Saccobasis polita (Nees) H. Buch var. arctica Konstant., Vilnet & Mamontov, 2022

Konstantinova, N. D., Vilnet, A. A. & Mamontov, Yu. S., 2022, How many taxa are in the genus Aaccobasis h. Buch? Evidence from integrative taxonomy, Arctoa 31 (2), pp. 166-180 : 177-179

Saccobasis polita (Nees) H. Buch var. arctica Konstant., Vilnet & Mamontov , var. nov. ( Figs. 3 , 4 , 5 :A, D)

Diagnosis. Differs from Saccobasis polita in smaller size, color of shoots without purple secondary pigmentation, often concave leaves that are not deccurent, noticeable admixture of two- or four-lobed leaves, mostly point- ed leaf lobes, and angulate sinuses, shorter cells on dorsal side of shoot and at leaf base.

Type: Norway: Svalbard, Wedel Jarlsberg Land, Recherchefjorden, Lagerneset, 77°31’31.8’’ N – 14°45’54’’ E, river terrace, patterned ground tundra on gentle slope, on frost boils, on side of micro-bumps and in crevices, dominate in mats with admixture of Anthelia juratzkana (Limpr.) Trevis. , Blepharostoma brevirete (Bryhn et Kaal.) Vilnet et Bakalin , Cephaloziella varians (Gottsche) Steph. , Odontoschisma macounii (Austin) Underw. , 19.VIII.2018, Konstantinova & Savchenko # K 45-3a-18 (KPABG-123662, holotype), paratypes all specimens cit- ed below. GenBank accession numbers are OP584670 for ITS1-2 and OP573508 for trn L-F.

Etymology. The name refers to the restriction of the variety to the difficult high arctic and subarctic mountains habitats.

The two varieties can be distinguished as follows:

l. Plants brown to dark and even black-brown, the color of the base of leaves usually does not differ from color leaves, small, (0.4–)0.6–1.2(–2) mm wide, cells on dorsal side of stem 2, rarely 3 times as long as wide, leaf lobes mostly pointed ending by one or sometimes two superposed cells, both sinuses mostly angulate, sometimes sharp at base and to 0.25(–0.3), leaf length, shoots often with noticeable admixture of two- or four-lobed leaves. Occurs on bare soil in crevices on frost boils in patterned ground tundras or on rocks in brook beds — S. polita var. arctica

1. Plants green to warm brown with purplish red secondary pigmentation and more deeply brown or purplish red brown basal part of leaves, mostly medium sized, (1.2–) 1.5–3 mm wide, cells of dorsal side are (2–)4–6 (–8) times as long as wide, leave lobes obtuse or round- ed, one sinuses often semi crescentic or even shallow crescentic just 0.1–0.15 leaf length, while the second can be angulate sometimes sharp at base and to 0.25(–0.3) leaf length, leaves mostly 3-lobed but single 4- or 2-lobed leaves occur. Occurs in moist humus-rich habitats — S. polita var. polita

