Bryophyte Flora of Avala Mountain (Central Serbia, Yugoslavia) (2003)

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    LINDBERGIA 28: 9096. Lund 2003

    Bryophyte flora of Avala Mt. (C. Serbia, Yugoslavia)

    M. Sabovljeviand T. Cveti

    Sabovljevi, M. and Cveti, T. 2003. Bryophyte flora of Avala Mt. (C. Serbia,Yugoslavia). Lindbergia 28: 9096.

    Avala Mountain (511m altitude) is covered by deciduous forests. The bryophyteflora had prevously been known to include 54 moss and 11 liverwort speciesand the present investigation adds 60 new mosses and 3 new liverworts. Six ofthese species are new records for Serbia, and three are second records. Moistand shady deciduous woods provide conditions for mesophytic mosses that arethe most numerous within the investigated area.

    M. Sabovljevi, Dept of Plant Ecology and Phytogeography, Inst. of Botanyand Botanical Garden, Faculty of Biology, Univ. of Belgrade, Takovska 43,YU-11000 Belgrade, Yugoslavia ([email protected]). T. Cveti, Dept ofBiology, Petnica Science Centre, PO Box 118, YU-14000 Valjevo, Yugoslavia.

    Bryology and bryophyte research in Yugoslavia hasbeen neglected for a long time. However, there aresome signs of bryology reviving lately (Paviet al.1998, Sabovljevi1999, 2000, Sabovljeviand Steva-novi1999, 2000, Sabovljeviet al. 1999). Nonethe-less, the bryophyte flora of Yugoslavia remains verypoorly known. Approximately 512 mosses and 119hepatic species have been recorded (SabovljeviandStevanovi1999, Sabovljevi2000), and detailed re-search often results in addition of species new to area.The bryophyte flora of Avala has previously been stud-ied very poorly even if it is close to the center of Ser-bian botany, Belgrade, and almost all records came

    Accepted 29 January 2003

    Copyright LINDBERGIA 2003

    Fig. 1. The position of the investigated area (Avala Mt.) inthe Federal Republic of Yugoslavia (dot on UTM map 34T.Mountain areas above 1000 m appear in grey. Abbrevia-tions: AL: Albania; BiH: Bosnia & Herzegovina; BG: Bul-garia; H: Hungary; HR: Croatia; MK: The Former Yugo-slav Republic of Macedonia; RO: Romania; YU: The Fed-eral Republic of Yugoslavia (Serbia and Montenegro).

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    from investigators passing by. Previous records attribut54 moss and 11 liverwort species (Jurii1900, Soka1949, Pavleti1955, Popovi, 1966)

    Background

    Avala Mountain is situated 16 km south of Belgrade(UTM: 34T DQ64, Fig. 1), between 203010203415E and 443817N444203N. An extinctvolcano, nowadays it is a quiet conical hill, coveredin woods from base to top (511 m). It is the first moun-tain arising from the huge Pannonian plain on its tran-sition to the southern mountain ranges (the Rhodopeanand the Dinaric). The mountain has a good hydrologi-cal network; 9 streams occur in the investigated area

    of ca 6 km2

    .ThegeologyofAvalamountainisveryvariable:lime-stones, marls, sandstones and serpentines. Rocks oc-cur in different metamorphic states throughout themountain, with a series of varieties from limestone tosiliceous rocks. Basic soil types include brown loesssoil, pseudogley, acidic brown soil and cambisol.

    The climate is mild-continental with a great conti-nental influence due to its exposure to the Pannonianplain. There is a short period of semidrought duringJuly (Fig. 2). January and February averages are be-low 0C and frost can appear from late October toearly April.

    Avala Mt. is mostly covered by forest vegetation,

    both native and planted. Meadow vegetation is alsopresent, but less extensive.

    The forest vegetation types are: 1. Fagetum mixtum silicicolumMii1963 2. Fagetum submontanum JovanovitilietosumMii

    1960 3. Querco-Carpinetum moesiacumRudski aculeatu-

    tosumB. Jovanovi1951 4. Orno-Quercetum petraeae Borisavljevi1955

    5. Quercetum frainetto-cerris RudskiaculeatutosumB. Jovanovi1951

    6. Orno-Quercetum virgilianae Gaji1963 7. planted Scottish and black pine,Abies concolor

    Lindl., Pseudotsuga glaucaMayr., Thuja orienta-lisL., Taxus baccataL.

