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IAWA Bulletin N.S., Vol. 8 (3),1987 245 ECOLOGICAL TRENDS in THE IAWA Bulletin n.s., Vol. 8 (3),1987 245 ECOLOGICAL TRENDS IN THE WOOD ANATOMY OF TREES, SHRUBS AND CLIMBERS FROM EUROPE by Pieter Baas and Fritz H. Schweingruber Rijksherbarium, P.O. Box 9514, 2300 RA Leiden, The Netherlands and Eidgenössische Anstalt für das forstliche Versuchswesen, CH-8903 Birmensdorf, Switzerland Summary Ecological trends for occurrence of certain established on a floristic basis. It is suggested vessel, tracheid and fibre characteristics. have that the ecological trends are the result of se­ been analysed for 505 species (belonging to lection for specialised wood anatomical fea­ 221 genera and 71 families) from Europe, Cy­ tures such as simple perforation plates, vascular prus, and Madeira. Macroclimatic gradients tracheids, and different vessel size classes in the from boreal, via temperate to mediterranean mediterranean region and dry and warm sites in are strongly related with a decreasing incidence general, and that primitive features such as scala­ of scalariform perforations, (almost) exclusively riform perforations, solitary vessels, and fibre­ solitary vessels, and fibre-tracheids (i. e., fibres tracheids are not selected against in cool boreal with distinctly bordered pits). In this sequence and mesic habitats. For some characters macro­ the incidence of different vessel size classes climatic factors (chiefly temperature) seem more (vessel dimorphism) and vascular tracheids in­ important than moisture availability, for others creases. Ring-porous tendencies and spiral ves­ moisture availability seems equally important. sei thickenings have their peaks in the temper­ The biological significance of the reported ate zone. The subtropical flora of Madeira trends is discussed in terms of safety and effi­ shows low values for the percentage of species ciency of xylem sap transport, and of evolution­ with any of the above attributes. ary correlative constraints. In the graded series from dry (including wet Key words: Vessel perforations, vessel distribu­ saline), via normal to mesic, the incidence of tion, tracheids, fibres, ecology, Europe. scalariform perforations strongly increases and that of vascular tracheids decreases. Other at­ Introduction tributes show a weaker or ambiguous relation­ In the final stages of the preparation of an ship with moisture availability: incidence of extensive atlas of the wood anatomy of trees, different vessel size classes and spiral vessel shrubs, and climbers from Europe, Cyprus, and thickenings decreases; incidence of fibre-tra­ Madeira (Schweingruber, in preparation) de­ cheids increase in dwarf shrubs and normal scribing and picturing c. 600 species of woody shrubs, but decreases in trees; ring-porosity and gymnosperms and angiosperms, an analysis of (alm ost) exclusively solitary vessels are inde­ ecological trends in the wood anatomy of the pendent of moisture availability. 505 species of native dicotyledonous trees, There is also a relationship of some of these shrubs, and climbers described has been carried wood anatomical attributes with plant habit. out to be incorporated as a special chapter The incidence of species with exclusively scala­ (Baas, in Schweingruber, in preparation). These riform perforations, ring-porosity, and (alm ost) species represent 221 genera and 71 families. exclusively solitary vessels, decreases gradually The results of this analysis seemed to justify a in the series from trees via shrubs to dwarf­ precursory publication in this Bulletin. shrubs, while the reverse is true for species with The analysis of wood anatomical trends in different vessel size classes, and vascular tra­ the ecologically and taxonomically diverse cheids. Fibre-tracheids show no clear relation­ European flora is a complement to recent anal­ ship with plant habit. Climbers in the European yses of local or regional floras from Israel (Baas woody flora score relatively low for incidence et al., 1983; Fahn et al., 1986; Baas &Carlquist, of scalariform perforations and solitary vessels, 1985), southern California (Carlquist & Hoek­ but high for ring-porous tendencies, different man, 1985), and Australia (Carlquist, 1977). vessel size classes, and spiral thickenings. Inci­ The present analysis is based on many more dence of fibre-tracheids and vascular tracheids species than these earlier studies, but is limited is more or less similar to that in erect shrubs. in its scope to qualitative characters of per­ Within ecologically diverse genera or families forate and imperforate tracheary elements. very little can be traced of these salient trends (text continued on page 259) Downloaded from Brill.com10/09/2021 04:50:05AM via free access 246 IAWA Bulletin n.s., Val. 8 (3),1987 Table 1. Species studied and their ecological and