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The Melliferous Potential of Forest and Meadow PlantCommunities on Mount Tara (Serbia)Author(s): Snežana Jarić , Marina Mačukanović-Jocić , Miroslava Mitrovićl , andPavle PavlovićSource: Environmental Entomology, 42(4):724-732. 2013.Published By: Entomological Society of AmericaURL: http://www.bioone.org/doi/full/10.1603/EN13031

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PLANT-INSECT INTERACTIONS

The Melliferous Potential of Forest and Meadow Plant Communitieson Mount Tara (Serbia)

SNEZANA JARIC,1 MARINA MACUKANOVIC-JOCIC,2 MIROSLAVA MITROVIC,1

AND PAVLE PAVLOVIC1

Environ. Entomol. 42(4): 724Ð732 (2013); DOI: http://dx.doi.org/10.1603/EN13031

ABSTRACT The apißora of 34 forest and meadow plant communities in Tara National Park wasstudied with the aim of assessing their melliferous potential and their contribution to bee pastureduring the vegetation period. The melliferous plants were analyzed individually from the aspect oftheir ßowering phenology, abundance, and the intensity of nectar and pollen production, as well asthe production of honeydew. The melliferous potential of each investigated plant community wastheoretically assessed on the basis of the coenotic coefÞcient of melliferousness incorporating aphytocoenotic analysis, the coenotic coefÞcients of nectar and pollen production, and the percentageof melliferous species in relation to the total number of species that characterize the association. Thehighest percentage of the melliferous species was noted in the meadow associationPetasitetumhybridi(70%) and the forest association Piceetum-Abietis serpentinicum (63.6%). The highest values of thecoenotic coefÞcient of melliferousness were established for the forest associationQuerco-Carpinetumiliricum, and the meadow association Rhinantho-Cynosuretum cristati. Trees notable for their hon-eydew production in good quantities were Pinus nigra Arnold, Picea sp. Fagus sylvatica Linnaeus,Populus tremula Linnaeus, and Quercus cerris Linnaeus. Because, the vegetation in the study area isforest dominated, forest bee pasture including early ßowering herbaceous and woody plants, is of thegreatest signiÞcance for the honey bee, both in the early spring because of pollen and nectarproduction, and in the autumn as a source of honeydew.

KEY WORDS melliferous, apißora, plant community, nectar, pollen

The vegetation of Tara National Park (NP) comprisesforest (deciduous, deciduous-coniferous, and conif-erous) and meadow plant communities. The conifer-ous forests include pure or mixed communities ofPinus sylvestris Linnaeus and Pinus nigra Arnold onserpentine soils (Erico-Pinetum nigrae serpentinicum,Erico-Pinetum sylvestris serpentinicum, and Pinetumsylvestris-nigrae serpentinicum), as well as fragments ofcommunities of spruce on limestone and serpentine(Piceetum-Abietis montanum, Piceetum-Abietis serpen-tinicum, Piceetum omorikae-Abietis serpentinicum, andPiceetum omorikae-Abietis calcicolum).

A mixed and relict beech, Þr, and spruce forest(Piceo-Abieti-Fagetum) extends across the highestmountain plateau. One of the main ediÞcators of themixed deciduous-coniferous forests is the endemo-relict species the Serbian spruce (Picea omorika[Pancic] Purkyne). The most frequent deciduous for-est communities are those of manna ash and hop horn-beam (Orno-Ostryetum carpinifoliae), oak forests(Quercetum cerris, Quercetum frainetto-cerris), andwalnut forests (Querco-Juglandetum, Parietario-Jug-

landetum, and Alno-Juglandetum), as well as differenttypes of beech forests (Fagetum submontanum, Fag-etum montanum, Aceri-Ostryo-Fagetum, etc.). Thoseareas of Tara NP not populated by forest are inhabitedby diverse herbaceous vegetation of meadows, pas-tures, and mountain wetlands. In accordance with thediversity of the habitat, the refugial character of thewhole Tara region, and the speciÞc history of the ßoraand fauna, there is great ßoristic diversity in the TaraNP with over 1,100 species of vascular ßora, which isapproximately one-thirds of the total ßora of Serbia(Gajic et al. 1992).