    Meadow cenoses are: 8.Xeranthemeto-IschaemetumR. Jovanovi1955 9. Festucetum vallesiacaes.l. R. Jovanovi195510. Poa angustifoliaAchillea collinacommunity11.Bromus arvensisBromus molliscommunityThe flora of the mountain includes 51 tree species and594 vascular plant species.

    Methods

    Fieldwork was performed mostly during 2001. SinceAvala mountain, and therefore the investigated area,covers ca 6 km2, with various vegetational, geologi-cal, climatic or hydrological characteristics, thetransect method has been used. This method enablesextensive surface coverage by random choice of habi-tats with various ecologies within the investigated area.Collected material is deposited in the Institute ofBotany and Botanic Garden of Belgrade UniversityHerbarium (BEOU), as well as in authors private col-lections.

    We followed the nomenclature of Corley et al.(1981) and Corley and Crundwell (1991) for mosses,

    except for Schistidium and Hypnum where we fol-lowed Blom (1996) and Smith (1997), respectively.Liverwort nomenclature follows Grolle and Long(2000). For the bryophyte distribution types we usedDll (1983, 1984, 1985, 1992) and Dll et al. (1999)and for ecological analyses Hallingbck (1996) andIngerpuu et al. (1994).

    Results and discussion

    The results presented here are based both on field work

    and literature data. We found 114 moss and 14liverwort species, of which 60 moss taxa and 3 liver-worts are new records for Avala Mt. Six species arenew for Serbia.

    Floristic analysis

    Despite the good hydrological network, only 11% liv-erworts were found on Avala mountain, and most ofthese are epiphytic on different tree species. Never-theless, such a low percentage is expected consider-Fig. 2. Climate diagram of Avala Mountain.

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    ing the low percentage of liverworts in bryoflora ofthe Federal Republic of Yugoslavia (19% of total Yu-goslav bryophyte species; Sabovljevi2000). Withinliverworts,Jungermanniales (72%) predominate fol-lowed byMarchantiales andMetzgeriales (both 14%).

    Within the classBryopsida,orderHypnalesis thebest represented comprising 34% of the bryophyteflora of Avala. This is no surprise, since woodland,with a mostly shaded and wet microclimate, predomi-nates. Such habitats are preferred by pleurocarpous

    mosses. The relative high number of Pottiales spe-cies (17%) is mainly due to the presence of favouritetourist picnic areas in urban and suburban zones ofthe mountain. Also, many rock walls, sunny roadsidesites and exposed rock outcrops provide good habi-tats for pottiaceous mosses (Fig. 3).

    Biogeography

    A biogeographical consideration of the main choro-logical elements represented in the study area has beenperformed by estimating percentages belonging to

    different distributional types. The taxa found belongto many different chorological types, but in view ofthe similarity of many of these elements we haveunited them for statistical and graphical purposes intoeight groups: temperate, boreal, subboreal, submedi-terranean, suboceanic, oceanic, mediterraneanoce-anic and subpontic. The absolute numbers of taxa be-longing to these large biogeographic groups are shownin Fig. 4.

    The most outstanding feature is the dominance oftemperate species (49%). This is expected consider-ing the climate, vegetation, lithology and geographi-cal position of the investigated area. Submediterra-

    nean (including submediterranean-suboceanic, 18%)and subboreal (12%) distribution types are second andthird among the bryophytes of Avala. These are theconsequences of the wide submediterranean north edgereaching this region and the altitude of 500 m in themild continental climate belt with various topogra-phy. The other chorological elements together makeup 21% of the flora.

    Ecological remarks

    Ecological preferences for humidity and acidity havebeen studied according to Hallingbck (1996) and

    Ingerpuu et al. (1994). Humidity preference analysisis shown in Fig. 5. It is presented as a percentage ofhygro- , meso- and xerophytic species and intermedi-ates (hygromesophytes, mesohygrophytes, mesoxerophytes and xeromesophytes).The most numer-ous are mesophytic species, (55% including meso

    Fig. 3. Distribution of bryophytes of Avala Mountain amongorders.

    Fig. 4. Distribution of bryophytes of Avala Mountain overchorological elements.

    Fig. 5. Humidity preferences of the bryophytes on AvalaMountain.

