Áðèîôëîðà Ñðåäíåðóññêîé Âîçâûøåííîñòè. I Bryoflora of the Central Russian Uppland
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Arctoa (2002) 11: 101-168 ÁÐÈÎÔËÎÐÀ ÑÐÅÄÍÅÐÓÑÑÊÎÉ ÂÎÇÂÛØÅÍÍÎÑÒÈ. I BRYOFLORA OF THE CENTRAL RUSSIAN UPPLAND. I ÍÀÒÀËÜß ÍÈÊÎËÀÅÂÍÀ ÏÎÏÎÂÀ1 NATALIA N. POPOVA1 Abstract The list of bryophytes of Central Russian Uppland, a separate physiogeographic land of the of the East Europe, is given. Brief essay of natural conditions and history of bryological investigation is provided. The major part of the Central Russian Uppland is situated in the forest-steppe zone, while its northern part is in area of deciduous forests and the southern one is in the steppe zone. Specific peculiarity of the region is the abundance of chalk and limestone outcrops on slopes to river valleys and ravines. The conspect of bryoflora includes 370 taxa; collections of author (of 1981-2002) and revised by author are marked with “!”. Species are annotated as follow: 1d – monoicous, 2d – dioicous, 1/2d – polyoicous; S+/- – sporophyte present/absent; LF – life form; C – coenoelement; E – ecology; G – geoelement; and frequency (rr – very rare, 1-5 localities, r – rare, 6-15, p – sporadic, 16-30, fq – frequent, 31 – 50, fqq – very frequent, >50). Other abbreviations are in forword to the species list. The analysis of bryoflora will be done in the second part of this paper. Ðåçþìå Ïðåäñòàâëåí ñïèñîê ìîõîîáðàçíûõ Ñðåäíåðóññêîé âîçâûøåííîñòè – îáîñîáëåííîé ôèçèêî-ãåîãðàôè÷åñêîé îáëàñòè íà òåððèòîðèè Âîñòî÷íîé Åâðîïû. Äàí êðàòêèé î÷åðê ïðèðîäíûõ óñëîâèé è èñòîðèÿ áðèîëîãè÷åñêèõ èññëåäîâàíèé. Áîëüøàÿ ÷àñòü Ñðåäíåðóññêîé âîçâûøåííîñòè íàõîäèòñÿ â ïðåäåëàõ ëåñîñòåïíîé çîíû, êðàéíèé ñåâåð – â çîíå øèðîêîëèñòâåííûõ ëåñîâ, þã – â ñòåïíîé çîíå. Ñïåöèôè÷åñêîé îñîáåííîñòüþ ïðèðîäíûõ óñëîâèé ÿâëÿåòñÿ îáèëèå ìåëîâ è èçâåñòíÿêîâ, îáíàæàþùèõñÿ ïî ñêëîíàì ðå÷íûõ äîëèí è áàëîê. Êîíñïåêò áðèîôëîðû âêëþ÷àåò 370 òàêñîíîâ è îñíîâàí íà ñáîðàõ àâòîðà 1981-2002 ãã., ðåâèçèè âñåõ äîñòóïíûõ êîëëåêöèé ïî äàííîìó ðåãèîíó è ëèòåðàòóðíûõ äàííûõ. Äëÿ êàæäîãî âèäà ïðèâåäåíà îáîáùåííàÿ ýêîëîãî-áèîëîãè÷åñêàÿ õàðàêòåðèñòèêà (ðåãóëÿðíîñòü ñïîðîíîøåíèÿ, ïîë, ýêîëîãè÷åñêèé òèï, ýêîëîãî- öåíîòè÷åñêàÿ ãðóïïà, æèçíåííàÿ ôîðìà, ãåîãðàôè÷åñêèé ýëåìåíò, âñòðå÷àåìîñòü), ñîñòàâëåííàÿ íà îñíîâå àíàëèçà ïîâåäåíèÿ âèäà â ðàññìàòðèâàåìîì ðåãèîíå. Äëÿ ðåäêèõ è ñïîðàäè÷è÷åñêè âñòðå÷àþùèõñÿ âèäîâ ïåðå÷èñëåíû âñå èçâåñòíûå ìåñòîíàõîæäåíèÿ. Àíàëèç áðèîôëîðû áóäåò ïîìåùåí âî âòîðîé ÷àñòè ñòàòüè. ÂÂÅÄÅÍÈÅ íî 250-300 òûñ. êì2, íàõîäÿòñÿ ñëåäóþùèå àäìè- Íà îáøèðíûõ ïðîñòðàíñòâàõ Âîñòî÷íîé íèñòðàòèâíûå îáëàñòè Ðîññèéñêîé Ôåäåðàöèè: Åâðîïû Ñðåäíåðóññêàÿ âîçâûøåííîñòü (48° Òóëüñêàÿ, êðàéíèé þã Ìîñêîâñêîé (Çàðàéñêèé, -53° ñ.ø. è 34°-40° â.ä.) ïðåäñòàâëÿåò ñîáîé Ñåðåáðÿíîïðóäñêèé, Êàøèðñêèé, Ëóõîâèöêèé ð- âïîëíå îáîñîáëåííóþ ôèçèêî-ãåîãðàôè÷åñêóþ íû), ÷àñòè÷íî Ðÿçàíñêàÿ (äî Ðÿçàíè íà âîñòî- îáëàñòü (Ðèñ. 1). Íàèáîëåå îò÷åòëèâî ãðàíè- êå), ÷àñòè÷íî Êàëóæñêàÿ (Ïåðåìûøëüñêèé, Òà- öû Ñðåäíåðóññêîé âîçâûøåííîñòè ïðîÿâëÿ- ðóññêèé, Êîçåëüñêèé, Óëüÿíîâñêèé ð-íû), Îðëîâ- þòñÿ íà âîñòîêå ïî ð. Äîíó è ð. Âîðîíåæó è ñêàÿ, ÷àñòè÷íî Áðÿíñêàÿ (êðàéíèé çàïàä – Ñåâ- íà ñåâåðî-âîñòîêå – ïî ð. Îêå (Ìèëüêîâ, ñêèé, Êîìàðè÷ñêèé, ÷àñòè÷íî Áðÿíñêèé, Áðàñîâ- 1961). Çàïàäíàÿ, þãî-çàïàäíàÿ è þæíàÿ ãðà- ñêèé, Íàâëèíñêèé, Ñóçåìñêèé ð-íû), Ëèïåöêàÿ íèöû èìåþò ñëàáîå âûðàæåíèå, ïîñêîëüêó (íà âîñòîêå äî äîëèíû ð. Âîðîíåæà), Êóðñêàÿ, Ñðåäíåðóññêàÿ âîçâûøåííîñòü (ÑÂ), ïîñòå- Áåëãîðîäñêàÿ, Âîðîíåæñêàÿ (çà èñêëþ÷åíèåì ïåííî ïîíèæàÿñü, ïëàâíî ïåðåõîäèò â ïðè- Îêñêî-Äîíñêîé ÷àñòè); à òàêæå Óêðàèíû: Ñóì- ëåãàþùèå íèçìåííûå ðàâíèíû.  ãðàíèöàõ ñêàÿ (äî ã. Ëåáåäèíà íà þãå), Õàðüêîâñêàÿ è äàííîé òåððèòîðèè, îáùåé ïëîùàäüþ ïðèìåð- Ëóãàíñêàÿ (íà þãå äî ð. Ñåâåðñêèé Äîíåö). 