The apißora of the researched area comprises ßow-ering plants and gymnosperms, whose ßoral and ex-traßoral products, above all nectar and pollen, are asource of essential nutrients for pollinators, particu-larly for the honey bee, as they are of existentialimportance in the development of colony and in theproduction of honey and other bee products(MacukanovicÐJocic 2010). The beginning and dura-tion of nectar ßow is determined by the biologicalcharacteristics of the plants and by climatic and geo-graphical factors. The quantity and quality of the avail-able nectar sources can be vastly different dependingon the season and blooming stage of the plant. Re-search of the ßora and vegetation of Tara has shownthat favorable conditions exist for continual bee pas-

1 Department of Ecology, Institute for Biological Research ÔSinisaStankovicÕ, Bulevar Despota Stefana 142, University of Belgrade,11060 Belgrade, Serbia.

2 Corresponding author: Faculty of Agriculture, University of Bel-grade, Nemanjina 6, 11080 Zemun, Serbia.

0046-225X/13/0724Ð0732$04.00/0 � 2013 Entomological Society of America

ture from spring to late autumn. During this period,major bee pastures yielding unißoral honeys alternatewith other nectar-producing plants of the Serbian api-ßora contributing to multißoral honeys.

Having reviewed the relevant literature, it was ev-ident that there are not many articles concerning theimportance of apißora and the contribution of certainmelliferous plants to the bee pasture of a speciÞcregion. Over the last two decades, as a contribution tothe development and advancement of apiculture, acertain amount of research has been devoted to var-ious aspects of studying the melliferous ßora of Serbiaand the wider region (Danon et al., 1990, 1994;Blazencic et al. 1994; Grabeljsek 1996; Macukanovic etal. 1996; Macukanovic and Grabeljsek 1996; Perisic etal. 2004; Farkas and Zajacz 2007; Grozeva 2011). Ourresearch offers a new approach to studying the mel-liferous signiÞcance and contribution of both mellif-erous plants and plant communities to bee pasture.The most important elements of bee pasture werestudied, such as nectar and pollen, with a particularlyemphasis on honeydew. On the basis of phytocoeno-logical analyses and the individual contribution ofmelliferous plants determined according to the inten-

sity of nectar and pollen production, the aim of thiswork was to assess the melliferous potential of plantcommunities, reßecting their importance to bee pas-ture in Tara NP.

Materials and Methods

The Research Area. Tara NP encompasses a largepart of Mount Tara (19,200 ha), which belongs to theDinaric massif and is located in western Serbia. MountTara is situated between 43o 52� and 44o 02� N, and19o15� and 19o38� E, and stands at 1,000Ð1,500 m abovesea level (Fig. 1). It was declared a National Park in1981as it is oneof themostdensely forestedmountainsin Europe, characterized by great biodiversity andnumerous endemo-relict plant and animal species.Forest ecosystems occupy �70% of the total area ofTara NP and are some of the best preserved, highestquality, and most productive in Europe (Gajic et al.1992).

A great ßoristic and phytocoenotic diversity of TaraNP vegetation is attributed to its position in the tran-sition zone between Fagion moesiacum and Fagionillirycum (Horvat et al. 1974). Various bedrocks, a rich

Fig. 1. A map of Serbia with the research area of Tara NP highlighted.

August 2013 JARIC ET AL.: THE MELLIFEROUS POTENTIAL OF PHYTOCOENOSIS ON MOUNT TARA 725

hydrographic network, a favorable altitude, and cli-mate also contribute to this diversity. For the purposesof this research, the 34 plant communities were ana-lyzed (of the 43 noted at Tara NP), including thosewith the greatest signiÞcance for bee pasture.

At Tara NP, the vegetation period in the river val-leys begins in the Þrst decade of April and lasts untilthe end of October, while at an altitude of over 1,000m, it begins 1 mo later and ends a month earlier. Theaverage duration of the vegetation period in the rivervalleys is �190 d, but only 130 d at altitudes of over1,000 m (Gajic et al. 1992).Melliferous Potential of Vegetation. The mellifer-

ous potential of individual plant species is not a rele-vant indicator of their quality unless other factors aretaken into consideration, such as: their frequency incommunities, ßowering phenology, and abiotic andbiotic factors (microclimate, soil, pollinators, and soforth). If they are very frequent, species with lowernectar and pollen production can be of greater im-portance to bees than better quality species that areencountered sporadically.

