Atlas of Stem Anatomy in Herbs, Shrubs and Trees: Volume 2

414 Pages·2013·71.12 MB·English

by  Fritz Hans Schweingruber

F. H. Schweingruber A. Börner E.-D. Schulze
Atlas of Stem Anatomy in Herbs, Shrubs and Trees
Volume I1
F. H. Schweingruber
A. Börner
E.-D. Schulze
Stem Anatomy
Atlas of
in Herbs, Shrubs and Trees
Volume I1
With contributions by
B. Kirchoff, M. Küchler and M. Nobis
With over 1500 colour illustrations
Prof. Dr. Fritz Schweingruber
Institute for Forest, Snow and Landscape Research WSL
Zürcherstrasse 111
Dr. Bruce K. Kirchoff
8903 Birmensdorf, Switzerland
Department of Biology
Annett Börner University of North Carolina at Greensboro
Greensboro, NC 27402-6170, USA
PO Box 808
Melrose Park SA 5039, Australia Dr. Meinrad Küchler
Dr. Michael Nobis
Prof. Dr. Ernst-Detlef Schulze
Institute for Forest, Snow and Landscape
Max Planck Institute for Biogeochemistry
Research WSL
PO Box 100164
Zürcherstrasse 111
07701 Jena, Germany 8903 Birmensdorf, Switzerland
ISBN 978-3-642-20434-0 e-ISBN 978-3-642-20435-7
DOI 10.1007/978-3-642-20435-7
Springer Heidelberg New York Dordrecht London
Library of Congress Control Number: 2012942068
© Springer-Verlag Berlin Heidelberg 2013
The photos on the following pages are published with the kind permission of the respective authors, whose names are indi-
cated in the figure legends:
Louis-M. Landry - pp. 33, 145, 118, 160, Per Arvid Åsen - p. 158 Pablo Necochoa - p. 40
203, 181, 199, 160 Faria Chowdhury - p. 106 Patrick Nicolas - p. 7
Marianne Lauerer - pp. 101, 143, 169, Brad Cotten - p. 40 Luigi Rignanese - p. 73
247, 256, 208 Hubertus Eicke - p. 118 Dave Riseborough - p. 145
Gregor Aas - pp. 129, 137, 262 Willem Frost - p. 169 Apollonio Tottoli - p. 98
Barry Rice - pp. 199, 254, 259 Grenville Godfrey - p. 33 Ann Van Roy - p. 162
Leonora Enking - pp. 7, 162 Chris Hendrickson - p. 280 Wolfgang Völkl - p. 84
Manuel Gil - pp. 205, 256 Wolfgang Katz - p. 173 Anne Vorstenbosch - p. 162
Patricio Novoa - pp. 38, 106 Elias Landolt - p. 118 BriAnna Weldon - p. 205
Barry Breckling - pp. 40, 98 Graeme Lyons - p. 199 John Winder - p. 280
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Cover illustrations (from right):
Cross-section of a dwarf shrub stem with successive cambia. Vessels and fibers are stained red, parenchyma cells are stained
blue. Chenopodium frutescens, Amaranthaceae, grows in the Mongolian steppes.
Cross-section of an old rhizome of an herb. The large red stained rays separate yellow stained radial vessel/fiber zones. Peuce-
danum venetum, Apiaceae, grows in the dry meadows of the Southern Alps.
Radial section of a liana stem. Radially arranged crystals in the vessel of a vine. Vitis vinifera, Vitaceae, grows in Mediterranean
riparian zones.
Cross-section of a water plant stem. Vessels in the center of the stems are surrounded by the phloem and an airconducting
tissue. The white dots represent calcium oxalate crystals. Myriophyllum alternifolium, Haloragaceae, grows in ponds.
The picture to the left is part of Peucedanum venetum.
All slides were stained with safranin and astra blue and photographed in
polarized light.
