Riparian Tree and Shrub Propagation Handbook

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Riparian Tree and Shrub Propagation Handbook
RIPARIAN TREE AND SHRUB
PROPAGATION HANDBOOK
RIPARIAN TREE AND SHRUB
PROPAGATION HANDBOOK
An Aid to Riverine Restoration in the Mediterranean Region
Edited by
María Aránzazu Prada and Daniel Arizpe
CIEFBanc de Llavors Forestals
Conselleria de Medio Ambiente, Agua, Urbanismo y Vivienda, Generalitat Valenciana
Avenida Comarques del País Valencià 114
46930 Quart de Poblet, Valencia, España
© 2008 Generalitat Valenciana
© texts, illustrations and plates: the authors
Illustrations: Faustino Díez
Unless otherwise specified
Emilio Laguna
Clematis flammula, Coriaria myrtifolia, Dorycnium rectum,
Flueggea tinctoria, Lonicera implexa
María Aránzazu Prada
Liquidambar orientalis, Myrtus communis, Populus orientalis,
Salix amplexicaulis, Salix pedicellata and cover
Plates: Gabriel SegarraMoragues
Salix plates
ISBN: 9788448249656
Legal Deposit no.: V38782008
Design: Essência ROFFdesign (http://essencia.roff/pt)
Layout and Printing: Gràfiques Vimar (http://www.vimar.es)
English edition: copy editing by Mary Georgina Hardinge
Preface
My opportunity to contribute began to take shape
when I became aware of the riparian ecological res
toration efforts at the Higher Institute of Agronomy
at the Technical University of Lisbon. That frame
work prompted me to launch a project to establish
a working group with other European institutions
that share the same concerns. The Ripidurable Pro
ject was presented in April 2003 and approved to
wards the end of that year. It was launched with the
goal of creating a forum for communication and co
llaboration between those responsible for the ma
nagement and restoration of riparian areas and the
academic and research institutions with knowledge
and expertise in those habitats. The group could co
rrelate the problems detected with the existing kno
wledge in pursuit of the solution to those problems
and develop specific tools, including some case stu
dies, which could be offered to society.
One of the problems that most caught my attention
was the great difficulty that institutions and com
mercial companies involved with the recovery of ri
parian ecosystems had in obtaining suitable plants
for this purpose. Generally, companies that market
plants do not have suitable reproductive material for
these interventions, from the standpoint of either
external quality or adaptability. Therefore, the solu
tion has historically depended on acquiring plants
from other countries and, in many cases, on orna
mental varieties or fastgrowing hybrid poplars.
The publication of this riparian species propagation
guide is intended as a tool to help resolve the afo
rementioned situation and stimulate the produc
tion of plant materials from local sources. The team
from the Forest Seed Bank of the Regional Govern
ment of Valencia has contributed significantly to
the preparation of this guide. This group combines
hard work and rigour with the institution's lengthy
experience in the management and effective short
term use of germplasm, as well as its valuable work
on longterm conservation, a legacy for future
generations.
The team that developed this guide has made a sig
nificant contribution to both scientific and techni
cal knowledge through their extensive experience
and current academic research. The guide can now
be used by companies and institutions that intend to
produce plants for the great task of pursuing ade
quate recovery of riparian ecosystems.
By 2015, following the Water Framework Directive,
rivers should be classified as having good ecological
status, in accordance with the reference conditions.
It is our fervent hope that this guide will help to en
sure that this status is achieved within the planned
time and with the care that nature deserves, by ai
ding the production of quality plants that safeguard
the genetic heritage of each species.
The authors of this guide deserve our gratitude for
sharing their experience and knowledge with us all.
The editors, Arantxa Prada and Daniel Arizpe, des
erve particular appreciation for the effort that they
have put into this project and for the perseverance
they have shown in making this book a reality.
Ana Mendes
Higher Institute of Agronomy
Technical University of Lisbon
5 Preface
The habitats that are perhaps most intensely modi
fied by man are riverine habitats. I became aware of
this situation during numerous visits to Portuguese
riparian environments in search of a Cetti’s warbler
or a kingfisher. This knowledge sparked in me a des
ire to contribute, in my own small way, to the reco
very of these degraded environments for the benefit
of nature, of mankind and ultimately, for the pre
servation of biodiversity.
Contributors
Maria Helena Almeida
Universidade Técnica de Lisboa, Instituto Superior de
Agronomia, Centro de Estudos Florestais, Tapada da
Ajuda, 1349017 Lisbon, Portugal
José Vicente Andrés
Avenida Salvador Allende 75, esc. 14, 4ºD, 50015
Zaragoza, Spain
Juan Añíbarro
Viveros Fuenteamarga SL, polígono 7, parcela 18,
47260 Cabezón de Pisuerga, Valladolid, Spain
Daniel Arizpe
CIEFBanc de Llavors Forestals, Área de Gestión de Re
cursos Forestales y Conservación Ambiental, Conselle
ria de Medio Ambiente, Agua, Urbanismo y Vivienda,
Generalitat Valenciana, Avenida Comarques del País
Valencià 114, 46930 Quart de Poblet, Valencia, Spain
Antonio del Campo
Dep. Ingeniería Hidráulica y Medio Ambiente, Escuela
Técnica Superior de Ingenieros Agrónomos, Univer
sidad Politécnica de Valencia, Camí de Vera s/n,
46002 Valencia, Spain
Esperanza Campos
CIEFBanc de Llavors Forestals, Área de Gestión de Re
cursos Forestales y Conservación Ambiental, Conselle
ria de Medio Ambiente, Agua, Urbanismo y Vivienda,
Generalitat Valenciana, Avenida Comarques del País
Valencià 114, 46930 Quart de Poblet, Valencia, Spain
Carla Faria
Universidade Técnica de Lisboa, Instituto Superior de
Agronomia, Centro de Estudos Florestais, Tapada da
Ajuda, 1349017 Lisbon, Portugal
Cándido Gálvez
Semillas Silvestres S.L., Carretera de Santa María de
Trasierra km 2, 14012 Córdoba, Spain
Jose Luis García Caballero
Junta de Castilla y León, Servicio Territorial de Medio
Ambiente León, Avenida Reyes Leoneses 145ºC
(Edificio Europa), 24071 León, Spain
Pablo Jiménez
Universidade Técnica de Lisboa, Instituto Superior de
Agronomia, Centro de Estudos Florestais, Tapada da
Ajuda, 1349017 Lisboa, Portugal
Fernando Martínez Sierra
Junta de Castilla y León, Servicio Territorial de Medio
Ambiente León, Avenida Reyes Leoneses 145ºC
(Edificio Europa), 24071 León, Spain
Eduardo PérezLahorga
Área de Gestión de Recursos Forestales y Conserva
ción Ambiental, Conselleria de Medio Ambiente,
Agua, Urbanismo y Vivienda, Generalitat Valenciana,
Calle Francisco Cubells 7, 46011 Valencia, Spain
Mari Carme Picher
CIEFBanc de Llavors Forestals, Área de Gestión de Re
cursos Forestales y Conservación Ambiental, Conselle
ria de Medio Ambiente, Agua, Urbanismo y Vivienda,
Generalitat Valenciana, Avenida Comarques del País
Valencià 114, 46930 Quart de Poblet, Valencia, Spain
María Aránzazu Prada
CIEFBanc de Llavors Forestals, Área de Gestión de
Recursos Forestales y Conservación Ambiental, Con
selleria de Medio Ambiente, Agua, Urbanismo y Vi
vienda, Generalitat Valenciana, Avenida Comarques
del País Valencià 114, 46930 Quart de Poblet,
Valencia, Spain
Jesús Rueda
Junta de Castilla y León, Dirección General del Medio
Natural, Calle Rigoberto Cortejoso 14, 47071 Valla
dolid, Spain
Pilar Ventimilla
CIEFBanc de Llavors Forestals, Área de Gestión de
Recursos Forestales y Conservación Ambiental, Con
selleria de Medio Ambiente, Agua, Urbanismo y Vi
vienda, Generalitat Valenciana, Avenida Comarques
del País Valencià 114, 46930 Quart de Poblet,
Valencia, Spain
7 Contributors
Neus Albert
CIEFBanc de Llavors Forestals, Área de Gestión de Re
cursos Forestales y Conservación Ambiental, Conselle
ria de Medio Ambiente, Agua, Urbanismo y Vivienda,
Generalitat Valenciana, Avenida Comarques del País
Valencià 114, 46930 Quart de Poblet, Valencia, Spain
Contents
Introduction
17 Species fact sheets
(M. Aránzazu Prada, Daniel Arizpe, Juan Añíbarro, Jesús
Rueda, Neus Albert, Esperanza Campos, Mari Picher, Pilar
Ventimilla, Cándido Gálvez, Carla Faria, Pablo Jiménez)
19
22
27
30
33
36
40
43
46
50
53
55
59
63
67
70
73
76
79
83
87
90
94
97
101
105
109
113
116
119
124
127
130
135
138
141
Fact sheet contents
Alnus glutinosa (L.) Gaertn.
Arbutus unedo L.
Celtis australis L.
Cercis siliquastrum L.
Clematis flammula L. y C. vitalba L.
Coriaria myrtifolia L.
Cornus sanginea L.
Crataegus monogyna Jacq.
Dorycnium rectum (L.) Ser.
Flueggea tinctoria (L.) G.L. Webster
Frangula alnus Mill.
Fraxinus angustifolia Vahl.
Hedera helix L.
Humulus lupulus L.
Laurus nobilis L.
Ligustrum vulgare L.
Liquidambar orientalis Mill.
Lonicera etrusca G. Santi y L. implexa Aiton
Myrtus communis L.
Nerium oleander L.
Pistacia lentiscus L.
Platanus orientalis L.
Populus alba L.
Populus nigra L.
Populus tremula L.
Prunus mahaleb L.
Prunus spinosa L.
Rubus ulmifolius Schott
Salix spp.
Sambucus nigra L.
Tamarix spp.
Ulmus minor Mill.
Viburnum tinus L.
Vitex agnuscastus L.