Melliferous plant species are classiÞed into fourcategories for each of the designated indices (Inp,Ipp), which express their degree of nectar and pollenproduction numerically (1 Ð poor melliferous specieswith minimal nectar or pollen production, 2 Ð good,3 Ð very good, and 4 Ð excellent) (Jasmak 1980; Ric-ciardelli DÕAlbore and Persano 1981; RicciardelliDÕAlbore 1997; Umeljic 1999, 2003).

The coefÞcient of nectar and pollen production(Cnp and Cpp) was calculated for each melliferousspecies by multiplying the mean value of its frequencyin the community n� by its nectar or pollen productionindex (Inp, Ipp).

Cnp(Cpp) � n� Inp(Ipp)

A phytocoenological analysis of the vegetation wasperformed using the Westhoff van der Maarel method(1973), by which the frequency of melliferous speciesin plant communities is determined on the basis of acombined scale of abundance and coverage (rangingfrom 1 to 9). The mean value of the abundance of eachspecies in the community was calculated as the ratioof its total abundance in the community to the numberof releves that deÞne it.

The importance of plant communities on Tara to beepasture was assessed on the basis of their coenotic co-efÞcient of nectar and pollen production (CCnp andCCpp). The CCnp and CCpp were calculated by addinguptheindividualcoefÞcientsofnectar(�Cnp)andpollenproduction (�Cpp) of all the species in the community.

By taking into account both nectar and pollen as themain elements of bee pasture, as well as the numberof melliferous plants related to the total number ofplant species, the coenotic coefÞcient of melliferous-ness (CCm) of a particular plant community, whichtheoretically indicates its melliferous potential, wascalculated as:

CCm � (CCnp � CCpp) � M/P,

where

M Ð the number of melliferous species in the com-munity.

P Ð the total number of species in the community.This formula is an expanded and modiÞed version of

the formula used previously in similar research(Danon et al. 1990).

The production of honeydew as one of the elementsof bee pasture was analyzed for certain species, butthis was not taken into account when calculating themelliferous potential of plant communities, because ofa lack of relevant and reliable data on its availabilityand of numerical values to describe its production.Because of the domination of forest communities, spe-cies producing honeydew constantly or occasionallyand to varying extents, are particularly discussed.

Results and Discussion

Floristic and phytocoenological investigation of thevegetation at Tara NP proved the presence of 188melliferous plants within 25 forest and nine meadowplant communities (Table 1). Considering the per-centage contributions of plant species according toindices of pollen and nectar production (Fig. 2), thedomination of species with a higher pollen productionwas noted for all categories of melliferous plants (Inp/Ipp � 2Ð4), apart from those with a Inp/Ipp of 1. Thepercentage of species with the highest pollen produc-tion index rating (Ipp-4) is almost twice as high asthose with the highest nectar production index rating(Inp-4). The proportion of nectarless species (Inp-0)is almost equal to the number of species producingnegligible quantities of pollen that are unattractive tobees (Ipp-0).

In the beginning of spring, forest bee pasture, whichis characterized by early ßowering herbaceous andwoody plant species, is of the greatest importance forthe early bee colony development. In forest commu-nities, melliferous species are distributed in severalvertical layers (forest ßoor, shrub layer, and treelayer) and they ßower successively throughout thevegetation period. The high percentage of melliferousspecies (�50%) was noted in the following commu-nities: Piceetum-Abietis serpentinicum (63.6%), Pin-etum sylvestris-nigrae serpentinicum (61.1%), Querco-Carpinetum illiricum (59.5%),Quercetum cerris (56.3%),Daphno blagayanae-Quercetum petrae (55.8%), Fag-etum submontanum (53.7%), Querco-Juglandetum(52.9%), and Quercetum montanum serpentinicum(51.6%) (Table 1; Fig. 3).

