Last Remnants of Riparian Wooded Meadows along the Middle Drava River (Slovenia): Species Composition Is a Response to Light Conditions and Management

Abstract

In the present study, we investigated the species richness and species composition in relation to light and management regime in remnants of wooded meadows within the riparian forests along the middle Drava River in Slovenia. 41 plots of still managed and, at different time periods (<5 yrs, 5–15 yrs, >15 yrs), abandoned riparian wooded meadows (RWM) were sampled. In addition to vegetation relevés, light intensity (PAR) was also measured in plots. Within the still-managed RWM, two floristically distinct types were recognized using TWINSPAN analysis: meadow-like and forest-like. Light intensity differed significantly between types. The CCA of active RWM showed a significant relation between species composition and light conditions. The number of species per relevé on active RWM was negatively correlated with light intensity – in contrast to North European wooded grasslands. This could be explained by the influence of species-rich riparian hornbeam forests that contribute many understorey species, in contrast to naturally mesotrophic meadows. CCA of both active and abandoned RWM demonstrated that light was a good predictor of RWM species composition and that abandonment caused profound changes in floristic composition. The species turnover during succession was more pronounced in less shaded meadow-like RWM where more light-requiring (grassland) species occurred. Species richness was the highest in active forest-like RWM. There were no significant differences in species richness between active meadow-like RWM and groups of abandoned RWM. The remains of riparian forests stretching along the flood plains of Central and Eastern Europe are considered one of the most natural ecosystems in the prevailing agricultural landscape. But naturalness could be in many cases only the consequence of abandonment of ancient land-use practices, like grazing of livestock, cutting between the trees, litter collecting, etc. Examination of the middle Drava River in Slovenia suggests the need to recognize the remains of ancient cultural landscape.

Appendix

A list of 129 vascular plant species recorded in the herb layer of the 41 relevés of riparian wooded meadows (RWM) along the Drava River in Slovenia