Vitis vinifera subsp. sylvestris (C.C. Gmelin) Hegi
147
149
Appendices
Variation and adaptation
152
Seed handling
(Helena Almeida, Carla Faria)
(M. Aránzazu Prada)
158
Nursery culture
162
Cutting propagation
165
169
171
172
174
190
195
(Antonio del Campo)
(Daniel Arizpe, M. Aránzazu Prada)
Stool beds
(José Luis García Caballero,
Fernando Martínez Sierra, Jesús Rueda)
Master certificate
(M. Aránzazu Prada)
Plant passport
(Eduardo PérezLahorga)
Populus spp. (diagnostic traits)
(José Vicente Andrés, M. Aránzazu Prada)
Salix spp. (distribution and diagnostic traits)
(José Vicente Andrés, M. Aránzazu Prada)
Tamarix spp (distribution and diagnostic traits)
(José Vicente Andrés, M. Aránzazu Prada)
Glossary
9 Contents
11
1
11
Introduction
Introduction
Riparian vegetation is of great ecological interest due
to the part it plays in numerous processes that have
to do with the quality of the physical environment
and the life cycles of the aquatic and terrestrial
fauna associated with fluvial systems. It connects
different habitats and improves the quality of other
terrestrial, aquatic or coastal/marine systems.
Fluvial systems in the Mediterranean region have
been greatly altered by human activity. Their chan
nels and riverside areas have been transformed, first
into farmland and more recently into urban deve
lopments. Human activity has also regulated flow vo
lumes, canalisation projects have destroyed the
connectivity between river courses and alluvial plains
and surface and underground water reserves are ove
rexploited. These alterations have directly or indi
rectly affected the natural riverside vegetation by
reducing its biodiversity, scattering its populations
and, in some extreme cases, making it disappear en
tirely from large stretches of rivers.
In the face of such general deterioration, restoring
fluvial ecosystems by recovering natural hydrological
regimes is an unavoidable task. Such interventions
may require plantation projects as a means of reco
vering riverside vegetation in the short term. Other
specific objectives might aim at enriching the flora or
introducing species that may have disappeared as a
result of human activity or that play a vital role in
plant/animal interactions. An additional objective
could be the development of tree coverage to com
pete with and eliminate alien species. Planting may
also be advisable in order to increase the genetic
base of populations, especially when these have suf
fered a decrease in the number of individuals or a
loss of geneflow rates due to human pressure, or
when inappropriate use has been made of reproduc
tive materials in the past, especially as regards spe
cies with vegetative propagation patterns. After
these interventions, however, the river itself will
eventually model the structure and the dynamics of
its riparian vegetation.
In any event, the generation of reproductive mate
rials for use in river restoration projects must try to
ensure the viability of the newly introduced popula
tions without impairing existing genetic resources.
This is achieved, first of all, by proper species selec
tion. Since these plantings are not made for com
mercial purposes, native taxa and materials of local
provenance should be favoured whenever possible.
The genetic base of the chosen materials should also
be as wideranging as available resources permit, in
order to enhance the adaptability of new popula
tions. Alien species – some of which have already
become naturalised in riverside areas of the Medite
rranean region – must be particularly avoided, as well
as species that may hybridise with the local ones.
This guide has been conceived as a support tool for
nursery managers and for others who are not spe
cialists but are involved, in one way or another, in
the production of riparian plant species for hydrolo
gical restoration projects. It offers useful data on the
production of seeds, parts of plants and plants be
longing to a large series of tree, shrub and lianoid
species that may be employed in fluvial systems of
the Mediterranean region. It provides information on
species that are dominant in the riparian vegetation
of the region, on species whose propagation may be
of interest due to their potential interaction with
some species of fauna and on species that are tradi
tionally used for hydrological restoration purposes.
Although some of the taxa included are not strictly
riparian, they originate in the Mediterranean shrub
and forest communities and can develop optimally
in these environments, especially in extremely dry
areas.
The information is presented in the form of fact she
ets that include details of the production of repro
ductive materials – from collection to conservation –,
the physical characteristics of seeds and the most
appropriate methods for producing plants generati
vely and vegetatively. General data on each taxon are
13 Introduction
The fluvial systems of the Mediterranean region, with
their specific dynamics and environmental condi
tions, less extreme than those of surrounding
systems, are home to a mosaic of habitats with a
high degree of biodiversity and serve as a migration
path for many species of flora and fauna. They also
play a critical role in the life of human communities,
which use their resources and also benefit from them
as leisure spaces.
also provided: geographical distribution, ecology, ta
xonomic identification features and reproductive
biology. These technical sheets include additional de
tails that may be of use for improved management of
reproductive materials. In particular, information
about intraspecific variation – and its implications
for collecting and using materials – is provided whe
never possible with a view to promoting the conser
vation of genetic resources.
A series of appendices deals with certain specific
matters that are directly related to the production
and use of forest reproductive materials, such as in
traspecific genetic variation and its relevance in
terms of population adaptability, practical aspects of
the production and conservation of seeds and parts
of plants and the European regulations applicable to
the production, transport and marketing of some
species included in the present guide.
Another appendix has tables and figures which aim
to make it easier to identify species of the genera Po
pulus, Tamarix and Salix that can be found in the Eu
ropean Mediterranean region. It was also considered
helpful to provide a glossary of the scientific and
technical terms employed that are not commonly
used by the intended users of this guide.
We hope that the guide will prove a useful manual
for plant producers and contribute to the conserva
tion and improvement of our Mediterranean riverine
environments, which are part of our natural and cul
tural heritage.
We would like to thank Christine Fournaraki (Medi
terranean Agronomic Institute of Chania, Greece),
Isabel Montávez (Intersemillas SA, Spain), Fabio Go
rian (CFSCentro Nazionale per lo Studio e la Con
servazione della Biodiversità Forestale, Italy), Despina
Paitaridou (Ministry of Rural Development and Food,
Greece), Jesús Martínez y Sisco Bosch (Banc de Lla
vors Forestals de la Generalitat Valenciana, Spain),
Ana Santos and Filipa Pais (Câmara Municipal de
MontemoroNovo, Portugal), José Luis García Ca
ballero (Junta de Castilla y León, Spain) and Valeria
Tomaselli (CNR Istituto di Genetica Vegetale, Italy)
for supplying valuable data included in the technical
sheets, and Francisco Sánchez Saorín, Miguel Cáno
vas and Manuel Balsalobre (Región de Murcia,
Spain), Pedro Sánchez Gómez (Universidad de Mur
cia, Spain), Begoña Abellanas (Universidad de Cór
doba, Spain) and Isabel Butler (Universidad de
Huelva, Spain) for their help in obtaining illustrations
of willows. We are grateful to Esther Tortosa, Jesús
Rueda and Ana Puertes for reviewing and correcting
the original Spanish version of the text.
We would also like to thank Antoni Marzo for giving
us the opportunity to take part in the Ripidurable
project and all our other friends at the Forest Seed
Bank (Banc de Llavors Forestals) of the Regional Go
vernment of Valencia who have in one way or ano
ther supported the present initiative, particularly
Raquel de Miguel and Gloria Ortiz. We want to give
special thanks to Esther Tortosa, without whose en
thusiasm and professionalism our endeavours would
not have been crowned with success.
Finally, we wish to express our affection, thanks and
satisfaction to all our colleagues in the Ripidurable
project for sharing their knowledge and information
and for generating a warm atmosphere throughout
the programme. We hope that we may continue to co
llaborate on future projects to preserve biodiversity.
The editors
Introduction 14
(“...
Father,
tell me
what they’ve done to the river
to stop it singing.
It slides by
like a dead barbel
under a foot of white foam.
“...
Pare,
digueume
què li han fet al riu
que ja no canta.
Rellisca
com un barb mort
sota un pam d'escuma blanca.
Pare,
que el riu ja no és el riu.
Pare,
abans que torni l'estiu
amagui tot el que és viu.
...”
Joan Manuel Serrat
(Pare)
15 Introduction
Father,
the river’s not the river any more.
Father,
before the summer comes again
hide every living thing
...”).
2
17
Species
Fact Sheets
A descriptive fact sheet has been prepared for each
taxon. The fact sheets are designed to facilitate rapid ac
cess to the information. They include the common names
in different languages as well as the scientific name.
highlighted, especially if the taxa are sympatric. It has
not been possible to avoid botanical terminology in the
description of the taxa but the meaning of the terms
can be consulted in the glossary at the end of the book.
Distribution and Ecology
For a more detailed description of the taxa, reference
works such as Flora Europaea or other national or re
gional flora can be consulted.
A map shows the distribution of the taxon in Europe
and in Asian and African countries bordering the
Mediterranean basin. The basic reference works for
these maps are those in Bolòs and Vigo (1989), Charco
(2001), the Atlas of Flora Europaea and the online “Pro
grama Anthos” database. The natural distribution of
some species that have been widely dispersed by man
is hard to specify; for this reason, the maps for Cercis
siliquastrum, Laurus nobilis, Platanus orientalis, Salix
fragilis and Vitis vinifera subsp. sylvestris should be
taken purely as a rough guide.
The map of tamarisk distribution in the eastern
Mediterranean follows the monograph on the genus by
Baum (1978), completed in some cases by information
from other works such as Boratyński et al. (1992), Pig
natti (1982) and Zohary (1972). Because of the lack of
precise information about the chorology of these
species in some Eastern European countries, these
maps should be taken as only a basic approximation to
their distribution.
The general distribution of the taxon is indicated
schematically, mentioning the regions in which it is
present according to the division established by Brum
mitt (2001), irrespective of its abundance. There is also
a list of Mediterranean basin countries that have areas
with a Mediterranean bioclimate in which the species
is present. This information has been taken mainly
from two online databases: the digital version of Flora
Europaea for its European distribution and the
“Germplasm Resources Information Network” (GRIN)
for its distribution elsewhere in the world.
The ecology of the species is indicated in a simple, sum
marised way to make it easy to understand.
Diagnostic traits
Information on relevant diagnostic features for identi
fying the taxon is provided as concisely as possible. Dif
ferences that distinguish the taxon from others with
which it might quite easily be mistaken have been
Reproductive biology
As the phenology of reproduction and reproduction
systems in the taxon is a determining cause of the ge
netic configuration of populations, the most relevant
data are indicated schematically. This information is re
garded as important for deciding on the right strategies
for collecting reproductive materials, for creating new
populations and for their subsequent management.