1 — Âîðîíåæ, 394043 Òèìèðÿçåâà 8, Âîðîíåæñêàÿ Ãîñóäàðñòâåííàÿ Ëåñîòåõíè÷åñêàÿ Àêàäåìèÿ — Voronezh State Silvicultural Academy, Timiryazeva str., 8, Voronezh 394043 Russia 102 Í. Í. ÏÎÏÎÂÀ ÈÑÒÎÐÈß ÈÇÓ×ÅÍÈß ÁÐÈÎÔËÎÐÛ ñàíèè øèðîêîëèñòâåííûõ ôîðìàöèé Êðàïèâåíñ- ÑÐÅÄÍÅÐÓÑÑÊÎÉ ÂÎÇÂÛØÅÍÍÎÑÒÈ êîãî ëåñíè÷åñòâà èìè âûÿâëåíî 39 âèäîâ Èñòîðèÿ èçó÷åíèÿ áðèîôëîðû Ñ íà÷èíàåòñÿ  ñòàòüå Í. Â. Ñàìñåëü (1968) ïî áðèîôëîðå ñî âòîðîé ïîëîâèíû XIX âåêà. Ðåçóëüòàòû ó÷åòà Ñåâåðî-Äîíñêîãî ðåëèêòîâîãî ðàéîíà èìåþòñÿ ñâå- èñòîðè÷åñêèõ ñâåäåíèé ñóììèðîâàíû ïî àäìèíè- äåíèÿ î ìõàõ íåáîëüøîãî ó÷àñòêà äîëèíû ð. Êðà- ñòðàòèâíûì îáëàñòÿì. Ïðèâîäÿòñÿ ðàáîòû, íàèáî- ñèâîé Ìå÷è â îêð. ñåë Øèëîâî è Âÿçîâî. Èç îá- ëåå ñóùåñòâåííûå äëÿ îöåíêè âèäîâîãî ðàçíîîá- ùåãî ÷èñëà âèäîâ (îêîëî 27), ñîáðàííûõ çäåñü, ïî- ðàçèÿ ìîõîîáðàçíûõ è òåíäåíöèé åãî èçìåíåíèÿ. ïîëíèëè ñïèñîê ìîõîîáðàçíûõ Òóëüñêîé îáëàñ- òè 5 âèäîâ (Blasia pusilla, Aneura pinguis, Encalypta ÒÓËÜÑÊÀß ÎÁËÀÑÒÜ vulgaris, Bryoerytrophyllum recurvirostrum, Äëÿ Òóëüñêîé îáëàñòè èìååòñÿ äîñòàòî÷íî îáøèð- Plagiochila porelloides) Êðîìå òîãî, íàìè îïðåäå- íàÿ è êâàëèôèöèðîâàííàÿ áðèîôëîðèñòè÷åñêàÿ ñâîäêà ëåíû íåáîëüøèå êîëëåêöèè ìõîâ È. Ñ. Øåðåìå- Í. Â. Öèíãåðà (1893).  íåé ïî îïðåäåëåíèÿì Ñ. Ã. Íà- òüåâîé èç îêð. ã. Òóëû è ïîñ. Äóáíû (îêîëî 20 âàøèíà ïðèâîäèòñÿ ñïèñîê 134 âèäîâ ëèñòîñòåáåëü- âèäîâ) è Å. Â. Âûøåãîðîäñêèõ èç Ñóâîðîâñêîãî è íûõ ìõîâ, ñîáðàííûõ â Òóëüñêèõ Çàñåêàõ (Ïîäãîðîä- Áåëåâñêîãî ð-íîâ (îêîëî 30 âèäîâ). Èññëåäîâàíèÿ, íîå, Âåíåâñêîå, Êðàïèâåíñêîå ëåñíè÷åñòâà), íà èçâåñò- ïðîâåäåííûå â Òóëüñêîé îáëàñòè Ïîïîâîé, ïîçâî- íÿêîâûõ îáíàæåíèÿõ ïî ð. Îñåòðó (ñ. Õðóñëîâêà, ñ. ëèëè äîáàâèòü ê ôëîðèñòè÷åñêîìó ñïèñêó 60 íî- Áÿêîâî), Åïèôàíñêîì áîëîòå, â ñîñíîâûõ áîðàõ ó ñò. âûõ âèäîâ. Òàêèì îáðàçîì, â òóëüñêîé áðèîôëîðå Ñâèíñêîé è ã. Àëåêñèíà, â îêðåñòíîñòÿõ ñîáñòâåííîãî ê íàñòîÿùåìó âðåìåíè óñòàíîâëåíî 215 âèäîâ èìåíèÿ ó ñ. Ìåëåõîâêà, à òàêæå â ßñíîé Ïîëÿíå. Ìîíî- (Ïîïîâà, 1999à). ãðàôèÿ ïðåäñòàâëÿåò ñîáîé ïîäðîáíûé êîíñïåêò áðè- îôëîðû ñ óêàçàíèåì êîíêðåòíûõ ìåñòîíàõîæäåíèé, ËÈÏÅÖÊÀß ÎÁËÀÑÒÜ ÷àñòîòû âñòðå÷àåìîñòè, ýêîëîãèè, âðåìåíè è ðåãóëÿð- Ïåðâûå ñâåäåíèÿ î ìõàõ Ëèïåöêîé îáëàñòè ïðè- íîñòè ñïîðîíîøåíèÿ ìõîâ. Íåêîòîðûå êðèòè÷åñêèå íàäëåæàò åëåöêîìó ó÷èòåëþ Ë. Ô. Ãðóíåðó (1873). âèäû ñíàáæåíû ïîäðîáíûìè êîììåíòàðèÿìè. Ãåðáà-  ñïèñêå ðàñòåíèé, ñîáðàííûõ â îêð. ã. Åëüöà èì ðèé Öèíãåðà ñîõðàíèëñÿ â ìåìîðèàëüíîì óãîëêå Íà- ïåðå÷èñëåíû 36 âèäîâ ìîõîîáðàçíûõ ñ óêàçàíèåì âàøèíà â Êèåâñêîì óíèâåðñèòåòå. òî÷íûõ ìåñòîíàõîæäåíèé è ýêîëîãî-öåíîòè÷åñêîé  îáîáùàþùåé ñâîäêå ïî ìõàì Ðîññèè Öèêåí- ïðèóðî÷åííîñòè. Èç ðåäêèõ è èíòåðåñíûõ âèäîâ – äðàòà (Zickendrath, 1900) ó÷òåíû âèäû, ñîáðàííûå Neckera pennata, Ditrichum flexicaule, Trichostomum â Òóëüñêîé ãóáåðíèè â 1890-å ãîäû Öèíãåðîì è À. crispulum (ïîñëåäíèé âèä óêàçàí â ëåñàõ, ÷òî äëÿ Ñ. Áàðêîâûì è íå âîøåäøèå â îïóáëèêîâàííóþ ýòîãî ñòåïíîãî êàëüöåôèòà íå ñîâñåì îáû÷íî). ñàìèì Öèíãåðîì ìîíîãðàôèþ; ñïèñîê ïîïîëíèëè Ñ îòêðûòèåì â êîíöå XIX âåêà Öèíãåðîì è Ä. Í. 5 âèäîâ, Trematodon ambiguus, Pyramidula tetragona Ëèòâèíîâûì ñêàëüíûõ è ãîðíî-ñòåïíûõ êîìïëåê- è äðóãèå, â òîì ÷èñëå Mnium submarginatum ñîâ â äîëèíå ð. Äîí è åãî ïðèòîêîâ áûëî íà÷àòî Navasch. et Zing., âèä, âïîñëåäñòèè îòíåñåííûé Ëà- ïðèñòàëüíîå èçó÷åíèå ôëîðû è ðàñòèòåëüíîñòè èç- çàðåíêî (1940) ê Mnium heterophyllum (Hook.) âåñòíÿêîâûõ îáíàæåíèé ñåâåðî-çàïàäíîé ÷àñòè Ëè- Schwaegr. Ýòî åäèíñòâåííîå ìåñòîíàõîæäåíèå âèäà ïåöêîé îáëàñòè, ïîëó÷èâøåé íàçâàíèå Ñåâåðî-Äîíñ- â Åâðîïå, åñëè íå ñ÷èòàòü Êàâêàçà. êîãî ðåëèêòîâîãî ðàéîíà. Íåîäíîêðàòíî ïîñåùàÿ âî  ñòàòüå À. À. Êðóáåðà (1900), ïîñâÿùåííîé êàð- âðåìÿ áîòàíè÷åñêèõ ýêñêóðñèé Ãàëè÷üþ Ãîðó, Ñ. È. ñòîâûì ÿâëåíèÿì â Ðîññèè, äëÿ ïðîâàëüíûõ áîëîò Ðîñòîâöåâ, Öèíãåð, Íàâàøèí, Â. È. Õèòðîâî íàðÿäó Òóëüñêîé ãóáåðíèè ïðèâîäèòñÿ îêîëî äåñÿòè âè- ñ ñîñóäèñòûìè ðàñòåíèÿìè ñîáèðàëè è ìõè. Ýòè äîâ ìõîâ. Ïîäîáíûå óíèêàëüíûå ëàíäøàôòû ïðè- ñáîðû, íå íîñèâøèå öåëåíàïðàâëåííîãî õàðàêòåðà, âëåêàëè âíèìàíèå íå òîëüêî çåìëåâåäîâ, íî è ñîõðàíèëèñü äî íàñòîÿùåãî âðåìåíè â ìåìîðèàëü- áîòàíèêîâ. Õàðàêòåðèçóÿ ðàñòèòåëüíîñòü ðåëèêî- íîì óãîëêå Íàâàøèíà â Êèåâñêîì ãîñóíèâåðñèòåòå âîãî êîìïëåêà Åïèôàíñêîãî áîëîòà, À.