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In Middle European Russia Род Bryum (Bryaceae, Musci)

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In Middle European Russia Род Bryum (Bryaceae, Musci) Arctoa (2000) 9: 155-232 THE GENUS BRYUM (BRYACEAE, MUSCI) IN MIDDLE EUROPEAN RUSSIA ÐÎÄ BRYUM (BRYACEAE, MUSCI)  ÑÐÅÄÍÅÉ ×ÀÑÒÈ ÅÂÐÎÏÅÉÑÊÎÉ ÐÎÑÑÈÈ VALERIJ IVANOVICH ZOLOTOV1 ÂÀËÅÐÈÉ ÈÂÀÍÎÂÈ× ÇÎËÎÒÎÂ1 Abstract A revision of the genus Bryum in Middle European Russia (= European Russia, exclud- ing the Caucasus, the Kola Peninsula and Karelia, and northern Arkhangelsk Province and the Komi Republic, and including the Ural Mountains to the northern border of Perm and Sverdlovsk/Ekaterinburg Provinces) documents the occurrence of twenty-nine species, in- cluding B. algovicum, B. amblyodon, B. alpinum, B. arcticum, B. argenteum, B. bicolor, B. bimum, B. caespiticium, B. capillare, B. creberrimum, B. cyclophyllum, B. elegans, B. funkii, B. intermedium, B. klingraeffii, B. knowltonii, B. kunzei, B. laevifilum, B. lonchocaulon, B. pallens, B. pallescens, B. pseudotriquetrum, B. rubens, B. schleicheri, B. subapiculatum, B. turbinatum, B. uliginosum, B. violaceum, B. weigelii. All species are described and illus- trated, selected specimens are cited (at least one for each province/republic), and a key is provided. Bryum bimum is treated as distinct from B. pseudotriquetrum, as is B. kunzei from B. caespiticium, and B. lonchocaulon (B. cirrhatum auct.) from B. pallescens. Ðåçþìå Ïðîâåäåíà ðåâèçèÿ ãåðáàðèíûõ ìàòåðèàëîâ ïî ðîäó Bryum ñ òåððèòîðèè ñðåäíåé ÷àñòè Åâðîïåéñêîé Ðîññèè (=Åâðîïåéñêîé ÷àñòè áåç Ñåâåðíîãî Êàâêàçà, Êîëüñêîãî ïîëóîñòðîâà è Êàðåëèè, ñåâåðà Àðõàíãåëüñêîé îáëàñòè è Ðåñïóáëèêè Êîìè). Ïîäòâåðæäåíî ïðèñóòñòâèå çäåñü 29 âèäîâ: B. algovicum, B. amblyodon, B. alpinum, B. arcticum, B. argenteum, B. bicolor, B. bimum, B. caespiticium, B. capillare, B. creberrimum, B. cyclophyllum, B. elegans, B. funkii, B. intermedium, B. klingraeffii, B. knowltonii, B. kunzei, B. laevifilum, B. lonchocaulon, B. pallens, B. pallescens, B. pseudotriquetrum, B. rubens, B. schleicheri, B. subapiculatum, B. turbinatum, B. uliginosum, B. violaceum, B. weigelii. Ïðèâîäèòñÿ êëþ÷ äëÿ îïðåäåëåíèÿ âèäîâ, è äëÿ êàæäîãî èç íèõ äàíû îïèñàíèÿ, èëëþñòðàöèè, ïåðå÷èñëåíû èçó÷åííûå îáðàçöû (ïî êðàéíåé ìåðå îäèí äëÿ êàæäîé èç îáëàñòåé/ðåñïóáëèê). Ïðèâîäÿòñÿ äàííûå â ïîëüçó âèäîâîé ñàìîñòîÿòåëüíîñòè B. bimum è B. kunzei, êîòîðûå â ðÿäå ïîñëåäíèõ ïóáëèêàöèé îòíîñèëè â ñèíîíèìû ê B. pseudotriquetrum è B. caespiticium ñîîòâåòñòâåííî. Òàêæå, B. lonchocaulon (B. cirrhatum auct.) è B. pallescens ïðèíèìàþòñÿ êàê ñàìîñòîÿòåëüíûå âèäû. INTRODUCTION mine how many species occur in this region and to The genus Bryum has a reputation as one of the identify their distinguishing characteristics. Spe- systematically most difficult genera of mosses. It is cies concepts are based on the publications of Lim- challenging to attempt identification of vast num- pricht (1895), Podpera (1959, 1962), Savicz-Lju- bers of specimens because of considerable intraspe- bitskaya (1954), Savicz-Ljubitskaya & Smirnova cific polymorphism; lack of, or imperfect preserva- (1970), Melnichuk, (1970), Nyholm (1993), Syed tion of, sporophytes; and insufficient material for (1973), Crundwell & Nyholm (1964), Demaret certainty regarding sexual condition. Also, the ge- (1986a, b), Smith (1978) and Smith & Whitehouse nus is one of the largest in the boreal and temper- (1978), among others, as well as on collections iden- ate bryofloras. These difficulties generally result in tified by Podpera, Warnstorf and other authori- a number of erroneous identifications. The main ties on European bryophytes (specimens in MW, goal of the present study is to revise the collections LE). from Middle European Russia in order to deter- MATERIAL AND METHODS 1 – Main Botanical Garden of Russian Academy of Sciences, Botanicheskaya 4, Moscow 127276 Russia – Ðîññèÿ 127276 Ìîñêâà, Áîòàíè÷åñêàÿ, 4, Ãëàâíûé áîòàíè÷åñêèé ñàä ÐÀÍ 156 V. I. ZOLOTOV The study area includes Middle European Rus- zling case is the group, B. pallescens–B. lon- sia (i.e., European Russia, excluding the Caucasus chocaulon–B. creberrimum; it is discussed in and northern territories: the Kola Peninsula, Kare- detail under the first species. lia, Northern half of Leningrad/St.-Petersburg Male and female plants of dioicous species are Province, and northern Arkhangelsk Province and often in separate tufts, but in some species both the Komi Republic, and including the Ural Moun- sexes can be often found in the same tuft (for tains up to the northern border of Perm and example, in B. pseudotriquetrum), at least if the Sverdlovsk/Ekaterinburg Provinces). collection is large. Male plants are usually slen- All Bryum collections from the study area in der, more loosely foliate (B. pallens, B. caespitici- MW, MHA, LE and VOR have been examined, as um), or almost identical with female plants (B. well as some from H and local Russian herbaria, schleicheri). Perigonia are usually about as long including Tver State University, Ekaterinburg Ped- as wide, and have a + capitate shape, but in some agogical University, Institute of Ecology (Ekat- species (B. schleicheri, B. pseudotriquetum) with erinburg), and Volgograd Pedagogical