Pylaisia condensata (Mitt.) A.Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1876

Ignatova, E. A., Kuznetsova, O. I., Shafigullina, N. R., Fedosov, V. E. & Ignatov, M. S., 2020, The genus Pylaisia (Pylaisiaceae, Bryophyta) in Russia, Arctoa 29 (1), pp. 135-178 : 163-165

7. Pylaisia condensata (Mitt.) A.Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1876 –77: 306. 1878 (Gen. Sp. Musc. 2: 372). — Stereodon condensatus Mitt., J. Proc. Linn. Soc. Bot., Suppl. 1: 93. 1859.

Figs. 21–23 ; 24A,B ; 25 :1–16; 34I.

Notes on taxonomy: This species for a long time has been called Pylaisia schimperi Cardot (described from Canada), until Crum et al. (1964) found its name to be illegetimate and thus requiring substitution with Pylaisia selwynii Kindb. (described from Ottawa, Canada), which Crum et al. (1964) placed in the genus Pylaisiella . Recently Arikawa (2017) found that Stereodon condensatus Mitt. described from West Himalayas (Kumaon) belongs to the same species as Pylaisia selwynii , thus he suggested the substitution of the latter with Pylaisia condensata . The present study found a rather considerable difference between the western North American and Asian plants, thus we suggest to retain the name Pylaisia selwynii for the former. The distribution of P. selwynii in North America needs further studies: at least one specimen from Alaska called P. selwynii in Genbank seems to belong to P. obtusa (a species superficially rather similar to P. selwynii ).

Morphological distinctions between the eastern North American plants referred here to ‘true P. selwynii ’ and P. condensata (=‘Eurasian P. selwynii ’) includes a smaller spores of the former species, 16–24 µm vs. 18–30 µm, with at least some spores ca. 30 µm found in most collections. Also the number of dorsal trabeculae on the exostome teeth was found fewer in North American P. selwynii than in Eurasian plants ( Fig. 24 ), although the latter character was studied by us on a limited material only.

Description. Plants small to medium-sized. Stems and branches curved, densely foliate; leaves homomallous, turned outwards substrate. Stem leaves broadly ovate to ovate-lanceolate, rather suddenly narrowed into a rather short acumen, 0.8–1.2(–1.4)× 0.5–0.6 mm; margins subentire; median laminal cells 25–40(–50)×6–9 µm, alar cells subquadrate to rectangular, homogeneous, in a rather large group, 20–25 cells long and 6–7(–10) cells wide. Branch leaves somewhat smaller. Inner perichaetial leaves erect or somewhat reflexed, oblong-lanceolate, to 1.6 mm long. Capsules ovoid. Peristome in dry state forms a low conus due to adherence of its elements. Exostome teeth 150–200 µm long, dorsal plates smooth below, coarsely papillose above; endostome adherent to exostome teeth to about teeth halves; segments narrow, split except their uppermost part; endostome moderately papillose. Spores 16–30 µm.

Differentiation and variation. Despite P. condensata is a variable taxon, in most of its range, where it cooccurs with P. polyantha , this species is easy to recognize in the field by strongly homomallous to falcate leaves, curved branches and ovate capsules that are essentially cylindrical in P. polyantha . In southern Russian Far East, the region where other East Asian species of the genus

21 22

occur, variation of P. condensata is greater and its delimitation is problematic. Most difficult is its separation from P. brotheri . These difficulties were discussed by Noguchi (1994), who mentioned among the distinctive characters of P. brotheri broader leaves, rather triangular than elongate alar regions, and broader capsules. Some of these character states, however, were found in three specimens that belong to the same clade as P. condensata . These specimens are marked in the haplotype network as CoB1, 2, 3 ( Fig. 4 ). They are not grouped to-

7 9

1510 11 12

Fig. 22. Pylaisia condensata (all specimens from Russia); 1–3: Co6, OK2754, Amurskaya Province, Norsky Reserve; 4–6: Co4, OK221, Khabarovsk Territory, Bureinsky Reserve ; 7–9: Co1, OK2744, Sakhalin; 10–12: Co3, OK448, Altai Mts; 13–15: Co5, OK2752, Primorsky Territory. 1–2, 4–5, 7–8, 10–11, 13–14: stem leaves; 3, 6, 9, 12, 15: branch leaves. Scale bar: 1 mm for all .

gether, thus we consider them as an infraspecific variation of P. condensata . Noteworthy, such morphotypes are especially similar in leaf shape to the type of P. condensata illustrated by Arikawa (2017).

Despite of certain doubts of Noguchi (1994) that P. condensata (referred that time to P. selwynii ) and P. brotheri are good species, Arikawa & Higuchi (2003) found that rbc L sequences of these species are distinct enough to maintain their status of distinct species.

Distribution: Pylaisia condensata is a widespread Eurasian species known from Japan, China, and India; in Russia it is common in the Far East, northward to Kamchatka, and is sporadically distributed in southern Siberia, while in European Russia it is resticted to northern regions, westward to the border with Finland.