Ê. Ñêâîð- è áûëè âêëþ÷åíû â ñâîäêó Öèêåíäðàòà (Zickendrath, öîâ (1948) ïåðå÷èñëÿåò 7 âèäîâ ìõîâ (âñå îíè 1900) ïî ìõàì Ðîññèè (âñåãî 16 âèäîâ, ñðåäè êîòî- ôèãóðèðîâàëè â ñâîäêå Öèíãåðà).  íàñòîÿùåå âðå- ðûõ Pottia lanceolata, Orthotrichum cupulatum, ìÿ îëèãîòðîôíûå áîëîòíûå êîìïëåêñû íà Åïè- Homalothecium sericeum è äð). ôàíñêîì áîëîòå óæå íå ñóùåñòâóþò.  îáîáùàþùåé ñâîäêå “Ïðèðîäà Îðëîâñêîãî Ðåçóëüòàòû èññëåäîâàíèé ñòåïíîé è ëåñíîé êðàÿ” (1925) ïîä ðåäàêöèåé Õèòðîâî (ðàíåå âñÿ ðàñòèòåëüíîñòè â âåðõîâüÿõ ðåê Ñêíèãè è Áåñ- çàïàäíàÿ ÷àñòü íûíåøíåé Ëèïåöêîé îáëàñòè âõî- ïóòû ïðèâåäåíû â ñòàòüÿõ Â.Â.Àëåõèíà (1944, äèëà â Îðëîâñêóþ ãóáåðíèþ) â ãëàâå “Ðàñòèòåëü- 1945), ãäå óêàçàíû è íåêîòîðûå âèäû ìõîâ, äîìè- íîñòü” ïîëíîñòüþ ïîìåùåí ñïèñîê ìõîâ Ãðóíåðà, èíèðóþùèõ â íàïî÷âåííîì ïîêðîâå. Âîñåìüþ íî- èç ÷èñëà íîâûõ äëÿ îáëàñòè óêàçàíû Orthotrichum âûìè âèäàìè, â îñíîâíîì ïå÷åíî÷íèêîâ, ïîïîëíè- fallax, Encalypta vulgaris, Bryum creberrimum. ëè ñïèñîê ìîõîîáðàçíûõ Òóëüñêèõ Çàñåê Ë. È. Ñïåöèàëüíûå áðèîëîãè÷åñêèå èññëåäîâàíèÿ îò- Àáðàìîâà è Ñ. Ô. Êóðíàåâ (1977); âñåãî ïðè îïè- äåëüíûõ èçâåñòíÿêîâûõ óðî÷èù Ñåâåðî-Äîíñêîãî Áðèîôëîðà Ñðåäíåðóññêîé âîçâûøåííîñòè. I 103 Ðèñ. 1. Òåððèòîðèÿ Ñðåäíå- ðóññêîé âîçâûøåííîñòè. Çàëè- òûå êðóæêè – ìåñòîíàõîæäå- íèÿ, îáñëåäîâàííûå àâòîðîì; îòêðûòûå êðóæêè – ìåñòî- íàõîæäåíèÿ, ïî êîòîðûì èñ- ïîëüçîâàíû ëèòåðàòóðíûå äàííûå. Fig. 1. Territory of the Central Russian Upland. Solid circles – localities studied by author; open circles – literature data. Îáëàñòíûå – Province öåíòðû ñentres ÌÎÑ Ìîñêâà – Moscow ÊËÆ Êàëóãà – Kaluga ÒÓË Òóëà – Tula ÎÐË Îðåë – Orel ÁÐÍ Áðÿíñê – Bryansk ËÈÏ Ëèïåöê – Lipezk ÊÓÐ Êóðñê – Kursk ÁÅË Áåëãîðîä – Belgorod ÂÎÐ Âîðîíåæ – Voronezh ÑÓÌ Ñóìû – Sumy ÕÀÐ Õàðüêîâ – Kharkov ËÓà Ëóãàíñê – Lugansk ðåëèêòîâîãî ðàéîíà áûëè ïðåäïðèíÿòû Ñàìñåëü Çíà÷èòåëüíî ìåíüøå ñâåäåíèé ïî áðèîôëîðå (1968). Ïðåäñòàâëåííûé åþ