Cover design: deblik Berlin, Germany
Camera-ready by Annett Börner, Adelaide, Australia
Printed on acid-free paper
Springer is part of Springer Science+Business Media (www.springer.com)
V
Acknowledgements
We have to thank many colleagues and institutions: Katharina Neumann (Frankfurt M., Germany) provided slides
from Northern Africa. Patrik Kuss (Zurich, Switzerland) pro-
The Federal Research Institute and the Max-Planck Institute vided many species from the Yamal Peninsula of Russia.
for Biogeochemistry provided a scientific infrastructure and
financial support at Birmensdorf (FS) and at Jena (EDS and Stephan Shiyatov (Yekaterinburg, Russia), Eugene Vaganov
AB). Many colleagues in Birmensdorf, in the mechanical work- and Vera Benkova (Krasnoyarsk, Russia) collected material on
shop (Arthur Kölliker), the carpenter shop (Sigi Witzemann, many expeditions in Siberia and helped with the identification
Albert Buchwalder), the IT-Departement (Bert Höwecke), the of plants. Victor Voronin (Irkutsk, Russia) provided an excellent
library (Christine Matter, Claudia Grütter-Berger) and friends collection of the cold steppes of the Lake Baikal.
supported the study. Silvia Dingwall and Melissa Dawes spent
much time to edit the texts. Willy Neuhaus was always willing Marina Mosulishvili introduced FS to the Flora of Georgia and
to explain FS mechanisms of the Excel-Format. identified all species from the Caucasus region.
We thank Dr. David Remington, UNC Greensboro USA for Fidel Roig jun. (Mendoza, Argentina) accompanied FS on an
his assistance with the statistical analyses in the Paedomorphosis excursion to the Andes and his father Fidel Roig sen. identified
chapter, and for helpful discussion of these results. many plants from Argentina and Chile. Davoud Parsa Pajouh,
Karadj, Iran, accompanied FS on excursions in Iran. Martin
Pieter Baas (Leiden, Netherlands), David Drew (Hobbart, Aus- Fisher, Muscat, Oman, identified many species from Oman.
tralia), Helmut Freitag (Kassel, Germany), Patrick Fonti and
Holger Gärtner (Birmensdorf, Switzerland), Rudolf Häsler (Zu- Vera Markgraf (Flagstaff, USA) and Hal and Miriam Fritts
rich, Switzerland), Heike Heklau (Halle, Germany), Christian (Tucson, USA) supported FS with the collection and identifica-
Körner (Basel, Switzerland), Dean Nicolle (Adelaide, Australia) tion of plants from Colorado and Arizona. Hansjorg Diez (Zu-
and Simcha Lev-Yadun (Haifa, Israel) made many substantial rich, Switzerland) provided many species from the Great Plains
critical remarks and suggestions to improve the scientific con- in USA and Germany and John Banks (Canberra, Australia)
tent. from Australia.
Many Botanical Gardens provided Material: Basel, Switzerland We also thank the following people for providing photos: Klaus-
(Bruno Erni), Bern, Switzerland (Christian Bühler), Ekaterin- Dieter Zinnert, Elias Landolt, Marianne Lauerer, Thomas Stüt-
burg, Russia (Sergei Shavnin), München, Germany (Susanne zel, Gregor Aas, Per Arvid Åsen, Barry Breckling, Faria Chowd-
Renner), Regensburg, Germany (Peter Poschlod), Zurich, Swit- hury, Simon S. Cohen, Brad Cotten, Hubertus Eicke, Leonora
zerland (Bernhard Hirzel and Peter Enz), Viera y Clavijo, Jar- Enking, Willem Frost, Manuel Gil, Grenville Godfrey, Chris
din Canario, Gran Canaria, Spain (David Bramwell), Jardim Hendrickson, Harmen Hendriksma, Ottmar Holdenrieder,
Botanico Lisboa, Portugal, Gärtnerei Ernst Rieger, Blaubeuren, Wolfgang Katz, Louis-M. Landry, Graeme Lyons, Tara Massad,
Germany. Gary A. Monroe, Pablo Necochoa, Patrick Nicolas, Patricio No-
voa, Angela Nüske, Barry Rice, Luigi Rignanese, Dave Risebor-
The xylarium of the Rijksherbarium Leiden provided some ma- ough, Birgit Schulze, Waltraud Schulze, Horst Thor, Apollonio
terial. Alan Crivellaro (Padova, Italy) provided a complete slide Tottoli, Ann Van Roy, Wolfgang Völkl, Anne Vorstenbosch,
collection from Cyprus. BriAnna Weldon, John Winder and Scott Zona.