The flowering and fruit ripening periods indicated nec
essarily cover a very broad range and present conside
rable interannual and intersite variations, especially
in widelydistributed species that grow in different cli
mate conditions.
The principal pollinating and dispersing agents are
mentioned, although in many cases other alternatives
for gene flow may exist. This situation is very common
among riparian species, where water may act as a sec
ondary disperser.
Variation and Hybridisation
Taxonomic observations are provided, such as the ex
istence of subspecies or the recognition of varieties and
natural hybrids. Likewise, information on the outcomes
of genetic studies is provided for some species to pro
mote an improvement in reproductive material han
dling and in conservation of the genetic resources of
the species.
Seed propagation
Tolerance to drying is indicated, as this is an aspect
which influences the treatment given to the seed lots
to a large extent. Practical information about collect
ing, producing and storing seeds is included. The clean
ing sequence to be employed, following the possible
seed handling procedures described in the appendix, is
19 Fact Sheet Contens
Fact Sheet Contents
indicated concisely. The storage conditions recom
mended – temperature (T), moisture content (MC) and
type of container – are the standard ones for short
and mediumterm storage of the materials, depending
on the type of seed.
The commonlyused treatments which have proved
most effective for stimulating germination are indi
cated. The length of the treatments is for guidance
only, as they may vary depending on the provenance of
the seeds. It should be said, however, that some of the
species included in this guide have seeds that are dif
ficult to germinate even when pretreated.
Useful information has been included about optimum
conditions for germinating the seeds, which can be
achieved by using incubators or cultivation chambers
where certain environmental factors can be controlled.
The optimum temperature is given; it may be continu
ous (20 ºC) or vary over a period of 24 hours (for ex
ample, 30/20 ºC). The seeds of some taxa germinate
well under different temperature conditions, which are
indicated as possible alternatives. In the case of the
varying temperatures, these are alternated: the lower
temperature is usually maintained for 16 hours and the
higher one for the remaining 8 hours of each 24 hour
period. The seeds of many species can germinate both
in the light and in the dark. Nevertheless, a photoperiod
of at least 8 hours a day is recommended, usually co
inciding with the higher temperature cycle in the case
of an alternating temperature treatment. In some
species, light stimulates germination, in which case this
requirement is expressly mentioned.
The data offered are only for guidance, as they may
vary greatly depending on the quality of handling,
cleanliness and storage conditions, as well as on the
specific characteristics of each batch of seeds, which
depend on the genotypes collected, the origin of the
seeds and the weather conditions each year.
Nursery production
Fact Sheet Contens 20
For mass producing nurserygrown plants, details are
given of the most appropriate sowing period and of
whether pretreatment is required to stimulate germi
nation. Guidelines are provided on suitable container
sizes and plant ages to obtain plants with a wellde
veloped root system that will withstand transplanting
and be capable of penetrating the soil rapidly after
planting. The containers recommended in the fact
sheets should have an antispiralling system. As regards
the large containers (3.5 litres), ones with a grid base
raised off the ground are recommended to help the
transplanted roots. Plant age is given as follows: 1/0 =
1 year, 2/0 = 2 years, 1/1 = 1 year in a 300 cm3 con
tainer + 1 year in a 3.5 l container. The use of plants
that are more than two years old is not recommended
and they should not be more than 150 cm high. The
approximate emergence time is indicated: it will vary
according to the batch, the nursery’s practices, the lo
cation of the nursery and the climate conditions that
year.
Some data for producing bareroot plants (sowing den
sity, dimensions) are provided for some species, al
though this traditional growing method has been
replaced by containergrowing, which makes it possi
ble to extend the plantingout period. The bareroot
plant dimensions given (stem circumference, total
height) are all maximum sizes.
Vegetative propagation
Information is provided about vegetative propaga
tion of the species using cuttings. This is the plant
production technique most commonly used for
restorations and forestations of Tamarix, Salix, Pop
ulus and some lianoid taxa. As plants of the other
species in this guide are usually produced from seed
rather than by vegetative means, most information
on their vegetative multiplication is experimental or
comes from the production of cultivars for ornamen
tal purposes.
The most suitable type of material the part of the stem
or set showing the best rooting ability, a suitable num
ber of internodes or cutting size and the best time of
the year to take the cuttings are indicated. The concen
tration of the watersoluble salt form of indolbutyric
acid (KAIB) in which to soak the cutting for 1 to 5 min
utes just before planting is given as a rough guide to a
starting point for finetuning a rooting protocol. Fol
lowing the indications of Mac Cárthaig and Spethmann
(2000), the species have been divided into four groups
according to the ease with which they form roots: those
that do not need treatment, those that only need it only
to accelerate the process (< 0.5 %), those that pose
medium difficulty (0.5 %) and those that are very diffi
cult to propagate (1 %).
For plant production by vegetative propagation, the
same kind of container as in the table for nursery pro
duction of plants from seed is recommended.
This guide does not aim to provide specific information
about plant production by micropropagation. This kind
of technique is relatively complex and costly and does
not seem justified when producing plants for hydro
logical restorations. However, bibliography on the sub
ject is provided for further reference.
References
To make the text easier to read, the basic bibliography
consulted systematically for describing species and infra
specific taxa and references to general works from which
propagation chart data have been taken have been
avoided in the text itself. All these references are men
tioned in the bibliography under the ‘General References’
heading. Other studies that have provided additional in
formation about different aspects, mostly published in
journals, are expressly cited in the text and are included
as a specific bibliography so that readers can go into the
subject in more depth if they wish.
References
Bolòs O, Vigo J (1989) Flora dels Països Catalans. Editorial Bar
cino, Barcelona
Boratyński A, Browicz K, Zieliński J (1992) Chorology of trees
and shrubs in Greece. Sorus, Poznań
Brummitt RK (2001) World Geographical Scheme for Record
ing Plant Distributions. Plant Taxonomic Database Standards
No. 2. Edition 2, August 2001. TDWG (online URL
http://www.tdwg.org)
Charco J (2001) Guía de los árboles y arbustos del norte de
África. Agencia Española de Cooperación Internacional,
Madrid
Flora Europaea. Royal Botanic Garden Edinburgh (online URL
http://rbgweb2.rbge.org.uk/FE/fe.html)
Germplasm Resources Information Network (GRIN) USDA
Agricultural Research Centre (online URL http://www.ars
grin.gov/)
Hulén E, Fries M (1986) Atlas of North European Vascular
Plants. North of the tropic of cancer. Koelz Scientific Books,
Königstein
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Pignatti S (1982) Flora d’Italia. Edagricole, Bolognia
Programa Anthos. Fundación Biodiversidad, Ministerio de
Medio Ambiente Real Jardín Botánico de Madrid, CSIC (on
line URL http://www.anthos.es/)
Zohari M (1972) Flora palaestina. Part two. Text. The Israel
Academy of Sciences and Humanities, Jerusalem
21 Fact Sheet Contens
Baum BR (1978) The genus Tamarix. The Israel Academy of
Sciences and Humanities; Jerusalem
Alnus glutinosa
(L.) Gaertn.
Distribution and Ecology
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus,
Western and Middle Asia, Siberia, Northern Africa
Mediterranean region: Portugal, España, Francia (incl.
Córcega), Italia (incl. Cerdeña, Sicilia), Croacia, Bosnia
y Herzegovina, Montenegro, Albania, Grecia, Turquía,
Libia, Túnez, Argelia, Marruecos
The alder prefers temperate to cool climates, al
though it can also withstand warmer temperatures
Diagnostic traits
Alnus glutinosa is a mediumsized deciduous tree, not
exceeding 25 m in height, with darkbrown fissured
bark. It can be distinguished from Alnus cordata, a na
tive of Albania, Corsica and Italy, by the shape of its
leaves. In the former these are obovate or suborbicu
lar, rarely elliptic, obtuse or retuse, and doubletoothed;
in the latter, they are suborbicular or cordate, generally
acute, and serrulate. Leaves are usually glabrous in
EN: black alder, common alder
EL: σκλήθρα, κλήθρα
ES: aliso, alno
FR: aulne glutineux, aulne noir
IT: ontano nero
PT: amieirocomum
Betulaceae
in conditions of adequate water intake. It grows on
clay, silty clay, sandy or alluvial materials and re
quires permanent moisture. Although it can be found
on substrata with varying pH levels, it prefers acid to
neutral soil types. The nodules of its roots, in sym
biosis with bacteria, are effective at fixing atmos
pheric nitrogen, allowing the tree to live on poor
terrain. It is found by watercourses and rivers, at the
bottom of valleys, in mixed deciduous forests, in
flooded areas and on wet slopes, either scattered or
in small populations.
both species, although tufts of hair may be present in
the nerve axils. The infructescence peduncle diameter
is smaller in A. glutinosa (0.5–1 mm) than in A. cordata
(2–3 mm). Alnus glutinosa also differs from Alnus in
cana, which is found in the centre, northeast and north
of Europe, in that in the latter the leaves are acumi
nate and pulverulent or tomentose, in young trees at
least, and the infructescences are sessile.
Alnus glutinosa 22
Sex expresion
■
monoecy
Flowering
pendulous male
catkins, erect female
catkins
■ from February to
June, before leaf
development
■
Pollination
■
■
anemophilous
selfincompatible
Two types of incompatibility systems seem to operate
in Alnus glutinosa: lowered fertilisation success of the
pollen parent against competition, when pollen of
Variation and Hybridisation
Alnus glutinosa can be pollinated by Alnus cordata.
However, only natural hybrids (A. × elliptica) have been
found in Corsica (Prat et al., 1992). It may often cross
breed with Alnus incana (A. × pubescens Tausch) in
areas where both species coexist. This hybrid exhibits
female catkins with short peduncles and leaves that
combine the features of both parent species.