A comparative analysis of the melliferousness of theforest communities proved that the Querco-Carpin-etum iliricumhas the highest CCpp and CCm, becauseQuercus petraea (Mattuschka) Lieblein, Carpinusbetulus Linnaeus, Corylus avellana Linnaeus, Cornussanguinea Linnaeus, and Acer campestre Linnaeus areprominent in terms of pollen production and abun-dance. The Picetum omorikae-Abietis serpentinicum as-sociation has the highest CCnp because of those plantscharacterized by a high intensity of nectar productionand dominate the forest ßoor (Erica carnea Linnaeus,Thymus pulegioides Linnaeus, andVacciniummyrtillis

726 ENVIRONMENTAL ENTOMOLOGY Vol. 42, no. 4

Linnaeus). The lowest CCnp was noted in the Quer-cetummontanum, because of the low diversity of mel-liferous species and the domination of woody plantsthat do not produce nectar (or just produce it innegligible quantities), but produce pollen or honey-dew (Fagus sylvatica Linnaeus) (Table 1). The asso-ciation Seslerio variae-Ostryetum has the lowest CCm,which can be explained by the low abundance of therecorded melliferous species and the lowest CCpprelative to the other communities.

As the coniferous forest communities mainly com-prise anemophilous woody plants, their valuable con-

tribution to bee pasture is in honeydew production(Fig. 5). For bees, the pollen production of conifers isrelatively low (Pinus sp.; Ipp-1) or even negligible(Picea abies [Linnaeus] Karsten andAbies albaMiller;Ipp-0) relative to deciduous trees (RicciardelliDÕAlbore 1997). However, conifers produce a greaterquantity of better quality honeydew compared withother woody species, which bees prefer (Binazzi 1978,Binazzi and Scheurer 2009). The production of hon-eydew depends directly on the population dynamicsof aphids (Rinchota, Homoptera), and to a great ex-tent on biotic and abiotic factors (Pechhacker 1976,

Table 1. Melliferous potential of investigated forest and meadow plant communities of Tara Mountain

No. Forest communities CCnp CCpp CCm % ms

1 Abieti-Fagetum 15.5 27.2 19.3 45.12 Alno-Piceetum omorikae 31.2 97.3 58.9 45.93 Daphno blagayanae-Quercetum petraeae 16.4 80.9 54.4 55.84 Erico-Pinetum nigrae serpentinicum 18.3 32.9 16.2 33.35 Erico-Pinetum sylvestris serpentinicum 19.9 45.4 26.2 406 Fagetum montanum 16.7 34.9 22.5 43.67 Fagetum submontanum 13.9 26.5 21.7 53.78 Orno-Ostryetum 45.2 59.1 43.9 42.29 Orno-Ostryo-Juglandetum 50.3 38.9 37.9 42.510 Ostryo-Pinetum nigrae 18.8 31.8 17.2 33.911 Parietario-Juglandetum 16.3 34.4 19.8 39.312 Piceetum Abietis montanum 12.4 24.5 18.1 49.113 Piceo-Abieti-Fagetum 16.4 34.7 23.5 46.114 Piceetum-Abietis serpentinicum 25.9 34.8 38.4 63.615 Piceetum omorikae-Abietis serpentinicum 56.4 64.6 51.5 42.616 Piceetum omorikae-Abietis calcicolum 48.4 99.5 61.8 41.817 Pinetum sylvestris-nigrae serpentinicum 39.9 38.9 48.3 61.118 Quercetum cerris 24.6 75.1 56.2 56.319 Quercetum montanum 3.7 40.5 18.3 41.420 Quercetum montanum serpentinicum 28 72.5 52.3 51.621 Querco daleschampii-Ostryetum 44.8 35.9 32.8 39.522 Querco-Carpinetum illiricum 30.9 105.5 81.2 59.523 Querco-Juglandetum 22.2 75.6 51.5 52.924 Seslerio variae-Ostryetum 16 21.2 12.4 33.325 Spiraeo canae-Quercetum petraeae 20.4 74.8 46.4 48.7

Meadow communities

1 Arrhenatheretum elatioris 33.2 33.3 33.2 502 Bromo-Plantaginetum mediae 33.5 40.6 33.6 45.53 Cariceto-Brometum erecti 30.8 36 28.9 43.34 Danthonietum calicinae 18.4 41 23.9 40.55 Eriophoretum latifolii 22 18 10.9 27.36 Lythro-Caricetum paniculatae 6.5 16.5 13.1 57.17 Petasitetum hybridi 18 44 43.4 708 Ranunculo-Nardetum strictae 15.7 19.2 9.9 28.69 Rhinantho-Cynosuretum cristati 56.8 58.8 53.9 46.6

CCnp, coenotic coefÞcient of nectar production; CCpp, coenotic coefÞcient of pollen production; CCm, coenotic coefÞcient of mellifer-ousness; % ms, percentage of melliferous species.