Ulf Büntgen (Birmensdorf, Switzerland) and Berhard Frauen- FS especially thanks his wife Elisabeth. Without her patience at
berger (Freiburg i.Br., Germany) provided many species from home and on countless field trips the present work would not
the east coast of Greenland. have been possible.
VII
Table of Contents
Acknowledgements ................................................V Ebenaceae ....................................................169
Frankeniaceae ..............................................173
Abbreviations ....................................................VIII
Garryaceae ...................................................176
1. Introduction to Volume 2 ...................................1
Hippuridaceae .............................................179
2. Monographic Descriptions .................................5 Hydrangeaceae ............................................181
Actinidiaceae ...................................................7 Lamiaceae ....................................................186
Adoxaceae ......................................................11 Lentibulariaceae ...........................................199
Apiaceae ........................................................17 Linnaeaceae .................................................203
Aquifoliaceae .................................................29 Myrsinaceae .................................................205
Araliaceae ......................................................33 Oleaceae ......................................................208
Asteraceae ......................................................38 Orobanchaceae ............................................216
Mutisieae .................................................40 Plantaginaceae .............................................226
Cardueae .................................................43 Polemoniaceae .............................................247
Lactuceae .................................................54 Rafflesiaceae ................................................252
Inuleae.....................................................64 Roridulaceae ................................................254
Gnaphalieae .............................................68 Sapotaceae ...................................................256
Calenduleae .............................................73 Sarraceniaceae ..............................................259
Astereae ...................................................76 Scrophulariaceae ..........................................262
Anthemideae ...........................................83 Solanaceae ...................................................271
Senecioneae .............................................91 Styracaceae ..................................................280
Heleniae ..................................................98 Valerianaceae ...............................................282
Heliantheae ...........................................101 Verbenaceae .................................................288
Eupatorieae ...........................................106
3. Ecological, Morphological, Taxonomical and
Tageteae .................................................108 Functional Significance of Stem Features Within
Summary ..............................................109 the Dicotyledons ............................................293
Balsaminaceae ..............................................113
4. Anatomical Adaptations to Environmental Condi-
Boraginaceae................................................117
tions ...............................................................355
Callitrichaceae .............................................127
Campanulaceae ...........................................129 5. Ontogeny of the Xylem ..................................371
Caprifoliaceae ..............................................137 6. Secondary Woodiness and Paedomorphosis ....383
Clethraceae ..................................................143
7. Conclusions ...................................................387
Convolvulaceae............................................145
Cornaceae ....................................................154 References ..........................................................389
Diapensiaceae ..............................................158 Alphabetic List of Species ...................................395
Diervillaceae ................................................160
Dipsacaceae .................................................162
VIII
Abbreviations
ae aerenchym mu mucilage
bpit bordered pit nu nucleus
ca cambium p perforation
cal callus, parenchymatic cells pa parenchyma
clu cell lumen, cell lumina ph phloem
co cortex phe phellem
cork phg phellogen
ct conjunctive tissue pit
cry crystal pith
csi collapsed sieve tubes
cu cuticula r ray
rd resin duct
di (ray) dilatation
ds dark-stained substances sc sclereid
duct sf septate fibers
shc sheath cell
ep epidermis si sieve tube, sieve element
en endodermis spit simple pit
ew earlywood
ewv earlywood vessel ta tannins
ewt earlywood tracheid te tension wood
tr tracheid
ft fiber tracheid ty tylosis
f fiber
ulcw unlignified cell wall
ge gelatinous fibers
gr growth ring v vessel
grb growth ring boundary vab vascular bundle
vat vascular tracheid
he helical thickenings vrp vessel-ray pits
ivp intervessel pit xy xylem
la laticifers
lf libriform fiber
lcw lignified cell wall
lw latewood
lwv latewood vessel
lwt latewood tracheid
1
1. Introduction to Volume 2
1.1 Phylogeny and Taxonomy
Volume 2, as a continuation of Vol. 1, provides monographic During the process of family characterization for Chapter 2,
descriptions for an additional 1295 species from 39 families. additional species have been collected, prepared and analyzed.
