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Atlas of Stem Anatomy in Herbs, Shrubs and Trees

Volume 1

Bearbeitet von Fritz H. Schweingruber, Annett Börner, Ernst-Detlef Schulze

1. Auflage 2011. Buch. vIII, 495 S. Hardcover ISBN 978 3 642 11637 7 Format (B x L): 21 x 29,7 cm Gewicht: 1554 g

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Number of , worldwide and in Europe Analyzed species: Aizoon canariense L. The Aizoaceae family includes 127 genera with 2500 species. cordifolia (L. fil.) N.E. Br. Most of the species grow in the tropics of the southern hemi- acinaciformis (L.) L. sphere. Mesembryanthemum crystallinum L. In Europe, there are 7 genera with 10 species. Only 2 species of Mesembryanthemum nodiflorum L. the geneus Mesembryanthemum are endemic. All other taxa are naturalized in Europe and the Canary Islands.

Analyzed material Aizoaceae

The xylem and phloem of 5 Aizoaceae species are analyzed here. Life forms analyzed: Studies from other authors: Semi-woody chamaephytes 2 18 genera Hemicryptophytes and geophytes 1 Therophytes 2 analyzed from different vegetation zones: Studies from other authors: Mediterranean 18 genera Arid 3 Subtropical 2 All species analyzed grow near the seashore in the Mediterra- nean and in subtropical climates. Mesembryanthemum crystallinum

Carpobrotus edulis Aptenia cordifolia Aizoaceae 36 verse section. Canary Islands.Canary zone, subtropical climate. Gran Canaria, site,thermophile succulent leaves, ruderal cm-high plantwith rays. Root collarofa 15 parenchymatic cells,connectedwithlarge ular bandsofxylem,phloemandadjacent Fig. 1. sue) ( zones are separatedby parenchymatic zones (conjunctive tis- sive cambia. Themore-or-less circular arrangedxylem/phloem Annual ringsare absent.Characteristicisthepresence ofsucces- Characteristics ofthexylem small andround ( cristallinum Islands.ife, Canary mophile zone, subtropical climate, Tener- site,ther- cm-high ,ruderal culent, 10 unlignified, 1-3-seriate. Root collarofsuc- parenchyma withlargecells.Raysmostly with small,radiallyorientedcells,adjacent mation process inmidFebruary. Phloem gential bandsofxylemandphloem,infor- Fig. 4. 500 250 Figs. 1-4 μ Stem withsuccessive cambia.Irreg- Stem withsuccessive cambia. Tan- μ m m , transverse section. ). Perforations are simpleandinter-vessel pitsare Aizoon canariense Fig. 5 Mesembryanthemum rv ). Fibres are thin-tothick-walled.The , trans- v r ca ph ph ct ct xy ca xy lands. tropical climate, Gran Is- Canaria,Canary site,thermophilezone,leaves, sub- ruderal a 15 cm-high annualplantwithsucculent small round inter-vessel pits.Root collarof Fig. 5. verse section. Islands.Canary phile zone, subtropical climate,Gomera, ing plantwithsucculentleaves, thermo- Rays are absent.Root collarofalonghang- jacent parenchymatic conjunctive tissue. gential bandsofxylem,phloemandad- Fig. 2. 500 50 μ μ Aizoon canariense Vessels and withsimpleperforations Stem withsuccessive cambia. Tan- m m Aptenia cordifolia absent in idioblasts intheinter-vascular bundlezone ( lignified ( absent ( be fairlylargewithoftenirregularly formedcells,1-3-seriateor axial parenchyma isabsentorparatracheal( , radialsection. p Figs. 2 Fig. 1 Aizoon canariensis , trans- , 4 ). Short rhaphides (20 μm) are bundledinafew ). Short rhaphides(20 and vrp f v ph ct xy Aizoon canariense cal climate,Gran Islands. Canaria,Canary site,thermophilezone, subtropi-ruderal high annualplantwithsucculentleaves, form isvariable. Root collar ofa15 cm- Fig. 6. acinaciformis Portugal.ranean zone, Algarve, culent leaves, garden onthecoast,Mediter- a several-meter-long chamaephytewithsuc- ioblasts withbundlesofrhaphides.Stem of chyma bands.Grey filledcells represent id- regular bandsofxylem,phloemandparen- Fig. 3. 6 100 ). Raycellwallsare mostlythinandun- 250 . μ Rays3to6-seriate.Cellsize and μ m Stem withsuccessive cambia.Ir- m , transverse section. cry , tangentialsection. r Fig. 3 Fig. 4 ). Crystals are ). Crystals r f ). Rayscan Carpobrotus

ph ct xy 37

Characteristics of the phloem

Outside the peripheral circle of vascular bundles is a parenchy- matic zone, which is delimitated by the lateral meristem and the phellem (Fig. 7-9). Many idioblasts with rhaphides are in both Mesembryanthemum species (Fig. 9).

UDSKLG phe

GHDG co Aizoaceae phg

co LGLREODVW

co ph

ct ph ca 100 μm 100 μm 100 μm xy Fig. 7. Bark with a small phloem with small Fig. 8. Bark with a small phloem with small Fig. 9. Bundles of rhaphides in idioblasts cells, large parenchyma cells and very thin- cells, a primary bark with large parenchyma in the parenchyma zone of the primary walled phellem cells (black and red). Root cells and an irregular, very thin-walled phel- bark. Root collar of a succulent 10 cm-high collar of a long hanging chamaephyte with lem. A lateral meristem consisting of a band plant, ruderal site, thermophile zone, sub- succulent leaves, wall, thermophile zone, of small cells is outside of the phloem. Root tropical climate, Gomera, Canary Islands. subtropical climate, Gomera, Canary Is- collar of a succulent 10 cm-high plant, rud- Mesembryanthemum nodiflorum, transverse lands. Aptenia cordifolia, transverse section. eral site, thermophile zone, subtropical cli- section, polarized light. mate, Gomera, Canary Islands. Mesembry- anthemum nodiflorum, transverse section.