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    15. Brachythecium populeum (Hedw.) B., S. & G. P(Soka 1949, Popovi1966)

    16. Brachythecium rivulare B., S. & G. (Soka 1949) Soilby the stream in the woods

    17. Brachythecium rutabulum (Hedw.) B., S. & G. (Soka1949, Popovi1966) Soil in the woods at the banksof stream, dark, very moist

    18. Brachythecium salebrosum (Web. & Mohr) B., S. &G. Rock occasionally flushed by stream or partiallysubmersed, shaded

    19. Brachythecium velutinum (Hedw.) B., S. & G. (Jurii1900, Simi, 1900) Stamped soil on paths throughforest and on tree roots

    20. Bryoerythrophyllum recurvirostrum (Hedw.) Chen.Sandy protosoil on asphalt patch; damp soil at theedge of asphalt patch

    21. a Bryum argenteum Hedw. var. argenteum(Popovi1966) Damp soil covered with herbaceousvegetation,protosoil between blocks in pavement, exposed, dry;exposed sandy protosoil on asphalt patch

    21. b Bryum argenteum Hedw. var.lanatum (P. Beauv.)Hampe idem-

    22. Bryum caespiticium Hedw. (Popovi,1966) Damp soilcovered with herbaceous vegetation

    23. Bryum capillareHedw. (Popovi, 1966) Compressedsoil between grass clods; soil and protosoil betweenblocks in pavements, exposed, dry; shaded soil at treebase

    24. Bryum subelegans Kindb., Forest litter, together withHypnum cupressiforme

    25. Calliergon cuspidatum (Hedw.) Kindb. P (Soka1949)

    26. Ceratodon purpureus (Hedw.) Brid. (Soka 1949,Popovi1966), Protosoil at the edges of the road; onsoil in meadow vegetation; soil at base of Carpinusbetulus L.

    27. Cirriphyllum crassinervium(Tayl.) Loeske & Fleisch.Soil in woods, near the stream, dark, moist

    28. Ctenidium molluscum (Hedw.) Mitt. (Popovi1966) 29. Dicranella cerviculata(Hedw.) Schimp. Soil in woods

    at tree bases, shady 30. Dicranella heteromalla(Hedw.) Schimp. Soil in base

    of tree or elsewhere in woods where shaded (some-times vertical); base of Crategus monogyna Jacq.,Carpinus betulus

    31. Dicranum scoparium Hedw. (Popovi1966) 32. Didymodon acutus(Brid.) K. Saito Soil in the base of

    Carpinus betulus 33. Didymodon insulanus(De Not.) M. Hill Exposed con-

    crete wall; sandy protosoils 34. Didymodon luridusHornsch. ex Spreng. Protosoil on

    the roadsides and bare exposed, dry soils 35. Didymodon vinealis (Brid.) Zander Protosoil on theroadsides and bare exposed, dry soils

    36. Encalypta ciliataHedw. P(Pavleti1955 ) 37. Enthosthodon fascicularis (Hedw.) Mll. (Jurii

    1900, Soka 1949) Soil, in woods dark, moist, soil incanalization hole, dark, very moist

    38. Entodon concinnus(De Not.) Par. Shaded, moist rock 39. Eurhynchium praelongum(Hedw.) B., S. & G. Dis-

    turbed soil, lawn, exposed 40. Eurhynchium pulchellum(Hedw.) Jen. Disturbed soil

    on lawn, in vegetation; vertical basic rocks andprotosoil

    41. Eurhynchium schleicheri (Hedw.) Jur., Soil in thewoods

    42. Eurhynchium speciosum (Brid.) Jur. (Popovi1966) 43. Eurhynchium striatulum(Spruce) B., S. & G. Soil in

    the woods 44. Eurhynchium striatum (Hedw.) Schimp. (Popovi

    1966) 45. Fissidens bryoidesHedw. (Popovi1966) Forest lit-

    ter; soil by forest stream 46. *Fissidens exilisHedw. Soil by forest stream 47. Fissidens taxifoliusHedw. (Soka 1949) Rock crev-

    ices with soil occasionally flushed by stream, shaded,soil under forest litter

    48. *Fissidens viridulus (Sw.) Wahlenb. Soil in forest,shaded

    49. Funaria hygrometricaHedw. (Pavleti1955, Popovi1966) Compressed soil between grass clods, togetherwithB. argenteum

    50. Grimmia orbicularisBruch. ex Wils. Exposed rockoutcrops

    51. Grimmia pulvinata (Hedw.) Sm. Damp soil; sandyprotosoil on asphalt patch; calcareous rocks and con-crete walls, exposed, dry