wood anatomical attributes. Legend: Macroclimate: b = boreal; t = temperate; m = mediterranean; s = subtropical oceanic (i.e., from Madeira); * = montane (j.e., highest localities at or above the timberline). See text for the classification of species with altitudinal or latitudinal ranges overlapping several categories. Moisture availability: d = dry; dh = saline (i.e., physiologically dry); n = normal; m = mesic. See text for classification of species covering several ecological categories. Habit: ds = dwarf shrub; s = shrub; t = tree; cl = climber, liana or scrambling shrub; ** = stern parasite. See text for classification of species with intermediate habit or covering more than one habit category. + = character present; ± = character intermediate or poorly developed (in case of perforation plates: mixed simple and scalariform perforations present; in case of ring-porosity: wood semi-ring-porous or semi-ring- to diffuse-porous); +/- = fibre-tracheids and libriform fibres present; - = character absent. -0 >, ~ '" & .... .~ '(j) -0'" os '" Ol '(j) -5 ~ Z' ~ "i ~ os Q) .; '" 0 '" -0'" -5 .... +' o '" '" Q) '0) os +' os Ei>, ~ Ei ~ t> ~ '" bD ..<:: .b .... 0 'U) ca Q) ~ " Q) o·~ '"Q).~ I::: "~ E <.:;+' o '-"> ~C/) E .-4 I::: .... .~ os o-... _rn •U'l ,...Q.) Q)cn > a c "o E .... .... ....... Q) ~ '3 Q) .... ~ Q) '" ;<;:l ~~ 0..Q)::l~", os .~ Species "os '0 os .... • .....-4 ......" ,,0..... " ~ " Q) ~ ~] ~~ '"os '"os Ei Ei ..<:: "'0.. .~ > (1) '"C1 ~-:5 t;: > > Aceraceae Acer campestre t d t + Acer granatense m* n + Acer heldreichii m n t + Acer hycranum m n t + Acer monspessulanum m n t + Acer obtusatum m n t + Acer opalus t n t + Acer platanoides m t + Acer pseudoplatanus m + Acer sempervirens m n + Acer tataricum m n t ± ± + Amaranthaceae Bosea cyprea m m cl Anacardiaceae Cotinus coggygria m d s + + + + Pistacia atlantica m d + + + + Pistacia lentiscus m d t ± + + + Pistacia terebinthus m d t + + + + Rhus coriaria m d s + + + + Apocynaceae Nerium oleander m m ± Aquifoliaceae Ilex aquifolium m + + + Araliaceae Hedera helix t n cl ± ± ± + Asc1epiadaceae Periploca graeca m n cl + + ± Periploca laevigata m d s ± ± ± ± Downloaded from Brill.com10/09/2021 04:50:05AM via free access IAWA Bulletin nos"~ Vol. 8 (3),1987 247 (Table 1 continued) VI "d & ~ ,~ VI 'S os VI "d 'S -5 ~ ~~ ~ ; VI os "' 0 VI '~ o VI VI "d -5 .b ~ S VI ßS>.Q) Q) 'S ..... r;- > VI b.O ..s:: <.> ... § VI I':: <.> 1: ''; Q) O'~ ..., E ~ ~ (1)..... ~ ..., ""...,,~ --00· .... <.> os ~ I':: VI > I':: ... ta o ... ... Q) Q) ...... Q) ~ E Q) ... VI ~ t; VI ~ & ~ os ....: Q) '3 <.> b.O 'fjJ Species os '0 .;~ :"ü ~gj '1""4........ " ..0... VI ~ <.> Q) I':: Q) >< "os S S ..s:: VlI'l. 'i: > Q) :.s'ü ~.s t-+= > ~ Berberidaceae Berberis aetnensis m* n s + + + + Berberis cretica m d s + + + + Berberis hispanica m m s + + + + Berberis vulgaris t d s + + + + Betulaceae Alnus cordata m m + Alnus glutinosa t m + ± ± Alnus incana t m t + ± ± Alnus orientalis m* m t + ± ± Alnus viridis t* m s + Betula etnensis m* n + Betula humilis b m ds + Betula nana b m ds + Betula pendula b n t + Betula pubescens b m t + Boraginaceae Echium angustifolium m n ds Echium nervosum s d s Lithodora fruticosa m d ds ± + + + Lithodora hispidula m d ds ± + + ± Moltkia petraea m d ds ± + + + Onosma erecta m d ds ± + ± Onosma fruticosa m d ds + + + Onosma graeca m d ds ± + ± Buxaceae Buxus balearica s d t + + + Buxus sempervirens t d t + + + Capparidaceae Capparis spinosa m d ds + ± Caprnoliaceae Lonicera alpigena n s ± ± ± ± + + Lonicera caerulea t* m s ± ± ± ± + + Lonicera caprifolium t m cl ± ± ± ± + + Lonicera etrusca m n cl ± ± ± ± + + Lonicera implexa m n cl ± ± ± ± + + Lonicera nigra t n s ± ± ± ± + + Lonicera periclymenum m cl ± + + Lonicera pyrenaica m* d ds ± ± ± ± + + Lonicera xylosteum t n s ± ± ± ± + + Sambucus maderensis s m Sambucus nigra m s ± ± ± Sambucus racemosa t m s ± ± ± Viburnum lantana t d cl + + + + Viburnum opulus n s + + + + Downloaded from Brill.com10/09/2021 04:50:05AM via free access 248 IAWA Bulletin n.s., Vol. 8 (3),1987 (Table 1 continued) Vl "CI ~ ~ ~ Vl .Ci) :Ei ",.~ - "CI ,.c: 'Z' ~.: ca .Ci) <> .0; 00'0 ~ ~ Vl ,.c: ...,Q) '" o rn rn _ "CI <> ...,... > .Ci) '" e ;g E .5;... Q) ~b/) ,.c: ...,... <> ... 0 Vl ~- > rnc:;:: <> '" ·C '" Q) o·~ 0 ..., ] '" ... ..., ... -- ~...:. Q,).~ ... 0 00· .... c:: eil > c:: ... a .: <> ..., Q) 0 ....~ '"... ~ '" E ..., d, '"5 <> ... Vl .~ <> b/) ~ ~ ~ ~ ca~ <> Species ·0 .!~ tfJ 'ü if: ~ .~.~ ... Vl Vl .: Q) ~ .~_ ~,.c: .J:J ~ '" Q) e'" e ,.c: ~~ ·C > (1) ""C:j u rn~ t+= '"> '"> (Caprifoliaceae) Viburnum tinus m n s + + + + Caryophyllaceae Arenaria ciliata b* n ds + Dianthus hispanicus m d ds ± ± + Dianthus rupicola m d ds ± ± + Silene pseudovelutina m n ds + Celastraceae Euonymus europaeus t m s ± + ± + +/- Euonymus latifolius t m s ± + ± + +/- Euonymus
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