The distribution pattern of the species, in relatively
small, isolated populations spread across an extensive
territorial range, is associated with highly significant
variations between provenances and individuals as re
gards quantitative features or characteristics of adap
tative relevance, (Weisgerber, 1974; DeWald and
Steiner, 1986; Krstinič, 1994; Baliuckas et al., 1999).
Geographical structuring of the genetic variation of the
species has been observed in studies using molecular
techniques (King and Ferris, 2000). At local level, a
relatively low rate of genetic variation has been esti
mated within populations, due to inbreeding (Kajba and
Gračan, 2003), which may have been encouraged by
the fact that sprouting from stumps occurs easily in
Fruiting
Ripening
■
■
black, lignified
infructescence,
persistent after
dehiscence
■ 1025 x 712 mm
from September
to November
■ dispersal by wind
other individuals is present, or the inability to fertilise
ovules when there is no competition (Steiner and Gre
gorius, 1999) as may happen in isolated trees.
the alder, especially in younger trees. In studies with
higher levels of intrapopulational diversity, Gömöry and
Paule (2002) established a spatial genetic pattern in
alder populations, probably due to limited seed disper
sal, which causes offspring to take root near their
progenitors.
Geographical structuring of genetic variation in this
species makes it advisable to use local populations as
a source of reproductive materials in restoration
projects. It is also advisable to promote genetic varia
tion in new populations by collecting material from a
great many individuals at different points within the
same region of provenance (Kajba and Gračan, 2003)
and to ensure that the material is taken from trees that
are quite distant from each other.
The extensive genetic variation that has been found for
features of productive interest is used to set up im
provement programmes, selecting superior genotypes
for these traits to create seed orchards (Krstinič and
Kajba, 1991).
23 Alnus glutinosa
Reproductive biology
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from September to
November
■ gathering by hand,
climbing or using
longhandled tools
sequence for dehis
cent fruits
■ seed weight / kg
fruit: 30260 g
■ purity: 4190 %
Black alder produces seeds every year, but good crops
occur once every 23 years. Climatic conditions
around the time of flowering initiation, in the sum
mer of the previous year and during the following
spring greatly affect fruiting (Suszka et al., 1994).
Collecting is carried out when the earliest fruits start
to open. Seeds collected before the fruits turn brown
12 g
require several months of afterripening to germinate
(McVean, 1953). The light weight of alder seeds
makes it difficult to remove empty seeds. If a kiln is
used to open the fruits, it is recommended that tem
peratures should not exceed 35 ºC; above this thres
hold seeds lose their viability.
Germination under controlled conditions
Pregermination treatments
■
■
fresh: without treatment
dried: prechilling (38 weeks)
Conditions
■
T: 5 ºC to 4 ºC
MC: 48 %
■ airtight container
■
30 / 20 ºC; 25 ºC
Germination Viability
■
3070 %
Seed lot quality and germination capacity are often
very low, as it is difficult to separate sound from empty
seeds.
Nursery production
Sowing season
■
autumn or early spring, without
treatment, or spring, with treatment
Nursery practice
bareroot: 1020 g/m2;
circumference up to 46 cm or
total height up to 100150 cm
3
■ forestpot 300 cm : 1/0
■ container 3.5 l: 1/1
■
Production in forestpot or container allows seedlings to
be inoculated with the actinobacteria Frankia and assu
Emergence
■
the first spring, complete within
35 weeks
res their nodulation prior to planting (Berry and Torrey,
1985) giving rise to plants with better development.
Alnus glutinosa 24
Type of cutting
■
■
hardwood mallet
semihardwood
Position along
the shoot
basal
terminal
Number of
internodes Size
25 cm
10 cm
It is advisable to apply rejuvenation treatments in
order to increase the percentage of rooting if adult
ortets are used, since results are highly conditioned
by the age of the parent plant (Krstinič, 1994; Martin
and Guillot, 1982; Psota, 1987). There is also great
clonal variation in rooting capacity (Good et al.,
1978). Kruger (1982) obtained high percentages of
rooting and survival and high quality root formation
using hardwood mallet cuttings. Cuttings are nor
mally reproduced in forestpots, in a mist environ
ment (Martin and Guillot, 1982).
References
Collecting time
winter
summer
Auxin
concentration
0.5 %
0.5 1 %
The capacity to produce shoots from the root, and,
therefore, the possibility of reproducing this species by
means of root cuttings has not been completely proved.
Some authors (McVean, 1953; Krstinič, 1994) suggest
that the alder has this capacity, although infrequently,
while Fayle (1996) questions this behaviour since it has
not been confirmed in the field.
There are several references to in vitro propagation tri
als that have achieved good results. (Garton et al.,
1981; Lall et al., 2005; Perinet and Tremblay, 1987;
Simon et al.,1985; Vergnaud et al., 1987).
General references
Specific references
Ball PW (1993) Alnus Miller. In: Tutin TG et al. (eds). Flora Eu
ropaea. Vol 1. 2nd edn. Cambridge University Press, Cambridge
Baulickas V, Ekberg I, Eriksson G, Norell L (1999) Genetic vari
ation among and within populations of four Swedish hard
wood species assessed in a nursery trial. Silvae Genetica
48:1725
Catalán G (1991) Semillas de árboles y arbustos forestales.
Ministerio de Agricultura Pesca y Alimentación. ICONA,
Madrid
Christensen KI (1997) Alnus Miller. In: Strid A, Tan K, (eds).
Flora Hellenica. Vol 1. Koeltz Scientific Books, Königstein
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Nicolás JL, Iglesias S, Alía R (2001) Fichas descriptivas de es
pecies. In: García del Barrio JM et al. (coord). Regiones de
identificación y utilización de material forestal de reproduc
ción. Ministerio de Medio Ambiente, Madrid
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea ANPA, Roma
Rocha Alfonso ML (1990) Alnus Miller. In: Castroviejo S et al.
(eds). Flora Ibérica. Vol 2. CSIC, Madrid
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Berry AM, Torrey JG (1985) Seed germination, seedling inoc
ulation and establishment of Alnus spp. in containers in
greenhouse trials. Plant and Soil 87:161173
DeWald LE, Steiner KC (1986) Phenology, height increment
and cold tolerance of Alnus glutinosa populations in a com
mon environment. Silvae Genetica 35:205211
Fayle DCF (1996) Sugar maple, black spruce and tamarack do
not reproduce vegetatively from roots. Forestry Chronicle
72:283285
Garton S, Hosier MA, Read PE, Farnham RS (1981) In vitro
propagation of Alnus glutinosa Gaertn. HortScience 16:758
759
Gömöry D, Paule L (2002) Spatial and microgeographical ge
netic differentiation of black alder (Alnus glutinosa Gaertn.)
populations. Forest Ecology and Management 160:3–9
Good JE, Bellis JA, Munro RC (1978) Clonal variation in root
ing of softwood cuttings of woody perennials occurring nat
urally on derelict land. International Plant Propagators Society
Combined Proceedings 28:192201
25 Alnus glutinosa
Vegetative propagation
Kajba D, Gračan J (2003) EUFORGEN Technical Guidelines for
genetic conservation and use for black alder (Alnus glutinosa).
International Plant Genetic Resources Institute, Rome
Prat D, Leger C, Bojovic S (1992) Genetic diversity among
Alnus glutinosa (L.) Gaertn. populations. Acta Oecologica
13:469477
King RA, Ferris C (2000) Chloroplast DNA and nuclear DNA
variation in the sympatric alder species, Alnus cordata (Lois.)
Duby and A. glutinosa (L.) Gaertn. Biological Journal of the
Linnean Society 70:147160
Psota V (1987) Rhizogenesis of stem cuttings in Alnus gluti
nosa (L.) Gaertn. and Quercus robur L. species as related to
dormancy and plant growth regulators. Acta Universitatis
Agriculturae, Facultas Agronomica 35:2744
Krstinič A (1994) Genetics of black alder (Alnus glutinosa (L.)
Gaertn.). Annales Forestales 19:3372
Krstinič A, Kajba D (1991) Possibilities of genetic gain for vig
orous growth of black alder (Alnus glutinosa (L.) Gaertn) by
clonal seed orchard. Sum. list 69:261271
Kruger H (1982) Vegetative vermehrung von Nadel und Laub
hölzen. Allgemeine Forstzeitschrift 910:243244
Lall S, Mandegaran Z, Roberts AV (2005) Shoot multiplication
in cultures of mature Alnus glutinosa. Plant Cell Tissue and
Organ Culture 83:347350
Martin B, Guillot J (1982) Quelques essais de bouturage de
l’aulne. Revue Forestière Francaise 34:381391
McVean DN (1953) Alnus glutinosa (L.) Gaertn. Journal of
Ecology 43:447466
Périnet P, Tremblay FM (1987) Commercial micropropagation
of five Alnus species. New Forests 3:225230
Simon L, Stain A, Côte S, Lalonde M (1985) Performance of in
vitro propagated Alnus glutinosa (L.) Gaertn. clones inocula
ted with Frankiae. Plant and Soil 87:125133
Steiner W, Gregorius HR (1999) Incompatibility and pollen
competition in Alnus glutinosa: evidence from pollination ex
periments. Genetica 105:259271
Suszka B, Muller C, BonnetMasimbert M (1994) Graines des
feuillus forestiers, de la recolte au semis. INRA, Paris
Vergnaud L, Chaboud A, Rougier M (1987) Preliminary analy
sis of root exudates of in vitromicropropagated Alnus gluti
nosa clones. Physiologia Plantarum 70:319326
Weisgerber H (1974) First results of progeny test with Alnus
glutinosa Gaertn. after controlled pollination. In: Proceedings
of the Join IUFOR Meeting, SO2.04.13. Session VI, Stockholm
Alnus glutinosa 26
Distribution and Ecology
General distribution: Southwestern and Southeastern
Europe, Western Asia, Northern Africa
Mediterranean region: Portugal, Spain (incl. Baleares),
France (incl. Corse), Italy (incl. Sardegna and Sicilia),
Croatia, BosniaHerzegovina, Albania, Greece (incl.