The major objectives, sampling design, sample preparation, and Therefore, the statistical analyses in Chapters 3 and 4 are
the anatomical characterizations are identical in both volumes. based on an extended dataset of 3347 species. An up-to-date
With the description of 2948 species within 124 families, Vol- table of all analyzed species can be found on the internet at
umes 1 and 2 cover all dicotyledonous classes and plant orders, http://www.wsl.ch/dendro/xylemdb/index.php.
excluding monocotyledons, as indicated in Fig. 1.1 on the next
page.
The scientific limitations of the dataset are described in Vol.1
on p. 2.
1.2 Geographic and Climatic Origin
Origin of sampled species
The majority of plants had been collected in Western Europe 1000
and Russia with fewer species collected in the Canary Islands 3 Africa
12 Australia, S-Asia
(Macaronesia), northern Africa, the Americas and Australia 52 S-America
(Fig. 1.2). ns 800 130 N-Africa, near East
o 157 Macaronesia
ati 325 N-America
The dataset covers all vegetation zones although the tropics are erv 600 2668 Europa, Russia, Georgia
s
b
represented by only a few species. The vegetation/climate zones o
are defined in Vol. 1 on p. 9. er of 400
b
m
u
N 200
Right: Fig. 1.2. Number of the analyzed species in relation to the
geographic origin and plant height. 0
2-10 10-25 25-50 50-100 100-150 150-300 >300
Plant height (cm)
F. H. Schweingruber, A. Börner, E.-D. Schulze, Atlas of Stem Anatomy in Herbs, Shrubs and Trees,
DOI 10.1007/978-3-642-20435-7_1, © Springer-Verlag Berlin Heidelberg 2013
2
Angiosperm Phylogeny Flowering Plant Systematics
A AMBORELLALES Amborellaceae
N GR NYMPHAELALES Cabombaceae Hydatellaceae Nymphaeaceae
TI AD AUSTROBAILEYALES Austrobaileyaceae Schisandraceae (incl. Illiaceae) Trimeniaceae
A E CHLORANTHALES Chloranthaceae
SM CANELLALES Canellaceae Winteraceae
A REPSOIGNYLR MAGNOLIDS PLMAAICAPUOGERRRNAAAOLLLELEEISASSLES DACHGAAnrceayoiodnsglmytrnoecoaoonnalcroraenteaeccrtacihhgeaeeiaaacaacceeceee aeaaeaeee EHLHPSauiaiepmupurenraoraruanmacrntcaeaadecantiaiaeeadecacr eaeeacaeeea e MMMSiayoprgnaisnirmutoicnilaiaaacccceeeeaaaaeeee
EA ALISMATALES AAlpisomnoagtaectoenaaec (einacel. LimBnuotcohmaarictaecaeeae) JPuonscidaogniniaacceeaaee RScuhpepuiacchezaeeri a c eae
Araceae Hydrocharitaceae Potamogetonaceae Zosteraceae
MONOCOTS PETROSAVIALES Petrosaviaceae
DIOSCOREALES Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae
PANDANALES Cyclanthaceae Pandanaceae Velloziaceae
LILIALES CAlosltcrhoiecmaceeriaaeceae LCioliarscieaaceeae PMeetlearnmthaiannceiaaceeae PSmhilielasciaacceeaaee
Amaryllidaceae (incl. Agapanthaceae, Alliaceae)
ASPARAGALES Asparagaceae (incl. Agavaceae, Hyacinthaceae, Ruscaceae)
n HXaypnothxoidrrahcoeeaaec eae (incIlr.i dAascpehaoed leaceae, HeLmaenraorciaaclleidaaeceae) Orchidaceae
o ARECALES Arecaceae
cti COMMELINDS PCOOAMLMEESLINALES BCCryoopmmemreaelicaliencaeaeaceeaeJuEncriaoHcceaaaeuemlaocedaoReraacpeaateePaocaecaePeaoentedTeyrpRiahecasectaeioeanea c(ienacel. SparXgaynridiaacceeaaee)