		Active		Abandoned
		G1	G2	<5 yr	5–15 yr	>15 yr
Grassland communities
Valeriana officinalis	Val off	43	.	.	.	.
Serratula tinctoria	Ser tin	14	.	.	13	.
Thalictrum flavum	Tha fla	29	.	13	13	.
Arabidopsis thaliana	Ara tha	71	9	25	63	43
Angelica sylvestris	Ang syl	57	9	13	13	29
Centaura carniolica	Cen car	71	36	50	38	14
Ajuga reptans	Aju rep	86	73	63	25	43
Galium mollugo	Gal mol	71	82	75	63	43
Cirsium oleraceum	Cir ole	57	27	38	25	14
Colchicum autumnale	Col aut	57	55	88	38	57
Heracleum sphondylium	Her sph	57	18	38	25	29
Brachypodium pinnatum	Bra pin	43	91	63	63	71
Pimpinella major	Pim maj	14	73	25	50	29
Anthriscus sylvestris	Ant syl	29	18	.	.	.
Filipendula ulmaria	Fil ulm	29	9	50	38	14
Vicia cracca	Vic cra	29	27	25	13	29
Orchis militaris	Orc mil	29	45	25	.	25
Plantago lanceolata	Pla lan	29	55	13	13	.
Crepis biennis	Cre bie	14	9	13	.	.
Euphorbia cyparissias	Eup cyp	14	27	25	13	14
Prunella vulgaris	Pru vul	14	45	25	.	.
Ranunculus acris	Ran acr	14	91	50	.	.
Trifolium pratense	Tri pra	14	64	.	.	.
Arrhenatherum elatior	Arr ela	.	100	13	.	.
Achillea millefolium	Ach mil	.	73	38	.	.
Briza media	Bri med	.	55	.	.	.
Trifolium repens	Tri rep	.	55	.	.	.
Centaurea jacea	Cen jac	.	45	.	.	.
Plantago media	Pla med	.	45	13	.	.
Bromopsis erecta	Bro ere	.	36	.	.	.
Lathyrus pratensis	Lat pra	.	36	.	.	.
Pimpinella saxifraga	Pim sax	.	27	.	.	.
Trifolium montanum	Tri mon	.	27	.	.	.
Leucanthemum ircutianum	Leu irc	.	18	.	.	.
Linum catharcticum	Lin cat	.	18	.	.	.
Lotus corniculatus	Lot cor	.	18		.	14
Salvia pratensis	Sal pra	.	18	.	.	.
Agrimonia eupatoria	Agr eup	.	9	13	.	.
Knautia arvensis	Kna arv	.	9	.	.	.
Sanguisorba officinalis	San off	.	9	.	.	.
Woodland and shrubland communities
Humulus lupulus	Hum lup	14	.	.	13	29
Euphorbia epithymoides	Eup epi	29	.	.	.	.
Polygonatum latifolium	Pol lat	43	.	13	.	.
Festuca gigantea	Fes gig	43	.	.	13	14
Geum urbanum	Geu urb	43	.	.	25	29
Paris quadrifolia	Par qua	43	.	13	13	29
Cerastium sylvaticum	Cer syl	71	.	13	13	.
Galeobdolon montanum	Gal mon	71	.	63	13	.
Stachys sylvatica	Sta syl	71	.	13	25	29
Anemone ranunculoides	Ane ran	71	18	13	13	.
Asarum europaeum	Asa eur	71	18	75	75	71
Viola riviniana	Vio riv	71	9	.	13	.
Brachypodium sylvaticum	Bra syl	100	18	75	63	100
Galanthus nivalis	Gal niv	100	18	13	38	14
Pulmonaria officinalis	Pul off	100	18	38	63	57
Salvia glutinosa	Sal glu	86	45	75	75	71
Knautia drymea	Kna dry	71	55	50	25	29
Clematis vitalba	Cle vit	71	45	13	63	43
Cruciata glabra	Cru gla	57	18	75	25	29
Dryopteris filix-mas	Dry fil	43	9	13	13	.
Lunaria rediviva	Lun red	43	36	.	25	.
Symphytum tuberosum	Sym tub	43	27	25	13	29
Gagea lutea	Gag lut	29	27	13	.	29
Fragaria vesca	Fra ves	14	9	38	13	14
Anemone trifolia	Ane tri	14	9	.	13	.
Cyclamen purpurascens	Cyc pur	14	18	13	13	.
Listera ovata	Lis ova	14	9	13	.	.
Primula vulgaris	Pri vul	29	82	63	50	14
Leontodon incanus	Leo inc	.	100	13	.	.
Viola hirta	Vio hir	.	64	25	13	43
Geranium phaeum	Ger pha	.	36	.	.
Clinopodium vulgare	Cli vul	.	18	13	.	14
Peucedanum oreoselinum	Peu ore	.	9	13	.	.
Carex alba	Car alb	.	9	13	25	14
Corydalis cava	Cor cav	.	9	13	25	43
Rubus fruticosus	Rub fru	.	9	25	38	43
Vinca minor	Vin min	.	9	.	13	14
Euonymus europaea juv.	Euo eur	.	.	38	75	71
Ligustrum vulgare juv.	Lig vul	.	.	25	25	43
Crataegus monogyna juv.	Cra mon	.	.	.	50	29
Ulmus laevis juv.	Ulm lae	.	.	13	13	29
Fraxinus excelsior juv.	Fra exc	.	.	.	13	43
Prunus padus juv.	Pru pad	.	.	.	13	29
Quercus robur juv.	Que rob	.	.	.	25	14
Acer campestre juv.	Ace cam	.	.	.	13	14
Rhamnus catharticus juv.	Rha cat	.	.	13	.	14
Frangula alnus juv.	Fra aln	.	.	.	.	29
Populus alba juv.	Pop alb	.	.	25	.	.
Lythrum salicaria	Lyt sal	.	.	.	13	.
Cornus sanguinea juv.	Cor san	.	.	25	13	29
Disturbed habitats
Impatiens glandulifera	Imp gla	14	.	25	.	14
Eupatorium cannabinum	Eup can	29	.	13	13	.
Stelaria montana	Ste mon	100	.	50	50	14
Erigeron anuus	Eri anu	71	9	75	25	14
Cardamine impatiens	Car imp	71	.	25	13	.
Alliaria petiolata	All pet	86	18	.	25	43
Solidago gigantea	Sol gig	100	45	75	100	86
Aegopodium podagraria	Aeg pod	86	100	88	50	100
Cruciata laevipes	Cru lae	43	18	13	13	14
Rudbeckia laciniata	Rud lac	57	18	63	63	14
Oxalis corniculata	Oxa cor	43	9	.	.	.
Carex hirta	Car hir	29	18	.	13	.
Glechoma hederacea	Gle hed	23	73	38	13	14
Cardamine hirsuta	Car hir	.	18	.	.	.
Lamium maculatum	Lam mac	.	9	.	13	.
Pastinaca sativa	Pas sat	.	9	.	.	.
Urtica dioica	Urt dio	.	.	38	50	.
Indifferent habitat preference
Oxalis acetosella	Oxa ace	14	.	13	.	14
Lysimachia vulgaris	Lys vul	43	.	25	13	100
Melica nutans	Mel nut	71	.	75	38	57
Rubus caesius	Rub cae	86	.	25	38	43
Dactylis glomerata	Dac glom	86	82	50	13	.
Anemone nemorosa	Ane nem	71	64	38	13	29
Taraxacum officinale	Tar off	71	27	13	.	.
Ficaria verna	Fic ver	43	18	13	.	14
Galeopsis tetrahit	Gal tet	43	9	25	13	43
Hedera helix	Hed hel	43	27	.	25	57
Lysimachia nummularia	Lys num	43	27	.	13	14
Symphytum officinale	Sym off	43	9	.	.	14
Vicia sepium	Vic sep	14	9	13	.	.
Cardaminopsis arenosa	Car are	.	27	.	.	29
Ranunculus repens	Ran rep	.	36	.	.	.
Solanum dulcamara	Sol dul	.	18	.	13	.
Sambucus nigra juv.	Sam nig	.	.	13	.	14
Robinia pseudacacia juv.	Rob pse	.	.	13	25	29
Deschampsia cespitosa	Des cae	.	.	38	13	.
Calamagrosits arundinacea	Cal aru	.	.	.	25	.
Campanula persicifolia	Cam per	.	.	25	.	.
Dactlylorhiza maculata	Dac mac	.	.	13	13	.

1. G1 – Twinspan cluster Nr. 1; G2 – Twinspan cluster Nr. 2. Species are grouped according to habitat preferences by Ellenberg et al. (1992).
2. Species values are percentage frequencies.