Kriti), Cyprus, Turkey, Lebanon, Tunisia, Algeria,
Morocco
This typical Mediterranean species prefers rather cool
soil and does not weather prolonged intense cold too
well. In the lower and warmer areas of its distribution
Diagnostic traits
Arbutus unedo is a perennial shrub or small tree up to
4 – 7 m height, with alternate, lanceolate leaves. In the
eastern area of its range of distribution it sometimes
shares territory with Arbutus andrachne, from which it
differs in that its bark is fissured, brownish in colour
Reproductive biology
Sex expression
■
hermaphroditism
Arbutus
unedo L.
EN: strawberry tree
EL: κουμαριά
ES: madroño
FR: arbousier
IT: corbezzolo
PT: medronheiro
Flowering
white flowers,
clustered in panicles
■ from October to
December
■
Pollination
■
range it can be found in shaded spots. It is more abun
dant on siliceous soils, but can also be found on cal
careous substrates. It resprouts well from the stump
after fire or cutting.
The strawberry tree grows in Quercus and Pinus forests
or among high shrubs together with other typical
species of the Mediterranean maquis. Although not
specifically a waterside species, its need for somewhat
cool soils and its interaction with animals make it
worth considering for use in areas of transition be
tween riparian and zonal vegetation.
and peels off in small flakes, its young twigs are often
glandulosesetose and it blooms in the autumn. In con
trast, the bark of A. andrachne is brilliant orangered in
colour, smooth and peels off in papery sheets, and it
blooms in the spring.
entomophilous
Fruiting
Ripening
■
■
globose red or
orange berry,
glandulose
■ 2025 mm
from October to
December
■ dispersal by
frugivorous
vertebrates
27 Arbutus unedo
Ericaceae
Variation and Hybridisation
Hybridisation of A. unedo and A. andrachne (A. × an
drachnoides Lint.) can often occur in areas where both
species coexist. The result is a fertile hybrid exhibiting
the brightlycoloured bark of A. andrachne and some
glandulose hairs on its young twigs. Also recognised is
Arbutus × androsterilis Salas, Acebes & Arco, a hybrid
of A. unedo and A. canariensis (Salas Pascual et al.,
1993) resulting from artificial sympatry.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from October
to December
■ gathering by hand
sequence for fleshy
fruits
■ seed weight / kg
fruit: 614 g
■ purity: 7097 %
The fruit production of strawberry trees varies greatly
from one year to the next (Herrera, 1988), since seed
23 g
quantity and quality are affected by the intensity and du
ration of summer drought periods (Chiarucci et al., 1993).
Germination under controlled conditions
Pregermination treatments
■
prechilling (412 weeks)
Conditions
■
T: 4 ºC
MC: 48 %
■ airtight container
■
15 to 20 ºC
Germination Viability
■
8099 %
Strawberry tree seeds have no problem in germinating,
but emergence is accelerated and made more homo
geneous by prechilling.
Nursery production
Sowing season
■
autumn, without treatment, or
spring, with treatment
Nursery practice
■
■
forestpot 300 cm3: 1/0
container 3.5 l: 1/1
Strawberry tree seedlings are very sensitive; care must
be taken to avoid the effects of frost and excessive
heat.
Emergence
■
the first spring, complete within
34 weeks
Arbutus unedo 28
Type of cutting
■
semihardwood
Position along
the shoot
terminal
Number
of internodes Size
2
The results obtained through vegetative propagation of
strawberry trees are very irregular. It has not been pos
sible to obtain rooting percentages higher than 50%
(Crobeddu and Pignatti, 2005; Pignatti and Crobeddu,
2005). According to Pignatti and Crobeddu (2005), it is
necessary to use material obtained from young parent
plants which have been repeatedly pruned to stimulate
vigorous sprouting. If this type of material is not used,
References
General references
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
GarcíaFayos P (coord) (2001) Bases ecológicas para la reco
lección, almacenamiento y germinación de semillas de espe
cies de uso forestal en la Comunidad Valenciana. Banc de
Llavors Forestals, Generalitat Valenciana, Valencia
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Nicolás JL, Iglesias S, Alía R (2001) Fichas descriptivas de es
pecies. In: García del Barrio JM et al. (coord). Regiones de
identificación y utilización de material forestal de reproduc
ción. Ministerio de Medio Ambiente, Madrid
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di
alberi e arbusti della flora mediterranea. ANPA, Roma
Villar L (1996) Arbutus L. In: Castroviejo S. et al. (eds). Flora
Ibérica. Vol 4. CSIC, Madrid
Webb DA (1972) Arbutus L. In: Tutin TG et al. (eds). Flora Eu
ropaea. Vol 3. Cambridge University Press, Cambridge
Specific references
Cervelli C (2005) Le specie arbustive della macchia medite
rranea. Un patrimonio a valorizzare. Collana Sicilia Foreste
26:39154
Chiarucci A, Pacini E, Loppi S (1993) Influence of temperature
and rainfall on fruit and seed production of Arbutus unedo L.
Botanical Journal of the Linnean Society 111:7182
Collecting time
june
Auxin
concentration
1%
the rooting probability seems to be nil. The best time to
harvest cuttings is at the end of the growth period.
After this time, the rootforming capacity diminishes
by 1020% (Cervelli, 2005).
The strawberry tree has been successfully propagated
by in vitro culture (Giordani et al., 2005; Mereti et al.,
2002; Morini y Fiaschi, 2000; Rodrigues et al., 2001).
Crobeddu S, Pignatti G (2005) Propagazione per talea di specie
mediterranee. Prove di substrato. Sherwood Foreste ed Alberi
Oggi 114:2731
Giordani E, Benelli C, Perria R, Bellini E (2005) In vitro germi
nation of strawberry tree (Arbutus unedo L.) genotypes: es
tablishment, proliferation, rooting and callus induction.
Advances in Horticultural Science 19:216220
Herrera CM (1998) Longterm dynamics of Mediterranean fru
givorous birds and fleshy fruits: a 12yr study. Ecological
Monographs 68:511538
Mereti M, Grigoriadou K, Nanos GD (2002) Micropropagation
of the strawberry tree, Arbutus unedo L. Scientia Horticulturae
93:143148
Morini S, Fiaschi G (2000) In vitro propagation of strawberry
tree. Agricoltura Mediterranea 130:240246
Pignatti G, Crobeddu S (2005) Effects of rejuvenation on cut
ting propagation of Mediterranean shrub species. Foresta
2:290295 (online URL http://www.sisef.it/)
Rodrigues AP, Sergio.PM, Teixeira MR, Pais MS (2001) In vitro
break of dormancy of axillary buds from woody species (Persea
indica and Arbutus unedo) by sectioning with a laser beam.
Plant Science 161:173178
Salas Pascual M, Aceves Ginovés JR, del Arco Aguilar M (1993)
Arbutus x androsterilis, a new interespecific hybrid between
A. canariensis and A. unedo from the Canary Islands. Taxon
42:789792
29 Arbutus unedo
Vegetative propagation
Celtis
australis L.
Distribution and Ecology
General distribution: Southwestern and Southeastern
Europe, Western Asia, Northern Africa
Mediterranean region: Portugal, Spain (incl. Baleares),
France (incl. Corse), Italy (incl. Sardegna and Sicilia),
Croatia, BosniaHerzegovina, Montenegro, Albania,
Greece, Cyprus, Turkey, Syria, Lebanon, Tunisia, Algeria,
Morocco
The European hackberry has been grown widely in the
Mediterranean basin since ancient times because of its
many uses, making it difficult to establish the precise
limits of its natural distribution range. It is often asso
Diagnostic traits
This species is a deciduous tree that can grow to a
height of 30 m. Its bark is smooth and greyish
coloured. Its leaves are sharply serrate, rounded or cor
date at the base and usually two to three times as long
as broad. The fruits are globose, with a strongly retic
ulaterugose endocarp.
It differs from Celtis tournefortii, which is distributed
throughout the south of Europe from Sicily to the
EN: European hackberry,
European nettletree
EL: μελικουκιά
ES: almez
FR: micocoulier
IT: bagolaro
PT: lódãobastardo
Ulmaceae
ciated with farming and stockbreeding and may be
found near rural buildings, alongside ditches and on
the edges of fields.
Spontaneous distribution occurs in the form of scat
tered individuals, in small pure stands or mixed with
other broadleaves species. The hackberry is found in
forests and ravines and on rocky, shady slopes, in semi
arid and subhumid Mediterranean environments. It
prefers cool, loose, rocky soils and is indifferent to sub
strate. It resprouts from the stump and the root after
cutting or fire.
Crimea, in that the latter is a bush or small tree not
usually exceeding 6 m in height, with leaves that are
less than twice as long as broad, and its endocarp has
four ridges. Morphological differences as regards Celtis
caucasica, which is found in Bulgaria, Yugoslavia and
western Asia, are less obvious, since C. caucasica also
grows into a tree, although its leaves are cuneate at
the base and its fruit is yellowish brown when ripe.
Celtis australis 30
Reproductive biology
Sex expression
■
andromonoecy
Flowering
Pollination
flowers small and
inconspicuous,
generally solitary
■ from March to May
■
■
anemophilous
Fruiting
Ripening
■
■
brownishblack,
spherical drupe
■ 812 mm
from October,
may remain until
late winter on
the tree
■ dispersal by
frugivorous
vertebrates
Variation and Hybridisation
There is no information on intraspecific variation and
hybridisation for this taxon.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from November to
winter
■ climbing, or using
longhandled tools,
or beating branches
sequence for fleshy
fruits
■ seed weight / kg
fruit: 320400 g
■ purity: 95100 %
100260 g
Germination under controlled conditions
Pregermination treatments
■
prechilling (8–12 weeks)
Conditions
■
20 / 10 ºC
T: 4 ºC
MC: 48 %
■ airtight container
■
Germination Viability
■
4096 %
European hackberry has dormancy that requires
prechilling.