u ZINGIBERALES CCaonstnaacceeaaee HLoewliciaocneiaaceeae MMaursaancteaaceeae SZitnregliibtzeiaracceeaaee
d
CERATOPHYLLALES Ceratophyllaceae
ro RANUNCULALES BCeirrcbaeeraidsatecreaacee ae ELaurpdtiezlaebaacleaaceeae MPaepnaisvpeerarmceaaceeae Ranunculaceae
t SABIALES Sabiaceae
n PROTEALES Nelumbonaceae Platanaceae Proteaceae
I OTS TROCHODENDRALES Trochodendraceae
DIC BUXALES Buxaceae (incl. Didymelaceae) Haptanthaceae
U
E GUNNERALES Gunneraceae Myrothamnaceae
DILLENIALES Dilleniaceae
SAXIFRAGALES CAletirncgidiaipcheyalela ceae DGaropshsnuiplahryiallcaecaeeae HPaaemoanmiaecleidaaeceae
Crassulaceae Haloragaceae Saxifragaceae
VITALES Vitaceae
ZYGOPHYLLALES Krameriaceae Zygophyllaceae
CELASTRALES C(ineclal.s Htriapcpeoacerateaceae, Brexiaceae) LPeaprnidaosbsoiatrcyeaaceeae
Achariaceae Euphorbiaceae Ochnaceae Podostemaceae
MALPIGHIALES CChlursyisaocbeaalea naceae HLiynpaecreicaaeceae PPahsysllaiflnotrhaacceeaaee RRahfizfloepsihaocreaaceeae
IOSIDS OXALIDALES BCErreuyptnhheralollioxatycaleacaceeea aee MaCClpouignnhnoianaricaaeccaeeeaaee PicrodEHenluadaercoaeccaaeerapeaceSaea l ic aceOaxealidVacioelaaceeae
COTS EUR RFAOBSAALLEESS FBCaaabrnbanecayebaaaceceeaaee EMPlooalreyagacagelnaaaceceeaaee QRUolumsilalaaccjaeecaaeeeae Surianaceae
DI Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae)
U
C EORE OSIDS CFAUGCAULREBSITALES BCBAeenatgissuuoolaanpcrihaienycaalelecaa eceaeea e FJCCuaooggrriylaaancrnoieadcacaaeecr aepeaa eceae MNCDouaytctrhiiuscocrabfacaicetgaeaacaeceeeaaee RTTeihctooraidpmeteneldlearacaceceeaaaeee
R GERANIALES Francoaceae Geraniaceae Ledocarpaceae Melianthaceae
Combretaceae Myrtaceae Penaeaceae (incl. Oliniaceae)
MYRTALES LMyethlarsatcoemaaet a(icnecal. eP (uinncicl.a Mceeamee, cSyolancneearaet)iaceaOen, aTgrarapcaecaeeae) Vochysiaceae
CROSSOSOMATALES CGreoissssoolsoommaatatacceeaaee SSttaacphhyyuleraacceeaaee Strasburgeriaceae
IIOSIDS SPAICPRINADMANLIAELSES BPAuincrarsacemarrandciiaaeccaeeeaa ee MNietrlaiarciaecaeea e SRauptaincdeaacee (ainecl. CneorSacimeaaero) ubaceae
EUR HUERTEALES Dipentodontaceae Gerrardinaceae Petenaeaceae Tapisciaceae
Bixaceae Malvaceae (incl. Bombacaceae, Sterculiaceae, Tiliaceae)
MALVALES CDiispttaecroecaaerpaceae Cytinaceae MNeuunrtaindgaiacceeaaee SThayrcmoelalaeenaacceeaaee
BRASSICALES BBaratassciecaaeceae CCaleroicmacaecaeeae LMimorninagnathceaaceeae SToavlvaaridaocreaaceeae
Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae
BERBERIDOPSIDALES Aextoxicaceae Berberidopsidaceae
SANTALALES BLoarlaannothpahcoeraaeceae MOliasocadceenaderaceae SOapnilitaaclaecaeeae (incl. VisScaccheoaeep)fiaceae
Aizoaceae Caryophyllaceae Molluginaceae Polygonaceae
Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae
CARYOPHYLLALES B(inacsle. Cllahceenaoep odiaceae)DDrroosseorpahcyellaaeceae NPhycyttaoglainccaacceeaaee STaimlinmaocnedaseiaceae
Cactaceae Frankeniaceae Plumbaginaceae Tamaricaceae
CORNALES CCourrntisaicaecaeea e GHyrudbrabniagceeaaceeae LNoyasssaacceeaaee
Actinidiaceae Ericaceae Myrsinaceae Polemoniaceae Sarraceniaceae