    52. *Habrodon perpusillus(De Not.) Lindb. Tree bark 53. Herzogiella seligeri (Brid.) Iwats. Trunk by forest

    stream 54. Homalothecium lutescens (Hedw.) H. Rob. (Popovi

    1966) 55. Homalothecium sericeum (Hedw.) B., S. & G. P

    (Soka 1949) 56. Hylocomium splendens (Hedw.) Schimp. (Popovi

    1966) 57. Hypnum cupressiformeHedw. (Soka 1949, Popovi

    1966) Exposed moist soil; soil in woods and bystreams, shaded; tree roots shaded

    58. Hypnum lacunosum(Brid.) G.F. Hoffman. Tree stump,

    exposed moist soil; soil in woods and by streams,shaded

    59. Hypnum resupinatumTayl. Bark at base ofAcer cam-pestre L., woods; soil, roots and bark at base ofFagussylvatica L.; base of Cornus mas L.

    60. Isothecium alopecuroides (Dubois) Isov. (Soka 1949,Popovi1966) Soil in woods

    61. Isothecium myosuroidesBrid. (Soka 1949) Soil bythe forest stream, together with Rhynchostegiumconfertum

    62. Leskea polycarpa Hedw. Roots of a tree stump inwoods; exposed soil

    63. Leucodon sciuroides(Hedw.) Schwaegr. (Soka 1949,Popovi1966) Quercus petraea(Matt.) Lieblein bark;tree stump; shaded, moist

    64. Neckera complanata(Hedw.) Hb. P(Soka 1949) 65. Orthotrichum anomalum Hedw. (Popovi1966) 66. Orthotrichum affineBrid. (Soka 1949) Carpinus

    betulustree. bark 67. Orthotrichum diaphanum Brid. Concrete wall, shaded,

    moist 68. Orthotrichum lyelliiHook. & Tayl. Quercus petraea

    bark 69. Orthotrichum pallens Bruch ex Brid. Quercus sp.

    bark, exposed 70. Orthotrichum pulchellumBrunt. Fraxinus ornus L.

    tree bark 71. Orthotrichum pumilumSw. exposed branch of Cas-

    tanea sativa Mill.

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    72. Physcomitrium sphaericum(Ludw.) Brid. Moist, darkplaces, on soil in woods

    73. Plagiomnium cuspidatum (Hedw.) T. Kop. (Soka1949, Popovi1966) Soil in woods near and on thebanks of the stream, dark moist

    74. Plagiomnium rostratum (Schrad.) T. Kop. P (Soka1949)

    75. Plagiomnium undulatum(Hedw.) T. Kop. P (Soka1949, Popovi1966)

    76. Plagiothecium cavifolium (Brid.) Iwats. Soil by thestream in the woods

    77. Plagiothecium laetumB., S. & G. Soil in wood, dark,moist

    78. Pogonatum aloides (Hedw.) P. Beauv. (Popovi1966) 79. Pogonatum urnigerum (Hedw.) P. Beauv. P (Soka

    1949) 80. Pohlia elongata Hedw. Forest litter, soil at base of

    Fagusby the stream, occasionally splashed by water 81. Polytrichum commune Hedw. (Popovi1966) 82. Polytrichum formosumHedw. Soil cliff by the stream

    83. Polytrichum piliferum Hedw. (Popovi1966) 84. Pottia intermedia(Turn.) Frnr. Pebbly soil by road,

    dry, exposed 85. Pottia truncata(Hedw.) B. & S. Damp soil covered

    with herbaceous vegetation 86. Pseudocrossidium hornschuchianum (K.F. Schultz)

    Zander Protosoils in urban and suburban zones andthe edges of the tracks

    87. *Pseudocrossidium revolutum(Brid.) Zander Proto-soils in urban and suburban zones and the edges ofthe tracks

    88. Pseudoleskeella catenulata(Schrad.) Kindb.Carpi-nus betulusbark, rock and concrete wall, dry

    89. Pseudoleskeella nervosa(Brid.) Nyh. (Popovi1966)Exposed concrete wall, Quercus petraea bark; Ex-posed sandy protosoil on asphalt patch

    90. Pteryginandrum filiforme Hedw. (Popovi1966) 91. Pylaisia polyantha(Hedw.) Schimp. (Popovi1966)

    Bark at base of Tilia, Carpinus and Quercus bark,shaded;

    92. Rhizomnium punctatum(Hedw.) T. Kop. (Soka 1949,Popovi1966) Rocks and soil occasionally flushedby stream or extremely moist, shaded

    93. *Rhynchostegiella teesdalei(B., S. & G.) Limpr. Trunkpartially submersed, dark; soil and rocks by foreststream.