Fig. 2. Percentage contributions of melliferous plants according to intensity of pollen and nectar production in allinvestigated plant communities of Tara NP. (A) Index of pollen production (Ipp � 0Ð4). (B) Index of nectar production(Inp � 0Ð4).

August 2013 JARIC ET AL.: THE MELLIFEROUS POTENTIAL OF PHYTOCOENOSIS ON MOUNT TARA 727

Ricciardelli DÕAlbore and Intopa 2000). In the re-search area, the following species were noted for hon-eydew production: A. alba, A. campestre, Acer pseu-doplatanusLinnaeus, Betula pendulaRoth, F. silvatica,P. abies, P. nigra, P. sylvestris, Populus tremula Lin-naeus,Quercus cerris Linnaeus, Salix caprea Linnaeus,andTiliaplatyphyllosScopoli.According toCarter andMaslen (1982), the most suitable forest honeydews forÔforest honeyÕ in Europe are known to originate mainly

from A. alba, P. abies, Larix decidua Miller, and P.sylvestris, but according to Kloft et al. (1985), otherconiferous plants, attacked by sapÐsucking insects, canalso be a good source of forest honey.

Some of the forests that dominate the Drina Basinin western Serbia and are the best preserved in Europecontain a rare endemic tertiary species, theP. omorika,or Serbian Spruce, which, based on our observations,is a good source of honeydew. This species belongs to

Fig. 3. Comparative distribution of forest plant communities of Tara NP according to their CCnp and CCpp (communitiesare listed in Table 1).

Fig. 4. Comparative distribution of meadow plant communities of Tara NP according to their CCnp and CCpp (com-munities are listed in Table 1).

728 ENVIRONMENTAL ENTOMOLOGY Vol. 42, no. 4

Fig. 5. Melliferous plants of spring and summer bee pasture in forest and meadow communities of Tara Mountain,providing nectar, pollen, and/or honeydew for insect pollinators: (a)Galanthus nivalis, (b)Cornus mas, (c) Salix caprea, (d)Primulavulgaris, (e) Scilla bifolia, (f)Corydalis solida, (g)Acerpseudoplatanus, (h)Coronilla emerus, (i)Taraxacumofficinale,(j) Tilia sp., (k) Trifolium pratense, (l) Lotus corniculatus, (m) Thymus pulegioides, (n) Melilotus albus, (o) Eupatoriumcannabinum, (p) Picea sp. Photo: Marina Macukanovic-Jocic. (Online Þgure in color.)

August 2013 JARIC ET AL.: THE MELLIFEROUS POTENTIAL OF PHYTOCOENOSIS ON MOUNT TARA 729

the Piceetum omorikae-Abietis calcicolum, Piceetumomorikae-Abietis serpentinicum, and Alno-Picetumomorikae communities and, because of its rarity andscientiÞc importance, has been placed under nationalprotection.

The majority of melliferous herbaceous and shrubspecies in coniferous forest communities appear onthe outer edges of the associations. Because of itslong period of ßowering (February to April), abun-dance, and nectar and pollen production (Inp-4;Ipp-4), the most signiÞcant species is E. carnea,which dominates in the early spring. This species isnumerous in the associations: Erico-Pinetum nigraeserpentynicum, Erico-Pinetum silvestris serpentyni-cum, Pinetum silvestris nigrae serpentynicum, Pice-tum-Abietis serpentynicum, Ostreo-Pinetum nigrae,Spireo canae-Quercetum petreae, and Seslerio variae-Ostrietum. Previous melissopalynological researchhas shown that unißoral honey can be obtained fromthis species (Grabeljsek 1996, Ricciardelli DÕAlboreand Intopa 2000).