ERICALES BCalelstharmacineaacee a e FEobueqnuaiceeriaaec eae PRroimriduulalacceeaaee SThtyeraacceaaceeae
Diapensiaceae Lecythidaceae Sapotaceae Theophrastaceae
GARRYALES Eucommiaceae Garryaceae (incl. Aucubaceae)
RIDS GENTIANALES GApeonctiyannaacceeaaee (incl. AsGcleelpsieamdaiacceeaaee) RLougbaianciaecaeeae
ASTE I SDIRETSAU LSAOMLAIANLAELSES GBHBACHyicyoyegbaddnsnlnrrnviooodtoenhslaleriavticaaaauceccccelaeaheeaeacyaaeeeaeeacee a(eincl. CMOLLOMeaulraoeonmsrnabttciiytcabauinnecuntiaiaealacaceachcreeeeiaaaaacceeeee)aaee PPSSPPaneaophladdulhrany aleHontmalniwaiagoapcnciccpeneileuaaeaaarceacieeede c (aa eai necaecee l(a.i neNc)ol.l aVSSCnetcaiarlrlbbolciateeprcinahceaeuhac)laaeecr aieaeacee ae
E BORAGINALES BCoordaogninaacceeaaee CEhorredtiaiacceeaaee (incl. LenHneolaiocteroapei)aceae WHyedllrsotpehdyiallcaecaeeae
AQUIFOLIALES Aquifoliaceae Cardiopteridaceae Stemonuraceae
Asteraceae Goodeniaceae Pentaphragmataceae
F(Fwioinwgl l.wro e1w2d..)i1bn i.ign oP lcCohlguoyidell.oeefdgu ea -ibnnneed rtbl iioHcn t.htidrl eegv/eose ylrisunb m(do2tie0/cpsa1 oto0sitfn)e ,gtr h/ ptisho sestt eufra1dm.yp.i dlifes II ESDIRETSAU PBEAAARSSPRCTIUAEAANLRCLIEALARSLOLYEENPSSIHAILAELSES CAPBECAparsraauacirlmayanlaiccapcilaleecroacyraennaepeaiucahaeleeicaaaecceaeeeaaee (incl. CGMMLooreyiblsonueedymllaoiianenccilatalehiracapaceeacea)ecaeeae ae (e incl. PRPSDeitoetytnusolinfsdsosaipnaenotcatareicaainaeccieaeeaeaa)ee
DIPSACALES CAdaopxriafocleiaaceeae DDiipesrvaicllaacceeaaee MLinonrianeaaceceaaee Valerianaceae
3
1.3 Life Forms
The dataset includes eleven life forms as described by Raunkiaer
(Ellenberg et al. 1992).
P: Phanerophytes. Woody plants that grow taller than 4 m T: Therophytes. Annuals. Plants whose shoot and root system
(trees). dies after seed production and which complete their life cycle
within one growing season (spring, summer, fall). Included
N: Nanophanerophytes. Woody, shrub-like 0.5-4 m high
are only specimens with one tree ring.
plants.
Liana: Mostly perennials, plants with extremely long shoots,
C: Chamaephytes. Herbaceous to semi-woody perennials. I
which need normally structural support by other species. n
Dwarf shrub-like plants whose mature branch or shoot sys- t
r
tem remains perennially 25-50 cm above ground surface. Climbers: Plants climbing with adhesive roots growing over o
other plants or objects. d
Z: Woody chamaephytes. Dwarf shrubs with less than 50 cm
u
height. Hydrophyte: Water plants where most parts live and root below c
t
H: Hemicryptophytes. Perennial herbaceous (including bienni- the water table. io
als) plants. With periodic shoot reduction to a remnant shoot Helophyte: Terrestrial plant rooting in wet ground. n
system that grows relatively flat on the ground. Here we in-
Succulent: Plant with water-storing tissues.
clude most of the geophytes (G) whose surviving shoot system
normally remains below surface. Also included are winter-
annuals, which perform their life cycle between fall to late
spring. These winter annual plants form two rings.