Selected specimens examined: EUROPEAN RUSSIA: Republic of Karelia: Kondopoga Distr. , 8-10 km W of Kondopoga, 28 Aug.1990 Smirnova s.n. (MHA9047582) . Leningrad Province: Podoporozhsk Distr., Terebsky Brook , 20 Aug. 2008 Kushnevskaya s.n. (MHA) . Vologda Province: Gryazovetsky Distr., 19 Aug. 2001 Ignatov & Ignatova 38 (MHA 9047555); Nikolsk Distr., Yug River ner Permas Village , 11 Aug. 2001 Ignatov & Ignatova 28 (MHA9047558); Nyuksenitsy Dustr., Sukhona River opposite Nyuksenitsy Settl, 13 Aug. 2001 Ignatov & Ignatova 31 (MHA9047559); Sokol Distr. , Shorega Creek , 22 Sept. 1990 Ignatov s.n. (MHA9047568) . Kostroma Province: Manturovo Distr., E of Unza River by the Hwy Kostroma-Verchnespasskoye, 18 Sept. 1990 Ignatov s.n. (MHA9047567); Sudislavl Distr. , 30 km N of Sudislavl, 19 Sept. 1990 Ignatov s.n. (MHA9047570) . Komi Republic: Syktyvkar, 1977 Zheleznova s.n. (MHA9047576) . Perm Territory: Gornozavodsk Distr., Basegi Reserve , Vilva Rive near Koroostelevka Creek mouth, 08 Jun. 1994 Ignatov & Bezgodov 215 (MHA9047577); Kosinsky Distr. , Gorki Village , 04 Aug. 2012 Bezgodov 252 (MHA9047580); Yurlinsky Distr. , 30 km W of Yurla, 2 Aug. 2012 Bezgodov 213 (MHA9047581) . Sverdlovsk Province: Alapaevsk , 31 Aug. 1928 Igoshina 38/2 (MHA9047634) . Bashkortostan Republic: Beloretzk Distr., Maly Iremel Mt., 31 Aug. 1990 Ignatova 2/1 (MHA9047579); Zilmerdak Range, Zuyakovo Settl., 1 May 1990 Muldashev 64 (MHA9047578) . Ivanovo Province: Provolzhsky Distr., Volga – Uvod’ channel, 21 Apr. 2014 Sorokin & Golubeva 1336 (MHA9047563). Marij-El Republic: Orshansk Distr. , Hwy from Yoshkar-Ola to Yaransk, near the border of Kirovsk, 16 Sept. 1990 Ignatov s.n. (MHA9047566) . Nizhny Novgorod Province: Bor Distr., Kerzhensky Reserve , 12 Sept. 1999 Ignatov s.n. (MHA9047554) . Tver Province: Nelidovo Distr., Central Forest State Reserve, Stulovo , 5 Aug. 1994 Ignatov s.n. (MHA9047574) . Yaroslavl Provonce: Danilov Distr. , 2 km W of Danilov Town, 21 Sept. 1990 Ignatov s.n. (MHA9047565) .

