 mm long to 2-2.5 mm wide x 7-20 mm long, simple or with solitary Frullania-type and minute ventral intercalary branches. Leaves imbricate, ovate to cordate with maximal width near the base, ca. x 0.75-1.0 as wide as long, subacute to obtuse, exceptionally a few leaves emarginated at the apex. Underleaves x 1.2-2 as wide as stem; 2/5-3/5 bilobed, with (2)3-6 tires of cells from the base of sinus to the rhizoid-initial zone; lobes acute or subacute; lateral margins en-
trictiammis 16. MNIUM foliis diftichis integerrimis.
Milium trichorpanis facie, foliôiis integris. Dill. m-j. V
236. /,31. f. s-Habitat in Sueciîe, Anglîae udis.
Fig. Description of Mnium trichomanis by Linné (1753: 1114) and its corresponding illustration from Dillenius (1741: pl. XXXI).
tire or with solitary angulations; cells of underleaves subisodiametric or partly elongated; rhizoid-initial zone oval ca. x 3 as wide as high. Leaf cells almost not differentiated near the margins and at the base; thin-walled, without distinct trigones, apical cells subisodiametric, ca. 25-40 x 30-45 pm; marginal cells 23-42 x 30-45(55) pm; median cells subisodiametric, (25)30-40 x 28-42 pm. Oil bodies blue or deep blue, (1)2-5 per cells, ca. 2-7 x 5-13 ^m, of (1)2-7 large segments, 2-5 pm each, in fresh conditions discernible with hand lens as dark or blue minute spots, present in all cells of underleaves and most cells of leaves (Plate I, 4, 5) [according to Buczkowska (2004: 140) and Buczkowska et al. (2004: Fig. 2g) oil bodies are absent in group of leaf cells and present in all cells of underleaves]. Gemmae not recorded in our collections, rare in Nordic countries (Damsholt, 2002), fairly frequent in Great Britain — 22-30 x 25-46 pm, with pale blue oil bodies (Paton, 1999). Studied plants sterile. According to Damsholt (2002) autoicous or paroicous, according Paton (1999) autoicous.
Ecology. The gatherings of C. azurea in the Leningrad Region and the Republic of Karelia have been made from forest floor with humus reach or peaty soil in spruce forests in the territories adjacent to rivers or lakes having ± humid microclimate.
Most broadly ecology of the species is described by Paton (1999) for Great Britain, where C. azurea occurs on moist or wet, acid to mildly base-rich loamy to coarse sandy soil, on peat, Sphagnum and other bryo-phytes, Molinia tussocks, wet stones and rocks, on thin peat on rock including limestone, and rarely on rotten logs.
Distribution in Russia. Northwestern European Russia: Rebublic of Karelia, Leningrad Region; Caucasus: Kabardino-Balkarian Republic (Potemkin, Zhashuev, 2010), Karachayevo-Circassian Republic (Ignatova et al., 2008), Krasnodar Territory (Konstantinova, Akatova et al., 2009); South Urals Perm Region (Konstantinova, Bezgodov, 2009); South Siberia: Kuznetskiy Alatau, Kemero Region (Konstantinova et al., 2003); Far East, Kuril Islands, Iturup Island, Sakhalin Region (Koroteeva, 2011, pers. com. 09.02.2017). The species should occur also in Primorye Territory. However, in report of Bakalin (2008) all cited specimens were collected on rotten wood, which is not characteristic for this species. Recent records from Sakhalin (Bakalin et al., 2012) and Kamchatka (Bakalin, 2009) are based on materials without oil bodies.
Variation
Despite only three identified alive specimens were studied they demonstrate the broad range of variability of the species comparable with the range described and illustrated by Paton (1999), Damsholt (2002), Buczkowska (2004) and Buczkowska et al. (2004).
Size of plants. Plants from the Leningrad Region are larger, ca. 2-2.5 mm wide, than plants from the Republic Karelia, which are 1.5-2 mm wide.