1.4 Anatomical Features
All features as described in Vol. 1, pp. 13- 32 have been used
for the characterization of the species described in Vol. 2. Two
additional features (Features 60.1 and 70.3) are used for the
characterization of species in Vol. 2 (Figs. 1.3-1.6).
60.1 Fibers absent. 70.3 Fibers absent in the stem
center.
250 µm 250 µm 250 µm 250 µm
Fig. 1.3. Euphorbia nicaeensis, Fig. 1.4. Arenaria ciliata, Fig. 1.5. Nonea erecta, Boragi- Fig. 1.6. Jovibarba hirta, Crassu-
Euphorbiaceae, herb. Caryophyllaceae, herb. naceae, herb. laceae, succulent plant.
4
1.5 Morphological and Ecological Interpretation of Features
In Chapters 3 and 4 of the present volume we relate the stem Lignification appears to be related to water, nutrient and phy-
anatomical features of the whole dataset to plant height, as it tohormone transport (Nardini et al. 2011), which in turn is
appears that static and hydraulic functions have been optimized related to plant size and leaf area. The wood anatomy would
in evolutionary processes, mainly in response to plant height. also be affected by climate seasonality (drought or frost), which
The actual APG III DNA-based taxonomic classification gener- results in tree rings with early- and latewood. A seasonality of
ally does not coincide with specific morphological or anatomi- water demand with bud break and flushing leaves will cause
n cal characteristics. The description of plant diversity in view of a differentiation in the dimensions of vessels even during a
o
the differentiation in xylem structures remains a major prob- single growing season. In seasonal climates, plants may avoid
i
t
c lem. The “life-form” classification of Raunkiaer (1904), which climatic stress by the shedding of leaves, short shoots (cladap-
u is still commonly used in vegetation science, does not treat stem tosis), branches, or the whole above-ground canopy (hemicryp-
d
features adequately. The Raunkiaer classification is based on the tophytes), or they may endure the climatic stress and have ever-
o
r position of the regenerating bud in relation to the soil surface as green foliage and perennial stems. This has major implications
nt a survival strategy in seasonal climates - an interpretation which for the storage of reserves in seasonal climates, and on stem
I has been questioned in the past (see Schulze 1982). Also, bud anatomy. Reserves could be stored in the leaves, stems, hypo-
height may be quite different from plant height considering the cotyl (including the root collar) or in the phloem and bark, and
flower stalks of herbaceous species. Lignification does not cor- this will affect the presence of parenchyma. Large vessels, which
respond with the Raunkiaer types of life forms (therophytes, are essential in deciduous species, would not be functional in
hemicryptophyte, chamaephytes and phanerophytes). The xy- evergreen trees occurring in seasonal climates (Schulze et al.
lem of small and large annual plants (e.g. Arabidopsis thaliana 2005). Thus, all kinds of transitions will exist between these two
and Helianthus annuus) can be fully lignified, as in trees. In con- adaptive techniques.
trast, some small plants, such as Silene acaulis and water plants,
consist primarily of un-lignified cells. Following these considerations, the degree of “woodiness” or
“herbaceousness” would be related to (1) plant size and height,
In contrast to Raunkiaer’s “life forms”, Monsi (1960) based a (2) the seasonality of climate, and (3) the persistence of foli-
structural description on the economy of resource use during age. In the following we will maintain the terms “woody” and
growth. He followed Clements (1920), who coined the words “herbaceous” because they are commonly used and easy to un-
“herbaceous” and “woody” as descriptors for plant structures. derstand, but in our definition we imply a dynamic range of
Monsi (1960) distinguished evergreen and deciduous species lignification, which could be species- or site-related. The degree
within both herbaceous and woody plants. However, the clas- of woodiness would be explained by the parameters of plant
sification of Monsi (1960) is questionable in an anatomical size/height, habitat seasonality, and the way reserves are stored
sense. Lens et al. (2011) describe all kinds of transitions be- in seasonal climates. Obviously, a more functional classification
tween more and less lignified plant stems. is needed to incorporate xylem anatomy.
In all monographic descriptions (Vol. 1 Chapter 5 and Vol. 2
Chapter 2) we have used Raunkiaer’s life form classification
because we were not aware that plant height correlates better
with anatomical features than life forms (see Chapter 3 in this
volume). Obviously, a more functional classification needs to
include xylem anatomy.