There are only a small number of early ßoweringspecies that are of importance for early spring beepasture in the forest ßoor of most of the deciduousforests in Tara NP, which are visited by bees searchingfor nectar and pollen before the foliation of woodyplants: Galanthus nivalis Linnaeus, Helleborus odorusLinnaeus,Corydalis solidaLinnaeus, Scilla bifoliaLin-naeus, Primula veris Linnaeus, Primula vulgaris Hud-son, Viola odorata Linnaeus, and Anemone nemorosaLinnaeus (Fig. 5). The signiÞcance of these specieslies not in intensive nectar and pollen production, butin their early appearance and their ßowering at a timewhen there are no other food sources in the surround-ing area. However, pollen production is more pro-nounced, which is important for early spring brooddevelopment.

At the beginning of spring, in the layer of shrubs andlow trees, S. caprea and Prunus species (Prunus avium(Linnaeus) Linnaeus, P. mahaleb Linnaeus) ßower,along withCornusmasLinnaeus,C. avellana, CoronillaemerusLinnaeus,Cotinus coggygriaScopoli, andUlmusglabraHudson. C. mas is a highly melliferous shrub orlow tree (Inp-2; Ipp-4) that appears sporadically(Querco-Juglandetum, Parietario-Juglandetum) andoffers bees the Þrst, precious source of nectar andpollen, very early in February or March (Fig. 5). Asthe hazel (C. avellana) is not characterized by nectarproduction, its ßowers are pollinated by wind, butbees visit them because of the high amount of pollen(Inp-0; Ipp-4) that they contain as early as February.Although its abundance is not high, it is present in asigniÞcant number of forest plant communities: Fag-etum montanum, Daphno blagayanae-Quercetum pe-trae, Spiraeo canae-Quercetum petrae, Querco-Carpin-etum illiricum, Picetum omorikae-Abietis calcicolum,and Piceo-Abieti Fagetum. Considering the ßoweringperiod of the hazel lasts for just over a week (Februaryor March), the ßowers often die quickly because ofsudden frosts and the bees do not manage to make useof it.

The important early spring melliferous species nextto rivers are the black poplar (P. tremula) and goatwillow (S. caprea). P. tremula, which grows sponta-neously in wet grounds, ßowers at the beginning ofspring, before or at the same time as foliation, and isvisited by bees gathering pollen and honeydew, be-cause there is no nectar in its unisexual ßowers (Inp-0;Ipp-3). It appears only sporadically in Fagetum sub-montanum, but has a signiÞcant frequency in the Pice-tum omorikae association in the layer of shrubs andtrees. While the poplar is an anemophilous species, thepollination of S. caprea (Fig. 5) occurs mostly withthe help of insects and to a somewhat lesser extent thewind and according to Tamura and Kudo (2000), ex-istence of two pollinating systems in willow speciescould be advantageous under certain conditions. Aspointed out elsewhere (Ricciardelli DÕAlbore 1997),the goat willow, more common along watercourses, isvisited by many Apoidea for the gathering of pollen inhigh percentages, nectar and frequently some honey-dew (Inp-4; Ipp-4).

In the tree layer, Juglans regia Linnaeus, a sponta-neously growing Tertiary relict that ßowers in spring,provides relatively large quantities of pollen (Inp-0;Ipp-2). According to Kloft and Kunkel (1985), it pro-vides only small amount of honeydew, which is notenough to attract bees. Similarly, Q. cerris, forminglarge woody habitats, is of importance for pollen col-lection (Inp-0; Ipp-4), but also for honeydew obtainedperiodically in good quantities. Spring-ßowering treesthat are very frequent and are characterized by highnectar and pollen production includeAcer species (A.campestre, A. platanoides, andA. pseudoplatanus). Ma-ple trees are frequently visited by bees searching forpollen (Ipp-3) and nectar (Inp-2) and occasionallyhoneydew (A. campestre, A. pseudoplatanus) (Ric-ciardelli DÕAlbore and Intoppa 2000). In line with thephenological calendar, the linden (Tilia sp.) ßowers atthe beginning of summer (Inp-4; Ipp-2). In Tara NP,it is found in theQuerco-Juglandetum, Picetum omori-kae-Abietis calcicolum, andPiceetumAbietismontanumassociations. There are three autochthonous, apicul-turally signiÞcant species of linden, which can befound in this region of Serbia. The large-leaved linden(T. plathyphyllos) is the Þrst to ßower, while the small-leaved linden (Tilia cordataMiller) ßowers at the endof June and beginning of July, and the silver linden(Tilia tomentosaMoench) ßowers during the Þrst 10 dof July.