Underleaves. Plants from the Leningrad Region have mostly broader and less deep divided underleaves with lateral angulations of margins, ca. 875-1050 ^m wide. Such large underleaves usually have rather narrow and sharp at the base y-like sinus and consist chiefly of subisodiametric cells. Smaller plants from the Leningrad Region have narrower underleaves, ca. 625 ^m wide, with broader U-like sinus. Smaller underleaves have often some elongated cells at the base and resemble underleaves of Karelian plants, which have even broader and often deeper sinuses and blunt lobes. The number of cell tires from the base of sinus to the rhizoid initial zone of underleaves varies greatly from usually 4-5(6) in larger plants to 3(2) in the smaller plants.
Leaves. Leaves of larger plants usually ± cordate, of smaller plants ± ovate, with maximal width near the base and chiefly subacute apexes. Rarely a few leaves with emarginate apex may be found.
Cells. Cells of leaves and underleaves are ± subisodiametric in larger plants, but often elongated in smaller plants.
Differentiation
Calypogeia azurea is a liverwort species with distinctive deep blue oil bodies. When oil bodies absent it may be confused with C. muelleriana, C. suecica, C. fissa or juvenile C. neesiana and C. integristipula with bilobed underleaves. C. azurea shares habitats with C. muelleriana, C. neesiana and C. integristipula. It differs from C. muelleriana in usually 2/5-3/5 bilobed underleaves with mostly subacute lobes; from C. fissa in often less deeply bilobed underleaves with (2)3-6 (vs. 2-4) tires of cells between base of sinus and rhizoid-initial zone, without regular marginal angulations or teeth and in chiefly entire or rare notched vs. often notched lobe apexes; from juvenile C. neesiana in marginal cells not forming a border; from juvenile C. integristipula in mostly ± cordate to ovate leaves about as wide as long vs. narrowly ovate leaves longer than wide.
The authors Potemkin and Borovichev made a lot of the liverwort flora research in the Northwestern European Russia since the 1984 and 2004 respectively. They paid special attention to the Calypogeia records but made no finds of C. azurea in the study area. This fact points at the pronounced rarity of this species in the Northwestern European Russia as well as the probably insufficient attention to its characteristic habitats. All authors' collections were made in the ground layer of spruce forests with high relative air humidity due to their location near lakes, rivers, or waterfalls. Special attention should be paid to such types of habitats and their conservation in future.
Acknowledgements
The authors are grateful to Nadezhda A. Konstantinova and Tatjana I. Koroteeva for their unpublished data on finds of Calypogeia azurea with oil bodies in Russia. Elmira G. Ginzburg is thankful to Gregory A. Isachenko for organization of the field trip to the place of collecting of C. azurea in the Leningrad Region. The two anonymous reviewers are thanked for their valuable comments and advices. Yuri B. Okolodkov (Universidad Veracruzana, Veracruz, Mexico) and Marcia M. Gowing (University of California at Santa Cruz, California, USA) kindly improved the style and corrected the English. The work of A. D. Potemkin and E. G. Ginzburg was carried out within the framework of the Koma-rov Botanical Institute research projects 012012255616 and «Herbarium collections of BIN RAS (history, conservation, investigation and replenishment». The work of E. A. Borovichev was partially supported by the Russian Science Foundation (project no. 15-14-10023) and the Russian Foundation of Basic Research (grant no. 15-29-02662).
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Литература
[Bakalin] Бакалин В. А. 1999. Печеночники Карелии. Arctoa. 8: 17—26.
Bakalin V. A. 2008. New liverwort records from Primorsky Territory. 2. Arctoa. 17: 221.
[Bakain] Бакалин В.А. 2009. Флора и фитогеография печеночников Камчатки и прилежащих островов. M.: 367 с.
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Plate I. Calypogeia azurea Stotler et Crotz from the Leningrad Region. 1 — part of shoot; 2-3 — leaves; 4 — underleaves with oil-bodies; 5 — median leaf cells with oil-bodies. Scale bars: 1-3 — 1 mm; 4 — 500 pm; 5 — 30 pm. All from Leningrad Region, Ginzburg 1109-3 (LE).
Plate II. Calypogeia azurea Stotler et Crotz from the Republic of Karelia. 1 — leaf; 2 — apex leaf cells; 3-5 — underleaves. Scale bars: 1 — 1 mm; 2 — 50 jm; 3-5 — 500 jim. All from Karelia Republic, Borovichev BE19-45-15 (INEP).