Sowing season
■
autumn, without treatment, or
spring, with treatment
Nursery practice
bareroot: circumference up
to 46 cm or total height up
to 100150 cm
3
■ forestpot 300 cm : 1/0
■ container 3.5 l: 1/1
■
Emergence
■
the first spring
31 Celtis australis
Nursery production
Vegetative propagation
Type of cutting
■
■
hardwood
semihardwood
Position along
the shoot
basal or middle
basal or middle
Number of
internodes Size
20 cm
10 cm
It is advisable to perform rejuvenation pruning on the
parent plants (Butola and Uniyal, 2005; Puri and
Shamet, 1988). Treatment with a high concentration
of auxins is indispensable to guarantee results of above
References
General references
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
GarcíaFayos P (coord) (2001) Bases ecológicas para la reco
lección, almacenamiento y germinación de semillas de espe
cies de uso forestal en la Comunidad Valenciana. Banc de
Llavors Forestals, Generalitat Valenciana, Valencia
Navarro C, Castroviejo S (1993) Celtis L. In: Castroviejo S et al.
(eds). Flora Ibérica. Vol 3. CSIC, Madrid
Nicolás JL, Iglesias S, Alía R (2001) Fichas descriptivas de es
pecies. In: García del Barrio JM et al. (coord). Regiones de
identificación y utilización de material forestal de reproduc
ción. Ministerio de Medio Ambiente, Madrid
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di
alberi e arbusti della flora mediterranea. ANPA, Roma
Collecting time
January to March
July
Auxin
concentration
0.5 1%
0.5 1%
50% (Shamet and Naveen, 2005). However, according
to trials carried out by Puri and Shamet (1988), the
concentration of indolebutyric acid could be reduced
to 0.01% if the treatment time is increased to 24 hours.
Tutin TG (1993) Celtis L. In: Tutin TG et al. (eds). Flora Euro
paea. Vol 1. 2nd edn. Cambridge University Press, Cambridge
Specific references
Butola BS, Uniyal AK (2005) Rooting response of branch cut
tings of Celtis australis L. to hormonal application. Forests,
Trees and Livelihoods 15:307310
Puri S, Shamet GS (1988) Rooting of stem cuttings of some
social forestry species. International Tree Crops Journal
5:6369
Shamet GS, Naveen CR (2005) Study of rooting in stem cut
tings of Khirk (Celtis australis Linn.). Indian Journal of Forestry
28:363369
Celtis australis 32
Distribution and Ecology
General distribution: Southwestern and Southeastern
Europe, Western Asia
Mediterranean region: France, Italy (incl. Sicilia), Croa
tia, BosniaHerzegovina, Montenegro, Albania, Greece
(incl. Kriti), Turkey, Syria, Lebanon, Palestine
Diagnostic traits
C. siliquastrum is a deciduous tree, 5 to 10 m in
height, with smooth bark. Its leaves are simple, alter
nate, orbicular or reniform, with a rounded apex. Its
flowers are pink or purplish and sprout directly from
the trunk and branches. C. siliquastrum may be easily
Reproductive biology
Sex expression
■
hermaphroditism
Cercis
siliquastrum L.
EN: Judas tree, lovetree
EL: κουτσουπιά
ES: árbol del amor, árbol de Judas
FR: arbre de Judée, gainier
IT: albero di Giuda, siliquastro
PT: olaia, árvore de Judas
Flowering
pale to deep pink
flowers, clustered in
racemes that growth
directly from the
stems and trunk
■ from March to May,
before leaf
development
■
Pollination
■
■
Lovetree is normally found on arid slopes or on the
banks of rivers, on calcareous substrates, although it
also tolerates moderately acid soils. It cannot with
stand prolonged periods of cold.
confused with many other Cercis varieties that are
used for ornamental purposes. Two of these are
C. canadensis, which has leaves with a pointed apex,
and C. chinensis, whose leaves are deeply acuminate
at the base.
entomophilous
selfcompatible
Analysis of storage proteins in seed sets of individual
trees indicates that the love tree is almost completely
Fruiting
Ripening
■
■
reddish to dark
brown legume
■ 60100 mm long
in July, remaining
many months on
the tree
■ dispersal by
gravity
autogamous, with less than 5% of crossfertilisation
(González and HenriquesGil, 1994).
33 Cercis siliquastrum
Fabaceae
Variation and Hybridisation
The subspecies C. siliquastrum subsp. hebecarpa
(Bornm.) Yalt., distributed in Asia Minor and Iran, has
nonglabrous calyces, pedicels and legumes.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
from late summer
gathering by hand or
beating branches
sequence for
dehiscent fruits
■ seed weight / kg
fruit: 300450 g
■ purity: 9598 %
Legumes can be collected any time after they turn dark
and the seeds turn brown. Although they remain closed
on the tree during most of the winter, early collection
2035 g
is prudent to minimize seed losses that might occur
from insect infestation.
Germination under controlled conditions
Pregermination treatments
mechanical scarification
mechanical scarification + prechilling
(412 weeks)
■ soaking in boiling water (1 minute)
■ soaking in water at 80 ºC and allowing
to cool for 24 h
■ scarification with concentrated
sulphuric acid (3060 minutes)
■
Conditions
■
■
Nursery production
Sowing season
■
spring, with treatment
Germination Viability
30 / 20 ºC
Love tree seeds experience dormancy due to the en
dosperm and to their impermeable coat (Riggio
Bevilacqua et al., 1985, 1988) and require scarification
and prechilling in order to germinate. The duration of
acid scarification should be determined by means of
tests for each seed lot. The application of gibberellic
■
7090 %
acid can break dormancy in previously soaked seeds,
but stratification for 16 weeks at 4 ºC is much more
efficient (Gebre and Karam, 2004). The use of acid may
also have a negative effect on the subsequent devel
opment of the seedling (Rascio et al., 1998).
Nursery practice
Cercis siliquastrum 34
bareroot; circumference up
to 46 cm or total height up
to 100150 cm
3
■ forestpot 300 cm : 1/0
■ container 3.5 l: 1/1
■
T: 4 ºC
MC: 48 %
■ airtight container
■
Emergence
■
the same spring, 24 weeks after
sowing
Type of cutting
■
semihardwood
Position along
the shoot
terminal
Number of
internodes Size
23
Lovetree is not easily reproduced from cuttings. Consi
derable variation in rooting ability depending on the
part of the shoot from which the cutting is taken and
the collection time has been observed (Karam and
Gebre, 2004).
References
General references
Ball PW (1968) Cercis L. In: Tutin TG et al. (eds). Flora Euro
paea. Vol 2. Cambridge University Press, Cambridge
Catalán G (1991) Semillas de árboles y arbustos forestales.
Ministerio de Agricultura Pesca y Alimentación. ICONA,
Madrid
Chamberlain DF, Yaltirik F (1970) Cercis L. In: Davis PH (ed).
Flora of Turkey and the Eastern Aegean Islands. Vol 3. Univer
sity Press, Edinburgh
Mac Cárthaigh D, Spethmann (eds) W (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea. ANPA, Roma
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Specific references
Bignami C (1984) Prove di micropropagazione di Cercis sili
quastrum L.. Informatore Agrario 40:103105
Collecting time
summer (July)
Auxin
concentration
1%
Micropropagation of lovetree has been tried with
some success using axillary buds (Bignami, 1984).
Gebre GH, Karam NS (2004) Germination of Cercis siliquas
trum seeds in response to gibberellic acid and stratification.
Seed Science and Technology 32:255260
González C, HenriquesGil N (1994) Genetics of seed storage
proteins in the love tree Cercis siliquastrum L. (Fabaceae). The
oretical and Applied Genetics 89:895899
Karam NS, Gebre GH (2004) Rotting of Cercis siliquastrum
cuttings influenced by cutting position on the branch and in
dolebutyric acid. Journal of Horticultural Science and
Biotechnology 79:792796
Rascio N, Mariani P, Dalla Vecchia F, La Rocca N, Profumo P
Gastaldo P (1998) Effects of seed chilling or GA3 supply on
dormancy breaking and plantlet growth in Cercis siliquastrum
L. Plant Growth Regulation 25:53–61
Riggio Bevilacqua L, Roti Michelozzi G, Serrato Valenti G
(1985) Barriers to water penetration in Cercis siliquastrum
seeds. Seed Science and Technology 13:175182
Riggio Bevilacqua L, Profumo P, Gastaldo P, Barella P (1988)
Cytochemical study on the dormancyimposing endosperm of
Cercis siliquastrum. Annals of Botany 61:561565
35 Cercis siliquastrum
Vegetative propagation
Ranunculaceae
Clematis flammula L.
Clematis vitalba L.
Clematis vitalba L.
Clematis flammula L.
EN: clematis, traveller’sjoy
EL: κληματίς
ES: clemátide
FR: clématite
IT: clematide
PT: clematis
Distribution and Ecology
C. vitalba:
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus, West
ern Asia, Northern Africa
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia,
BosniaHerzegovina, Montenegro, Albania, Greece,
Turkey, Syria, Lebanon, Algeria
C. flammula:
General distribution: Southwestern and Southeastern
Europe, Caucasus, Western Asia, Northern Africa
Mediterranean countries: Portugal, Spain (incl. Balea
res), France (incl. Corse), Italy (incl. Sardegna, Sicilia),
Croatia, BosniaHerzegovina, Montenegro, Albania,
Greece (incl. Kriti), Turkey, Cyprus, Syria, Lebanon, Is
rael, Libya, Tunisia, Algeria, Morocco
C. vitalba. C. flammula 36
Clematis vitalba and C. flammula may be among the
climbers that make up the riparian vegetation. C. vi
talba requires greater amounts of moisture and is com
mon in broadleaved forests and deciduous spiny
thickets in Eurosiberian environments, although it is
also found in shady and cool areas in the Mediter
ranean region. C. flammula likes warmer weather and
is restricted to the Mediterranean coast; it is also found
in hedges, thickets and forests, on sunny open ground.
Diagnostic traits
Clematis vitalba and C. flammula are perennial lianas
with woody stems, at least in their lower parts. They can
be distinguished because the former has 1pinnatisect
leaves with ovate, cordateovate or ovallanceolate
leaflets, while the leaves of the latter are mostly bipin
natisect, sometimes tripinnatisect, with ovate, lanceolate
or linear leaflets. The perianth segments in C. flammula
are white and glabrous on the inside; in C. vitalba, they
are greenish white and pubescent on both sides.