Herbaceous plants of signiÞcance for summer pas-ture, found on the edges of the investigated forestcommunities are: Ajuga reptans Linnaeus, Calaminthaofficinalis Moench, T. pulegioides, Campanula rapun-culoides Linnaeus, Campanula persicifolia Linnaeus,Campanula trachelium Linnaeus, Calamintha clinopo-dium Bentham, Salvia glutinosa Linnaeus, SymphytumtuberosumLinnaeus, Stachys rectaLinnaeus, Trifoliumalpestre Linnaeus, Trifolium repens Linnaeus, Trifo-lium pratense Linnaeus, Trifolium montanum Lin-naeus, Trifolium campestre Schreber, Pulmonaria offi-cinalis Linnaeus, Lotus corniculatus Linnaeus andGentiana asclepiadea Linnaeus.

730 ENVIRONMENTAL ENTOMOLOGY Vol. 42, no. 4

The meadow communities are of signiÞcance at theend of spring and during the summer when the de-ciduous forests are for the most part in leaf, while atthe end of the summer and beginning of autumn thecontribution of forest bee pasture once again becomesrelevant because of honeydew. Thanks to the ßoristicdiversity and the successive ßowering of good mellif-erous plants, bees can use the meadow pasture forlonger, during the vegetation period. In meadow veg-etation of Tara NP, the greatest contribution to beepasture (in terms of abundance and pollen and nectarproduction index ratings) is made by species from theFabaceae and Lamiaceae, whose populations are mostnumerous. It is clear that species producing high quan-tities of pollen dominate, as is conÞrmed by the CCppvalues for individual (Fig. 4) communities.

The meadow communities with the highest CCmare Rhinantho-Cynosuretum cristati and Petasitetumhybridi, although the reasons for this differ (Table 1).The greatest contribution to the melliferous potentialof the Rhinantho-Cynosuretum cristati is made by spe-cies, which account for only half of the total numberof species that characterize it (46.6%), but which areprominent in terms of both abundance, and high nec-tar and pollen production index ratings, such as:Carum carvi Linnaeus (Inp-2; Ipp-4), Centaurea jaceaLinnaeus (Inp-3; Ipp-3),L. corniculatus(Inp-4; Ipp-2),T. pulegioides (Inp-4; Ipp-2), Vicia cracca Linnaeus(Inp-4; Ipp-3),T. repens (Inp-4; Ipp-4), andT. pratense(Inp-4; Ipp-4). As opposed to Cynosurus cristatus Lin-naeus (fam. Poaceae), which is most abundant, but asan anemophilous species is insigniÞcant for bee pas-ture, the second ediÞcator Rhinanthus rumelicus Lin-naeus (Inp-2; Ipp-2) is abundant melittophilous spe-cies, recorded in all the phytocoenological releves.Among the other anemophilous species that can be asource of pollen in certain circumstances, there arePlantago media Linnaeus and Plantago lanceolata Lin-naeus, which appear in relatively small numbers in theRhinantho-Cynosuretum cristati. The Petasitetum hy-bridi had the lowest total number of species, but theproportion of melliferous plants was high (70%),which contributed to a high CCm and gave a distortedpicture of melliferous signiÞcance of this communitycompared with other ones in the research area. Re-gardless of the high value of CCm calculated, thiscommunity is of no great importance for bee pastureas it is fragmentary, covering small areas and apartfrom the ediÞcator Petasytes hybridus (Linnaeus)Gaertner, Meyer and Scherbius, no single species oc-curs in high abundance.Ranunculo-Nardetum strictae had the lowest co-

enotic coefÞcient of melliferousness (CCm-9.9) as itcontained only 26.8% melliferous species. However,the lowest coenotic coefÞcient of both nectar andpollen production (CCnp-6.5; CCpp-16.5) was notedin the Lythro-Caricetum paniculatae, where 57.1% ofthe recorded species were melliferous ones (Table 1).However, these were mainly present only sporadicallyand were poor nectar and pollen producers, withLyth-rum salicaria Linnaeus (Inp-2; Ipp-4) and Ranunculus

repens Linnaeus (Inp-1; Ipp-3) making the greatestcontribution to the melliferousness of this community.