Ecology. The type variety is a hydrophilic basiphile occurring in moist habitats as a pioneer directly on damp, mainly basic, sediments and rocks or on humus-rich soil and peat covered basic rocks, e.g. on banks of rivulets or in beds of streams drying up, in Alnus fruticosa Rupr. tickets, in swampy and peaty areas with basic seepages both in ravines and on rock outcrops, at base of cliffs, in swampy depression with Ledum palustre L. and Alnus fruticosa Rupr. in forests near sea-side, in sedge-moss and sedge subshrubs tundra on edges of pools, in pools in hummocky boggy tundra. The species often occur in pure turfs or mats or mixed with other bryophytes. Most often its associates are bryophytes of rich or eutrophic mires or stream sides Aneura pinguis (L.) Dumort., Blepharostoma trichophyllum (L.) Dumort., Harpanthus flotovianus (Nees) Nees , Odontoschisma elongatum (Lindb.) A.Evans , Scapania uliginosa (Lindenb.) Dumort. , Trilophozia quinquedentata (Huds.) Bakalin , Campylium stellatum (Hedw.) C.E.O. Jensen , Loeskypnum badium (Hartm.) H.K.G. Paul , Meesia uliginosa Hedw. , Scorpidium revolvens (Sw. ex anon.) Rubers,, Warnstorfia sarmentosa (Wahlenb.) Hedenäs , and weak calciphytes like Mesoptychia gillmanii (Austin) L. Söderstr. & Váňa , M. heterocolpos (Thed. ex Hartm.) L. Söderstr. & Váňa. Saccobasis polita var. arctica is found in more xeric habitats. All specimens from Svalbard and Franz Josef Land were collected in crevices in patterned ground tundra on frost boils. Specimens from Trans-Baikal Territory were found on rocks in river valley in forest zone near timberline. Associates of Saccobasis polita var. arctica are arctic and montane species like Anthelia juratzkana (Limpr.) Trevis. , Blepharostoma brevirete (Bryhn & Kaal.) Vilnet & Bakalin , Jungermannia polaris Lindb. , Cephaloziella varians (Gottsche) Steph. , Odontoschisma macounii (Austin) Underw. , Marsupella arctica (Berggr.) Bryhn & Kaal. and Schljakovianthus quadrilobus (Lindb.) Konstant. & Vilnet.

Distribution. Saccobasis polita is an arctic-montane species ( Fig. 6 , 7). It is most widespread in the oceanic and suboceanic areas with predominance of basic rocks. In Europe it occurs scattered in mountains of central and western including north-western Europe and Iceland and the British Islands ( Paton, 1999; Damsholt, 2002) in the west to Svalbard in the north where the species is present as var. arctica . In the European part of Russia the species is not rare in the Murmansk Province, but much rarer both to the south where it is recorded only in the north of Republic of Karelia ( Bakalin, 1999) and to the east where it occurs in Nenets Autonomous District (Potemkin, 2008) and Urals (Zinovjeva, 1973). In Asia the species is not rare in mountains in oceanic and suboceanic areas of Far East of Russia, particularly in Chukotka Autonomous Areas and Kamchatka Territory ( Konstantinova et al., 2009; Bakalin, 2010) and occurs scattered in humid areas of mountains of South ( Váňa & Ignatov 1995; Konstantinova et al., 2003) and Baikal Siberia (authors’ unpublished data, see specimens examined), where it probably is not rare in appropriate localities. In the north of Asia the species is found in single localities on eastern slopes of Urals (Storozheva, 1986, Konstantinova & Lapshina, 2017), on Putorana Plateau ( Bakalin et al., 2016) and several localities in the Taimyr Peninsula (KPABG, isling.org/hepatics). The species is probably more widespread in poorly studied northern and mountain regions of Asia and will be found more to be common in future studies.

In the western hemisphere the species is rather common in West and South Greenland ( Damsholt, 2013), not rare in western and northern part of North America from Alaska and Yukon to British Columbia but rather rare in east America from Ellesmere I. to Newfoundland and Quebec ( Schuster, 1969).