Reproductive biology
Sex expression
■
hermaphroditism
Flowering
flowers clustered in
paniculiform cymes
■ from May to August,
occasionally later
■
In addition to these two species, other climbing Clema
tis varieties are found less frequently in the Mediter
ranean region: C. viticella L., with purplish flowers, and
C. campaniflora Brot., which is limited to the centre
and the west of the Iberian Peninsula and has solitary
or two to four clustered flowers and bracteoles that are
welded together to form an involucre below the flower.
Pollination
■
entomophilous
Fruiting
Ripening
■
■
achene with a
persistent feathery
plume
■ 23 mm (plume
length: up to 3.5 cm
C. flammula; up to
5.5 cm C. vitalba)
from September
to November
■ dispersal by wind
Variation and Hybridisation
There is no information on intraspecific variation and
hybridisation for these taxa.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from October
to December
■ gathering by hand
friction for feathery
style removal
■ purity: 99100 %
■
57 g C. flammula (achenes)
13 g C. vitalba (achenes)
T: 4 ºC
MC: 48 %
■ airtight container
■
37 C. vitalba. C. flammula
Seeds are not usually extracted from the achenes.
Germination under controlled conditions
Pregermination treatments
■
manual removal of seeds from fruits or
mechanical scarification + prechilling
(8–24 weeks)
Conditions
■
20 / 10 ºC; 20 ºC
Clematis seeds have morphophysiological dormancy
and prechilling is required for germination. Cold strat
ification for 8 to 12 weeks seems to be adequate for
the germination of Clematis vitalba seeds (Bungard et
Nursery production
Sowing season
■
autumn or early spring, without
treatment, or spring, with treatment
Vegetative propagation
Type of cutting
■
semihardwood
or softwood
Position along
the shoot
indifferent
Germination Viability
■
al., 1997). Prechilling can be replaced by applying al
ternating temperatures of 5 ºC for 12 hours and 15 ºC
for 12 hours in an incubator (Vinkler et al., 2004).
Nursery practice
■
forestpot 300 cm3: 1/0 or 2/0
Number of
internodes Size
12
Propagation of the genus Clematis is carried out in
summer using shoots that have grown in the spring of
the same year. Use of softwood cuttings with a pair of
leaves is recommended. Münster (2000) recommends
making a cut in the lower 2 cm of the cutting, leaving
the cambium visible, to accelerate the formation of
roots. One of the leaves may be removed to save space
and to avoid fungal infections by the genus Botrytis.
Another form of propagation is the socalled ‘Japan
ese method’, which also uses cuttings with a node but
somewhat longer and more vigorous, since the cuts are
made in the internodes immediately above and below
6595 %
Emergence
■
in the first spring, may be
complete in the following
autumn
Collecting time
summer
Auxin
concentration
without or < 0.5 %
the node. In this method, the probability of fungi
reaching the buds is much lower than in the conven
tional method. However, it has the disadvantages of
occupying more space and providing less material from
each parent plant (Gunn, 2005). Kreen et al. (2002) rec
ommends using perlite as a substrate and rooting in a
mist environment.
Rooting microcuttings obtained by in vitro propaga
tion seems to be more effective than the traditional
method using softwood cuttings (Kreen et al., 2002).
C. vitalba. C. flammula 38
General references
Fernández Carvajal MC (1986) Clematis L. In: Castroviejo S
et al. (eds). Flora Ibérica. Vol 1. CSIC, Madrid.
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Strid A (1967) Clematis L. In: Stris A, Tan K (eds). Flora Hel
lenica. Vol 2. ARG Gantner Verlag KG, Ruggell
Tutin TG and Akeroyd JR (1993) Clematis L. In: Tutin TG et al.
(eds). Flora Europaea. Vol 1. 2nd edn. Cambridge University
Press, Cambridge
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Specific references
Bungard RA, Daly GT, McNeil DL, Jones AV, Morton JD (1997)
Clematis vitalba in a New Zealand native forest remnant: does
seed germination explain distribution? New Zealand Journal of
Botany 35:525534
Gunn S (2005) Clematis from cuttings. Plantsman 4:8183
Kreen S, Svensson M, Rumpunen K (2002) Rooting of Clema
tis microshoots and stem cuttings in different substrates. Sci
entia Horticulturae 96:351357
Münster K (2000) Clematis. In: Mac Cárthaigh D, Spethmann
W (eds). Krüssmans Gehölzvermehrung. Parey/Blackwell Wis
senschaftsverlag, Berlin
Vinkler I, Muller C, Gama A (2004) Germination de la Clé
matite (Clematis vitalba L.) et perspectives de maîtrese
préventive au forêt. Revue Forestière Française 56:275286
39 C. vitalba. C. flammula
References
Coriaria
myrtifolia L.
μυρτόφυλλος, κοριάρια η
μυρτόφυλλη
ES: emborrachacabras, garapalo
FR: corroyère, redoul
IT: coriaria, sommacco provenzale
PT: coriaria
Distribution and Ecology
General distribution: Southwestern and Southeastern
Europe, Northern Africa
Mediterranean region: Spain (incl. Baleares), France,
Italy, Algeria, Morocco
Coriaria myrtifolia is a species that requires moderately
moist soils. It is indifferent to the nature of the sub
Diagnostic traits
This species is a semideciduous bush that reaches a
height of 1 to 2 m, with simple, entire, opposite leaves.
Reproductive biology
Sex expression
■
andromonoecy
Flowering
greenish flowers,
clustered in raceme
■ from March to June
■
Coriariaceae
EN: coriaria
EL: βυρσοδεψική η
Pollination
■
■
strate and grows in partial shade or under full sunlight.
It is found in the Mediterranean region, from the coast
to the mountain areas, in riversides, ravines, bramble
patches and moist hedges. It has a strong root system
that acts in symbiosis with bacteria to fix atmospheric
nitrogen, allowing it to vegetate on nutrientpoor soil.
Its fruit is very striking, both in shape and colour, but
highly toxic to humans.
anemophilous
selfcompatible
Coriaria myrtifolia 40
In spite of the selfcompatibility of the species, cross
fertilisation between individuals is enhanced by the
fact that male flowers appear before hermaphrodite
flowers in each single plant (Thompson et al., 1995).
Fruiting
Ripening
■
■
black achene,
surrounded by
keeled, fleshy pieces,
initially red, black
when ripe
■ about 4 mm
from July to
September
■ dispersion by
frugivorous
vertebrates
Thus it would seem that the fruit collected from a sin
gle mother plant would tend to result from pollination
by different individuals.
Variation and Hybridisation
Coriaria is the only genus of the Coriariaceae family,
which has a conspicuously disjunct world distribution
(Yokoyama et al., 2000). C. myrtifolia is the only species
that is present in Europe.
Seed propagation
Seed handling and storage
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from late summer to
early autumn
■ gathering by hand
sequence for fleshy
fruits
■ seed weight / kg
fruit: 1034 g
■ purity: 99100 %
1113 g
Germination under controlled conditions
Pregermination treatments
■
mechanical scarification + immersion
in a 500 ppm gibberellic acid solution
(4 days) + prechilling (4 weeks)
Nursery production
Sowing season
■
spring, with treatment
Conditions
■
■
Germination Viability
25 / 20 ºC
light
■
Nursery practice
■
■
forestpot 300 cm3: 1/0
container 3.5 l: 1/1
For mass production in a nursery, pretreatment could
be simplified to mechanical scarification and sowing
in autumn or in early spring, although emergence could
T: 4 ºC
MC: 48 %
■ airtight container
■
8099 %
Emergence
■
the same spring, complete
within one to two months
be slow. Inoculation with nitrogenfixing microor
ganisms noticeably improves the development of the
plants (Martinez et al., 1997; Cañizo et al., 1978).
41 Coriaria myrtifolia
Tolerance to desiccation: ORTHODOX
Vegetative propagation
Type of cutting
■
semihardwood
Position along
the shoot
basal or middle
Number of
internodes Size
10 15 cm
Vegetative propagation of Coriaria myrtifolia offers
better results when using softwood cuttings harvested
during vegetative rest (autumnwinter). However, root
ing of Coriaria myrtifolia cuttings in autumnwinter
must be done in a protected environment, maintaining
a temperature of 20 ºC (Melgares de Aguilar et al.,
References
General references
Webb DA (1968) Coriaria L. In: Tutin TG et al. (eds). Flora Eu
ropaea. Vol 2. Cambridge University Press, Cambridge
Specific references
Cañizo A, Miguel C, RodríguezBarrueco C (1978) The effect of
pH on nodulation and growth of Coriaria myrtifolia L. Plant
and Soil 49:195198
MartínezSánchez JJ, Orozco E, Selva M, Gilabert J (1997) Ob
tención de planta de Coriaria myrtifolia L. en vivero. Ensayos
de inducción a la nodulación en sustrato estéril. Montes
50:4044
Collecting time
autumn winter
Auxin
concentration
none or < 0.5 %
2005). If these conditions cannot be met, it is prefer
able to harvest the cuttings in early spring, when tem
peratures begin to increase. Melgares de Aguilar et al.
(2005) obtained 85 % survival rates when cuttings
were collected in the spring, compared to 100% for
cuttings obtained in the autumn.
Melgares de Aguilar J, González D, Navarro A, Bañón S, Gar
cía F (2005) Influencia de la estacionalidad en el enraiza
miento de esquejes de Coriaria myrtifolia L. V Congresso
Ibérico de Ciências Hortícolas ; IV Congresso Iberoamericano
de Ciências Hortícolas Vol 1:457461. Asociación Portugesa
de Ciencias Hortícolas, Porto
Thompson PN, Gornall PN, Gornall FLS RJ (1995) Breeding
systems in Coriaria (Coriariaceae). Botanical Journal of the
Linnean Society 117:293304
Yokoyama J, Suzuki M, Iwatsuki K, Hasebe M (2000) Molecu
lar phylogeny of Coriaria, with special emphasis on the dis
junct distribution. Molecular Phylogenetics and Evolution
14:11–19
Coriaria myrtifolia
42
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus,
Western Asia
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia,
BosniaHerzegovina, Montenegro, Albania, Greece,
Turkey
Diagnostic traits
Cornus sanguinea is a deciduous shrub, 1.5 to 6 m in
height, with dark red twigs and opposite, ovate or el
liptic, entire leaves. Its flowers, which have dull white
or pale yellowishwhite petals, come out after its leaves
Reproductive biology
■
hermaphroditism
Flowering
white flowers,
clustered in large
corymbiform cymes
■ from April to July,
sometimes also in
autumn
■
Pollination
■
The common dogwood is a species that requires a cool
environment. In the Mediterranean region it thus finds
refuge in shady areas, ravines, riverbanks, and humid
thorny thickets. In less dry regions it is found on the
edges of forests and glades and in deciduous thickets.