ASIAN RUSSIA: Khanty-Mansi Autonomous District: Surgut Distr., Yugansky State Reserve , 20 Jul. 1987 Onipchenko s.n. (MHA9047636). Tyumen Province: Uvatsky Distr. , Moil River , 14 Sept. 2013 Bezgodov 203 (MW). Novosibirsk Province: Salairsky Mts. , Korotovo Settl., 26 Jun. 1993 Pisarenko op00912 (MHA9047591). Altai Republic: Teletzkoe Lake , Yailyu, 06 Aug. 1988 Zolotukhin s.n. (MHA9047626); Ulagan Distr. , 6 km upstream from Chulyshman River mouth, 17 Aug. 2012 Ignatov & Ignatova 12-528 (MHA9047623); Chemal Distr. , Chemal River , 12 Jul. 1993 Ignatov 34/135 (MHA); Amurskaya Province: Zeysy Reserve , Teplyj Klyuch, 22 Sept. 1979 Petelin 289 (MHA9047637). Krasnoyarsk Territory: Evenkia, Baikit Distr. , Central-Siberian Reserve , 19 Jul. 1992 Szerbina 141 (MW); Taimyr Distr. , Khatanga settl. outskitrs, vicinity of Kotuykan River mouth, 9 Aug. 2011 Fedosov 11-390 (MW). Republic of Sakha /Yakutia: Khangalassky Distr. , National Park “Lena Pillars”, 05 Aug. 2016 Ignatov & Ignatova 16-434 (MHA9021361); Tomponsky Distr. , Sette-Daban, Okraina Ridge, between Ulakh and Nadezhda Creeks, 17 Jul. 2016 Ignatov & Ignatova 16-957 (MHA9022653). Irkutsk Province: East Sayan Mts , Udinsky Range, Kara-Buren’ River , 13.VIII.1961 Bardunov s.n. (MHA9047632); Slyudyanka Distr. , Slyudyanka Creek , 08 Jun. 2005 Ignatov & Kazanovskii s.n. (MHA 9047604). Zabaikalsky Territory: ca. 40 km W of Chita Town, 4 Aug. 1988 Bochkin et al. s.n. (MHA9047633). Amurskaya Province: Zeisky Reserve , Tukuringra Range, 21 Sept. 2012 Dudov Br_12_059 (MW9061951). Khabarovsk Territory: Sarapulskoe Settl., 22 Aug.1988 Bochkin et al. s.n. (MHA 9047644); Verkhnebureinsky Distr. , upper course of Bureya River , 04 Aug. 1997 Ignatov 97-195 (MHA9047607); Bureinsky Reserve , Levaya Bureya River 2 rm upstream Chapkhoz Creek mouth, 16 Aug. 1989 Grigor’eva 89-М-90 (MHA9047642); Sovgavan Distr. , Botchi Reserve , Mulpa River , Ignatov & Ignatova 13-239 (MHA9047583); Badzhal Mts. , Yarap River valley , 14 Aug. 2016 Fedosov 16-48 (MW9130180). Primorsky Territory: Lazovsky Reserve , Perekatnaya River , 21 Sept. 1977 Bardunov s.n. (ex IRK, MHA); Vladivostok Area , Okeanskaya Station , 20 Aug. 1952 Voroshilov 6748 (MHA9047638); Dalnegorsk Distr. , 5 km N of Sedaya Mt., Ignatov & Ignatova 13-1496 (MHA9047617); Lazo Distr. , Elomovsky Klyuch Creek , Ignatov et al. Pr-5-13-13 (MHA9047594); Vadivostok, Bolshaya Sefdanka Creek , Ignatov et al. 06-2453 (MHA); Olga Distr. , waterfall on Milogradovka Creek , 21 Aug. 2007 Ignatov 07-434 (MHA9047600); Partizansk Distr. , Olkhovaya Mt. , 4 Oct. 2006 Ignatov et al. 06-2551 (MHA); Partizansk Distr. , Chandolaz Mt. , Ignatov et al. 06-2718 (MHA9047596); Ussurijsky Reserve , Komarovka Creek , Ignatov & Ignatova 06-3426 (MHA). Chuguevka Distr. , Izvilinka Creek ca. 5 km downstream Bereznyaki village, 15 Aug. 2007 Ignatov 07-502 (MHA9047602). Sakhalinskaya Province: Kuril Islands , Shikotan Island , Nyushko N-1-06a (MHA9047590); Sakhalin, Tymovsky Distr. , Tym’ River , Ignatov & Teleganova 06-424 (MHA9047589). Kamchatsky Territory: Kamchatka, Pravy Kikhchik River basin, 22 Jul. 2004 Czernyadjeva 12 (MHA9047603) .

Figures

Fig. 21. Pylaisia condensata (1–12: Co10, OK 446, Russia, Ivanovo Province) and P. selwynii (13, 15–23: Se2, OK548, U.S.A., Ochio; 14: OK2740, U.S.A., Missouri). 1–3, 15–17: branch leaves; 4, 13–14: capsules; 5, 12: habit, dry; 6, 21: median laminal cells; 7, 22: upper laminal cells; 8, 23: basal laminal cells; 9–11, 18–20: stem leaves. Scale bars: 5 mm for 5; 2 mm for 4, 12–14; 1 mm for 1–3, 9–11, 15–20; 100 µm for 6–8, 21–23.

Fig. 23. Pylaisia condensata, SEM images of peristomes, from European Russia, A–B from Kostroma; C–D from Co10, OK 446, Russia, Ivanovo Province.

Fig. 24. Peristomes of Pylaisia under light microscope, showing endostome split along the keel nearly to the top of segments and its extensive adherence to exostome teeth. A–B: Pylaisia condensata, Co10, European Russia, Ivanovo; and C: P. selwynii (USA, Missouri, Allen 13400, Mosses of the Interior Highlands Exsiccatae #325; LE).

Fig. 25. Pylaisia brotheri (1–15, B, OK2799, Russia, Primorsky Territory,Ussurijsky Reserve) and P. condensata (16–24, CoB2, OK2796, Rus- sia, Primorsky Territory, Chandolaz Mt.). 1–2, 16: capsules; 3–4: habit, dry; 5–8, 18–20: stem leaves; 9, 24: basal laminal cells; 10, 23: me- dian laminal cells; 11: upper laminal cells; 12– 14, 21–22: branch leaves; 15–16: perichaetial leaves. Scale bars: 5 mm for 4; 2 mm for 1–3, 17; 1 mm for 5–8, 12–16, 18–22; 100 µm for 9– 11, 23–24.

Fig. 4. TCS haplotype network based on ITS dataset (105 accessions) for Homomallium and Pylaisia. Missing haplotypes are shown as small black circles upon haplotype interlinks. Abbreviations and specimen details are in Table 1, the distribution of P. polyantha, P. coreana, P. steerei, and P. curviramea is shown in Fig. 5.