Universi- especially numerous antheridia, perigonia are wide, ty. Altogether, approximately 700 specimens were i.e., noticeably wider than long. studied. Several species cited in publications per- Plants grow as separate individuals scattered taining to Middle European Russia were not avail- on the substrate (often in B. funkii, sometimes able for study, and these are cited in the section in B. argenteum), or form loose (B. weigelii), “unconfirmed records”. compact (B. bimum), or dense tufts (B. caespit- Measurements in species descriptions are based icium). I call a tuft ‘compact’ if a complete shoot on ten specimens from different parts of the terri- can be pulled off the tuft from above, whether tory, if this number was available. The number of dry or wet. Correspondingly, I call a tuft ‘dense’ specimens studied is indicated in parentheses af- if a shoot usually breaks in attempts to pull it ter “specimens studied”. Species known only from from above, and the separation of a single shoot sterile plants are supplemented with descriptions requires first a splitting of a tuft and then the of sporophytes (and sometimes perigonia and careful removal of a plant from the opening. The perichaetia) based on other, mostly European col- compact or dense states are generally dependent lections, which are listed in Appendix 1. All data on the density of rhizoidal tomentum, but they from outside of our area are in square brackets. are not correlated in all cases. Therefore, tuft The SEM analyses of peristome and spores in- density can be treated as a separate character. clude several species from outside of the study The height of tufts is a complex parameter re- area, and a number of these are used to illustrate sulting from the length of female shoots, the length particular character states. of subterminal shoots, and the mode of growth ANALYSES OF CHARACTER STATES (shoots are mostly straight, or sometimes ascen- Sexual condition traditionally has been con- dant, as in B. rubens, B. laevifilum, or shady phe- sidered one of the important characters in Bryum notypes of B. capillare, and includes both living taxonomy, because most species are invariable and dead, partly decayed parts). Tufts vary from in this respect. In some groups, the differences low (less than 0.5 mm, as in B. violaceum) to tall in sexual condition were treated as an impor- (10-12 cm, as in B. pseudotriquetrum, B. schle- tant basis for species segregation, e.g., with icheri, B. weigelii). Bryum pseudotriquetrum–B. bimum, B. palle- Tuft color can be pure green (B. capillare), scens–B. lonchocaulon–B.creberrimum, and B. light green (B. rubens), dirty green (often in B. capillare–B. torquescens. However, recent au- turbinatum), pale green (often in B. weigelii), yel- thors have expressed doubts about the signifi- low-green (B.cirrhatum), brownish-green (some- cance of this character, which (along with oth- times in B. bimum), brownish-yellow (sometimes er evidence) has led some to reduce the status in B. elegans), pinkish to vine-red (often in B. of B. bimum to a variety or a synonym with B. pallens), glaucous (sometimes in B. laevifilum and pseudotriquetrum, and to treat B. lonchocau- B. turbinatum), whitish (B. funkii), silvery (B. lon (as B. cirrhatum) as synonymous with B. argenteum), or variegated with a mixture of green, pallescens. Sexual condition was found, in the yellow-brownish and purplish (B. alpinum). Tufts present study, to be fairly stable. The most puz- can be dull (B. weigelii) or glossy (B. alpinum). The genus Bryum (Bryaceae, Musci) in Middle European Russia 157 Most Bryum species have leaves + evenly ar- tively rare character state, known in certain spe- ranged along the stem, except the leaves around cies groups not represented in our study area gametangia are larger and more densely crowded, (e.g., B. calophyllum), and occasionally in spe- or comose (from the Latin adjective comosus, which cies with normally holodontous peristomes (e.g., is used sometimes as a species epithet in vascular B. creberrimum). In most species, the teeth are plants to describe characteristically crowded up- gradually tapered in the transition zone, rarely per flowers). The largest comose leaves (Fig. 69, p. more abruply tapered (B. bicolor, Fig. 7), and 168) are most characteristic for individual spe- pale-yellow to brownish, reddish and orangish. cies, and they are used in the descriptions and An important diagnostic character of tooth measurements of “upper leaves”. Leaves from the is the number of ventral trabeculae: they were lower part of stems are described additionally count up to the upper well-developed one (i. e. for species where they are numerous. Inner comose uppermost non-projecting PPL transversal cells leaves become smaller and narrower as the mar- walls were not included). gins are more recurved, and often form a gradual The part of an exostome tooth below the lower- transition to outer perichaetial leaves. most ventral trabeculae is referred to as the ‘fun- The thickness and porosity of laminal cell dus’, and usually is considered important in the walls were evaluated in median cells between taxonomy of Bryum. The fundus can be broadened, the costa and leaf border.
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