Because of geographical and environmental char-acteristics of the research area, as well as melliferouspotential of vegetation, beekeeping within the TaraNP is in the phase of expansive development. Con-sidering the isolation of the mountain from all kinds ofpollution, in the last 10 yr, more attention is paid toorganic beekeeping. According to the Beekeepers As-sociation “Spruce,” around 400 hives are placed, andthe average honey yield per hive is 8Ð30 kg, dependingon altitude and weather conditions. Annual honeyproduction averages �7Ð8 tons, and it is believed thatit could be 10 times higher, indicating underutilizationof melliferous potential of the vegetation. Because, thestudy area is characterized by forest-dominated veg-etation, it is evident that honeydew is produced everyyear, reaching maximum in dry years, when the au-tumn is warm and without precipitation, and the av-erage summer temperature ranges between 35Ð40C.Meadow bee pasture is more used by honey bees inyears with increased rainfalls during the summer.

The importance of the woody species recorded inTara NP for bee pasture (C. mas, C. sanguinea, C.avellana, Tilia sp., Acer sp., J. regia) has been substan-tiated in other parts of Serbia and the Balkans bymeans of Cnp levels (Perisic et al. 2004, Grozeva 2011,Grabeljsek 1996). The results of earlier researchesconcerning apißora of meadow communities inthree regions (Suva Planina, Kragujevac, and VelikaMorava) showed that the association Agropireto-Festucetum pratensis in the Kragujevac area containedthe most melliferous plants (58.4%), while there wereleast in the Cariceto-Dryadetum octopetale (10.3%) onSuva Planina (Macukanovic et al. 1996, Danon et al.,1990). Furthermore, previous research on SuvaPlanina, done by Danon et al. (1990), revealed thehighest CCnp in the Trifolieto-Agrostidetum vulgaris(CCnp-69.0), and the lowest in the Cariceto-Dryade-tum octopetale (CCnp-1.62).

In comparison to the results of previous research,which was based solely on nectar production and theproportion of melliferous species (Blazencic et al.1994, Macukanovic et al. 1996, Macukanovic andGrabeljsek 1996), this research includes also pollenproduction, as well as a review of the plants thatproduce honeydew, which gives an altogether truerreßection of the melliferous potential of the speciesand communities of this mountain region. However, itis quite unlikely that the melliferous potential of anassociation can be equated to beesÕ actual utilizationof it, which largely depends on the surface area thecommunity covers, climatic and orographic factors,the distance from bee colonies, and so forth. Evenwhen a community is composed entirely of mellifer-ous species contributing considerably to a high CCm,it does not mean that it is important to bees, if theabundance, and nectar and pollen production of thosespecies are low or the community is fragmentary,extending over small areas. Finally, to gain a completeinsight into the melliferous potential of apißora and toestablish the quality of certain melliferous species and

August 2013 JARIC ET AL.: THE MELLIFEROUS POTENTIAL OF PHYTOCOENOSIS ON MOUNT TARA 731

their attractiveness to bees in competing bee pastureduring thevegetationperiod, it isnecessary toconductmelissopalynological analyses of honey originatingfrom this region. Monitoring the ßowering phenologyparallel with determining the spectrum of pollen typesin honey samples in terms of relative frequencies,would reveal the dominant and most attractive beeforage plants and the most important honey sourcesduring the vegetation season. Therefore, this articlepresents preliminary theoretical investigation of themelliferous potential of apißora and vegetation of TaraNP, which will be continued by further melissopa-lynological study aiming to determine botanical originof honey and the signiÞcance of certain plants for beepasture in this area.

Acknowledgments

This work was supported by the Ministry of Education,Science and Technological Development of Serbia, grants173018 and 46009.

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Received 30 January 2013; accepted 22 May 2013.

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