Characteristic features of taxa Present features in relation to the number of analyzed species IAWA code frequency The presence or absence of ray-like radial strips of thin-walled Total number of analyzed species 5 2 growth rings indistinct or absent 5 parenchyma, the size and distribution of earlywood vessels, as 5 diffuse-porous 5 well as the presence of rhaphides can differentiate species. There 10 vessels in radial multiples of 4 or more common 3 is not enough material to present a definite classification neither 40.2 earlywood vessels: tangential diameter 20-50 μm 5 in relation to species nor to growth forms. 50.2 200-1000 vessels per mm2 in earlywood 5 58 dark-stained substances in vessels and/or fibers present (gum, tannins) 1 61 fiber pits small and simple to minutely bordered Ecological trends and relations to life forms (<3 μm = libriform fibers) 5 70 fibers thin- to thick-walled 5 Since all species analyzed grow in dry regions an ecological 75 parenchyma absent or unrecognizable 3 grouping could not be recognized. 79 parenchyma paratracheal 2 96 rays exclusively uniseriate 1 97 ray width predominantly 1-3 cells 2 98 rays commonly 4-10-seriate 4 Discussion in relation to previous studies 105 ray: cells upright or square 2 117 rayless 2 Carlquist 2007 analyzed much material and studied in detail 133 successive cambia, Caryophyllacea type 5 133.2 successive cambia, concentric continuous 5 the ontogeny of successive cambia of 11 perennial species of 11 134.1 conjunctive tissue thin-walled 5 genera representing a wide range of growth forms. The present 149 raphids present 2 study does not include the whole range of anatomical struc- R1 groups of sieve tubes present 1 tures. Not represented are species with vascular strands (Stayle- R10 phloem not well structured 5 ria neilii) or species with scalariform inter-vessel pits. R11 with rhaphides 2 P1 with medullary phloem or vascular bundles 1 38 Amaranthaceae

Number of species, worldwide and in Europe Analyzed species: Achyranthes aspera L. The Amaranthaceae family, including Chenopodiaceae, has 170 Achyranthes sicula (L.) All. genera with 2400 species. Cosmopolitan and especially charac- Aerva javanica Juss. ex Schult. teristic of disturbed, arid or saline habitats. Aerva persica Merr. Agathophora alopecuroides (Del.) Fenzl. In Europe there are 38 genera with 106 species. The majority are Amaranthus blithum L. Amaranthus cruentus L. represented by the genera Salsola (25 species), Chenopodium (23 Amaranthus deflexus L. species), Atriplex (19 species), and Amaranthus (12 species). Amaranthus hybridus L. Amaranthus lividus L. Amaranthus retroflexusL. Amaranthus standleyanus Covas Analyzed material Amaranthus viridis L. Anabasis brevifolia C.A. Mey The xylem and phloem of 15 genera with 62 species are ana- Arthrocneumum fruticosum (L.) Moq. lyzed here. Arthrocneumum glaucum (Del.) Ung. Arthrocneumum macrostachya Moris et Delponte Life forms analyzed: Studies from other authors: Arthrocneumum perenne (Mill) Moss. Atriplex canescens (Pursh) Nutt.

Amaranthaceae Nanophanerophytes 0.5-4 m 15 30 species Atriplex dimorphostegia Kareilin & Kiriloff Woody chamaephytes 11 Atriplex glauca Maire Semi-woody chamaephytes 3 16 genera Atriplex halimus L. Atriplex patula L. Liana 3 1 species Atriplex portulacoides L. Hemicryptophytes and geophytes 3 Atriplex prostrata DC Therophytes 27 1 species Atriplex saggitata Bork. Atriplex semibaccata R.Br. Plants analyzed from different Studies from other authors: Bosea cypria Boiss.ex Hook vegetation zones: Bosea yervamora L. Alpine and subalpine 1 Cheneloides tomentosa Botsch Hill and mountain 12 Chenopodium album L. Chenopodium bonus-henricus L. Mediterranean 12 2 species Chenopodium frutescens C.A. Mey Arid 22 23 Chenopodium glaucum L. Chenopodium hybridum L. Subtropical 10 2 species Chenopodium murale L. Tropical 2 species Chenopodium polyspermum L. Chenopodium strictum Roth Chenopodium urbicum L. Cornulaca monacantha Del. Einadia nutans Scott Haloxylon articulatum (Moq.) Bunge Haplopeplis perfoliata (Forsk.) Bg. ex Schweinf. Kochia prostrata DC Krascheninnikovia ceratoides (L.) Gueldenst. Krascheninnikovia lanata (Pursh) A. Meeuse & Smit Mairena pyramidata (Benth.) Paul G.Wilson Noea mucronata Ascherson et Schweinf. Patellifolia patellaris Scott et al. Patellifolia procumbens F. L. et W. Polycneum arvense L. Salsola foetida Del. Salsola genistoides Juss. ex Poir. Salsola kali L. Salsola oppositifoila Desf. Salsola vermiculata L. Salsola verticillata Schousboe Suaeda fruticosa Forsk. Suaeda pruniosa Lange Suaeda vera L.F. Gmelin Suaeda vermiculata Forsk. Traganum moquinii Webb Haloxylon ammodendron (photo: W. Schulze) 39 Amaranthaceae

Amaranthus caudatus (photo: Zinnert) Amaranthus lividus

Salsola kali Chenopodium bonus-henricus