Selected specimens examined: Saccobasis polita . RUSSIA, Republic of Karelia, 17.VIII.1989, Konstantinova #215-2-89 (KPABG-2911). Rybachiy Peninsula, 19.VII.1978, Konstantinova # 83-2-78 (KPABG-6138); Monche tundra, 28.VI.2009 Borovichev # BE37-21-09 (KPABG-19158); Paz River Valley, 02.VIII.1994, Kostina s.n. (KPABG-8622); Khibiny Mountains, 09.VIII.1998, Konstantinova #39-1-98 (KPABG-6937), ibid. 10.VIII.1996, # 21-3b-96 (KPABG-6189), ibid. 20.VIII.2000, #310-1-00 (KPABG-8167); Iokan’ga River Valley , 18.IX.1997, Konstantinova # 60-2-97 (KPABG-6203) . Khanty-Mansi Autonomous Area , 14.VII.2018, Lapshina s.n. (KPABG-122280), ibid. 10.VIII.2015, Lapshina #1771 (KPABG-121323) . Kemerovo Province, 28. VI.2000, Konstantinova #61-1-00 (KPABG-101870) . Krasnoyarsk Territory, Taimyr Peninsula 23.VII.2021 Lapshina # YSU-MH-04421 ( YSU, KPABG); Eastern Sayan , 5.VIII.1993, Vasiljev ( KPABG 103028 ) Eastern Sakhalin Province, 11.IX.2009, Bakalin # S-61-37 - 09 , Hep. Ross. Exs., Fasc. VI, No. 143 (KPABG-116071) ; Chukotka Autonomous Okrug ( Area ), 15.VII.1988, Kuzmina #D1(20) (KPABG-100003) . Kamchatka Territory , 01.VII.2004, Bakalin # K-24-16-04 (KPABG-106564) . AUSTRIA, as Tritomaria polita (Nees) Jørg. , 23.VIII.1882, Breidler, Hep. exic. S. O. Lindberg, Fasc. II, No. 462 (KPABG-s.n.) . USA, Alaska, Matanuska-Susitna Borough , all as Saccobasis polymorpha : 04.VII.1992, Konstantinova #88-2-92 (KPABG-123744), ibid. 04.VII.1992, Konstantinova # 94-1-92 [KPABG-123771], ibid. 07.VII.1992, Konstantinova, # 120-9-92 [KPABG-123867] .

Saccobasis polita (Nees) H.Buch var. arctica Konstant., Vilnet, Mamontov (paratypes): RUSSIA, Franz Josef Land , Ziegler (Tsigler) Island, 25.VII.2019, Savchenko # CA19-30 - 5 b (KPABG-122768). Trans Baikal Territory, Mamontov # 525- 1-2 (KPABG-119591), ibid. # 526-2-2 (KPABG, MHA). NORWAY: Svalbard, Sørkapp Land, Stormbukta, 15.VIII.2018, Konstantinova & Savchenko # K11-1 a-18 (KPABG-123564); Prins Karls Forland, 06.VIII.2016, Savchenko # CA16-34 - 2 a (KPABG-121469); Wedel Jarlsberg Land, Recherchefjorden, 19.VIII.2018, Konstantinova & Savchenko # K45-2 b-18 (KPABG-123661), ibid. 19.VIII.2018, # K45-1 c-18 (KPABG-123658); Haakon VII Land, 07.VIII.1974, Frisvoll # TRH74005 (KPABG-122658, dupla).

Figures

Fig. 3. Saccobasis polita var. arctica: A-D – habit. E-P – leaves. A-G, I-K, M, P from Mamontov 526-2-2 (KPABG, MHA). H from Savchenko CA-14-8a (KPABG, MHA). L, N, O from Savchenko CA-16-34-2a (KPABG, MHA).

Fig. 4. Saccobasis polita var. arctica: A-E – habit. F – part of shoot showing leaf insertion, dorsal view. G, H – leaf lobe apices showing gemma formation and mature gemmae. I, J – leaves. K, L – stem cross sections. A, C-E, L from Savchenko CA-16-34-2a (KPABG, MHA). B, H-K from Konstantinova K45-3a-18 (KPABG, MHA). F from Mamontov 526-2-2 (KPABG, MHA). G from Savchenko CA-14-8a (KPABG, MHA).

Fig. 5. Saccobasis polita var. arctica: A, D – basal leaf cells. Saccobasis polita: B, C, E – basal leaf cells. A, D from Mamontov 526-2-2 (KPABG, MHA). B, C, E from Bakalin 01.VII.2001, Hep. Ross. Exs., No. 19 (MHA).

Fig. 6. Distribution map of Saccobasis taxa. A: S. polita s.l. based on sequenced and studied specimens. Black circles – sequenced specimens of Saccobasis polita var. arctica, unpainted circles – studied specimens of S. polita var. arctica, black squares – sequenced specimens of S. polita var. polita, unpainted squares – studied specimens of Saccobasis polita var. polita. B: distribution map of Saccobasis polita s.l. based on all available data. Black circles: S. polita var. arctica; squares S. polita var. polita.