    94. **Rhynchostegiella tenella(Dicks.) Limpr. Rock oc-casionally flushed by stream or partially submersed,shaded

    95. *Rhynchostegium confertum(Dicks.) B., S. & G. Soiland rocks by ephemeral streams in woods, sometimes

    covered with herbaceous vegetation; dry concrete wall 96. Rhynchostegium megapolitanum(Web. & Mohr.) B.,S. & G. Soil covered with herbaceous vegetation, to-gether withEurhynchium praelongum

    97. Rhynchostegium riparioides(Hedw.) Card. (Popovi1966) Calcareous rock and brick, submersed in for-est stream

    98. Rhytidiadelphus triquetrus (Hedw.) Warnst. (Popovi1966)

    99. Schistidium apocarpumHedw. (Jurii140.1900, Simi 1900, Soka 1949) Protosoil on con-crete wall, exposed

    100. Schistidium brunnescensLimpr. Basic rock and con-crete wall in the woods, very dark, not very moist

    101. **Schistidium crassipilumBlom Wet and shaded con-crete by the road and rock outcrops

    102. Taxiphyllum wissgrilli(Garov) Wijk. & Marg. Soilby the stream in the woods

    103. Tetraphis pelucida Hedw. (Popovi1966)104. Thuidium recognitum(Hedw.) Lindb. P(Soka 1949)105. Thuidium tamariscinum (Hedw.) Schimp. (Popovi

    1966)106. Tortella tortuosa (Hedw.) Limpr. (Popovi1966)107. Tortula intermedia(Brid.) De Not. Wood soils among

    tree roots and shaded rock fissures with soil108. Tortula latifolia Bruch ex Hartm. Exposed sandy

    protosoil on asphalt patch;109. Tortula muralisHedw. Dry concrete wall, exposed

    or not110. Tortula ruralis (Hedw.) Gaertn., Meyer & Scherb.

    (Popovi1966) Protosoils by roads, shady, dry; ex-posed rock

    111. Tortula subulataHedw. (Pavleti1955, Soka 1949,Popovi1966) Soil, roots and bark at base of Fagus

    sylvatica112. Trichostomum brachydontium Bruch Calcareous

    rocks and concrete walls, exposed or shaded but dry;soil in rock features in woods.

    113. Trichostomum crispulumBruch Shaded, soil by treestump; rock cervices in exposed and shaded places

    114. Weissia brachycarpa(Nees & Hornsch.) Jur. P (Jurii1900, Soka 1949)

    Marchantiopsida:

    1. Apometzgeria pubescens (Schrank) Kiwah. (Popovi1966)

    2. Calypogeia azureaStotler& Crotz P(Soka 1949) 3. Frullania dilatata(L.) Dum. (Popovi1966) Quercus

    petraeabark, bark at base of field maple tree 4. Frullania tamarisci(L.) Dum. Quercus sp.bark 5. Lophocolea bidentata(L.) Dum. Tree stump, shady,

    moist; soil by ephemeral stream in woods; shaded soilin base of Carpinus betulus

    6. Lophocolea heterophylla(Schrad.) Dum. Tree stumpin woods, brick; dark

    7. Marchantia polymorphaL. (Popovi 1966) Basicrocks and soil at the edge of the forest stream

    8. Metzgeria furcata(L.) Dum. P(Pavleti1955, Popovi1966)

    9. Plagiochila porelloides (Torrey ex Nees) Lindenb.(Popovi1966) Calcareous rocks submersed or oc-casionally flushed by the stream

    10. Porella cordaeana(Hb) Moore (Pavleti1955, Soka

    1949) roots and soil at tree bases by the stream 11. Porella platyphylla(L.) Pfeiff. (Popovi1966) Soil inthe base of Quercus, shaded; tree trunk in the woods;base of Cornus mas

    12. Radula complanata (L.) Dum. Tree bark, usuallyQuercus

    13. Reboulia hemisphaerica(L.) Raddi. P(Soka 1949) 14. Scapania undulata (L.) Dum. (Popovi1966)

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