It needs soils that are relatively rich in nutrients but
grows on substrates with different pH levels. It toler
ates calcareous materials quite well and grows easily
on heavy soils.
have grown, in contrast to Cornus mas, a species with
yellow or greenish flowers that is found widely in west
ern Asia and Europe.
entomophilous
Fruiting
Ripening
■
■
■
black globose drupe
58 mm
from July to
October
■ dispersal by
frugivorous
vertebrates
43 Cornus sanguinea
Distribution and Ecology
Sex expression
Cornus
sanguinea L.
EN: common dogwood
EL: αγριοκρανιά
ES: cornejo
FR: cornouiller sanguin
IT: corniolo
PT: sanguinho legítimo
Cornaceae
Variation and Hybridisation
Two subspecies have been described: C. sanguinea
subsp. sanguinea and C. sanguinea subsp. australis. The
latter is found in Southeast Europe and Southwest Asia.
The difference between these subspecies has to do with
the type of indumentum on the back of the leaves,
which consists of simple, more or less curly hairs in C.
sanguinea, while in C. australis the hairs are navicular
and parallel to the nerves.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from August to early
autumn
■ gathering by hand
sequence for fleshy
fruits
■ seed weight / kg
fruit: 172317g
■ purity: 100 %
Cornus sanguinea crops vary both interannually and
between populations, but in general the fruittoflower
ratio is very low, due to the high incidence of fruit
abortion throughout the stages of its development
(Krüsi and Debussche, 1988). However, in cases of high
3055 g
flower mortality (resulting from predation, for exam
ple), the proportion of aborted fruit is lower (Guitián
et al., 1996). Fruits should be collected as soon as they
are ripe, to reduce losses to birds.
Germination under controlled conditions
Pregermination treatments
preheating (8 weeks) + prechilling
(812 weeks)
■ scarification with concentrated
sulphuric acid (120 minutes) +
prechilling (12 weeks)
■
Nursery production
Sowing season
■
autumn, without treatment, or
spring, with treatment
Conditions
■
■
30 / 20 ºC; 20 / 10 ºC
light
Nursery practice
■
■
T: 4 ºC
MC: 48 %
■ airtight container
■
forestpot 300 cm3: 1/0
container 3.5 l: 1/1
Germination Viability
■
8096 %
Emergence
■
the first spring, may be
completed in the second spring
Cornus sanguinea 44
Vegetative propagation
Type of cutting
■
hardwood
Position along
the shoot
indifferent
Number of
internodes Size
20 cm
Vegetative propagation of Cornus sanguinea is readily
achieved with hardwood material collected in the win
ter. It is not necessary to apply hormones, although
their use makes the rooting time uniform.
References
General references
Ball PW (1968) Cornus L. In: Tutin TG et al. (eds). Flora Euro
paea. Vol 2. Cambridge University Press, Cambridge
Catalán G (1991) Semillas de árboles y arbustos forestales.
Ministerio de Agricultura Pesca y Alimentación. ICONA,
Madrid
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns Ge
hölzvermehrung. Parey Buchverlag, Berlin
Nieto Feliner G (1997) Cornus L. In: Castroviejo S et al. (eds).
Flora Ibérica. Vol 8. CSIC, Madrid
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea. ANPA, Roma
Collecting time
winter
Auxin
concentration
none
Experiments have been conducted on in vitro propaga
tion of other species of the genus Cornus (Edson et al.,
1994; Kaveriappa et al., 1997).
Specific references
Edson JL, Wenny DL, LeegeBrusven A (1994) Micropropaga
tion of Pacific dogwood. HortScience 29:13551356
Guitián J, Guitián P, Navarro L (1996) Fruit set, fruit reduc
tion, and the fruiting strategy of Cornus sanguinea (Cor
naceae). American Journal of Botany 83:744748
Kaveriappa KM, Phillips LM, Trigiano RN (1997) Micropropa
gation of flowering dogwood (Cornus florida) from seedlings.
Plant Cell Reports 16:485489
Krüsi BO, Debussche M (1988) The fate of flowers and fruits
of Cornus sanguinea L. in three contrasting Mediterranean ha
bitats. Oecologia 74:592599
45 Cornus sanguinea
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Crataegus
monogyna Jacq.
Distribution and Ecology
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus,
Western Asia, Northern Africa
Mediterranean region: Portugal, Spain (incl. Baleares),
France (incl. Corse), Italy (incl. Sardegna and Sicilia),
Croatia, BosniaHerzegovina, Montenegro, Albania,
Diagnostic traits
The hawthorn is a small prickly tree, 5 (10) m in height,
with thorns 7 to 20 mm in length. Its leaves exhibit
large variations in shape and size within the same in
dividual, from deeplylobed to entire blade. The genus
Crataegus, like other species of Rosaceae, is of great
taxonomic complexity.
C. monogyna can be distinguished from the other species
of the genus that are found in Mediterranean Europe by
EN: hawthorn, white thorn
EL: τρικουκκιά, μουρτζιά
ES: espino albar, majuelo
FR: aubépine, noble épine
IT: biancospino, marucca bianca
PT: pilriteiro, espinheiroalvar
Rosaceae
Greece (incl. Kriti), Cyprus, Turkey, Syria, Lebanon,
Israel, Tunisia, Algeria, Morocco
This species has a broad ecological range. It is found at
the edges and in glades of deciduous forests and in
deciduous thorny thickets. In more arid regions, it is
limited to riparian zones and shady areas.
the shape of its leaves or the pilosity of its different
structures. Lobes that are entire or have a few acute
teeth and entire stipules differentiate it from C. laevi
gata, whose leaves have serrulate lobes and serrate stip
ules. The twigs, leaves, pedicels and receptacles of C.
heldreichii, C. azarolus and C. pycnoloba, species with
limited distribution in the eastern Mediterranean region,
are tomentose, lanate or sericeous, whereas in C. monog
yna they are glabrous or have straight, patent hairs.
Crataegus monogyna 46
Reproductive biology
Sex expression
■
hermaphroditism
Flowering
white or pale pink
flowers, clustered in
corymbs, 4 to 11
flowers per
inflorescence
■ from March to June
■
Pollination
■
■
entomophilous
selfcompatible
Variation and Hybridisation
The complexity of the genus Crataegus is a result of
polyploidy (Talent and Dickinson, 2005), hybridisation,
introgression and apomixis. Crataegus monogyna in
cludes a complex, widely varied number of plants. Nu
merous subspecies or varieties have been determined by
different authors on the basis of qualitative or quanti
tative traits relating to leaves, flowers or fruits. Natural
hybrids of C. monogyna and C. azarolus or C. laevigata
have been described. The introgression between C.
monogyna and C. laevigata seems to have been con
firmed by molecular techniques (Fineschi et al., 2005).
Genetic diversity between and within populations, de
termined by molecular techniques, would seem to be
Fruiting
Ripening
■
■
■
red pome
610 mm
from August to
November
■ dispersal by
frugivorous
vertebrates
low and to lack spatial structuring, possibly as a result
of extensive fruit dispersal by animals (Fineschi et al.,
2005). According to the results of these determinations,
material from individuals of distant populations could
potentially be collected and mixed, although it is ad
visable to be cautious and to keep to collection units
located within the limits of the same region of prove
nance, seed zone or ecological unit. Such conservative
practice is supported by the results obtained when
using material of different origins in a series of refor
estation projects under varying ecological conditions,
where local populations have been shown to adapt bet
ter to the climate and to be more resistant to disease
(Jones et al., 2001).
Seed propagation
Seed handling and storage
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from late summer to
early autumn
■ gathering by hand
sequence for fleshy
fruits
■ seed weight / kg
fruit: 150230 g
■ purity: 99100 %
C. monogyna seems to exhibit a marked tendency to
fruit abortion, especially at the beginning of the fruit
development cycle, although the incidence varies be
tween individuals (Guitián et al., 1992). Material col
lection should not focus too heavily on the most
productive plants, but must be well spread out among
the entire tree population.
55180 g
T: 4 ºC (23 years)
MC: 48 %
■ airtight container
■
Collecting at the end of the summer, when the fruit is
reddish but has not yet ripened completely, can shorten
seed germination time.
47 Crataegus monogyna
Tolerance to desiccation: ORTHODOX
Germination under controlled conditions
Pregermination treatments
preheating (416 weeks) + prechilling
(1236 weeks)
■ mechanical scarification + prechilling
(48 or more weeks)
■ scarification with concentrate sulfuric
acid (30120 minutes) + prechilling
(48 or more weeks)
■ mechanical or acid scarification +
preheating (412 weeks) + prechilling
(1220 weeks)
■
Conditions
■
Germination Viability
30 / 20 ºC
■
70100 %
Hawthorn seeds display deep embryo dormancy and a
thickened endocarp and require both scarification and
prechilling to germinate.
Nursery production
Sowing season
■
late summer, without treatment,
seeds from not completely ripened
fruits immediately after collecting
these, or spring with treatment
Vegetative propagation
Type of cutting
hardwood
semihardwood
■ root
■
■
Position along
the shoot
basal or middle
basal or middle
Nursery practice
Emergence
forestpot 300 cm3: 1/0
■ container 3.5 l: 1/1
79 months after late summer
sowing, may be completed in
the following spring
■ 2 months after spring sowing,
may be completed in the second
spring
■
Number of
internodes Size
15 cm
10 cm
58 cm
Crataegus monogyna 48
Vegetative propagation of Crataegus is not common
practice. However, it is possible to use this technique
with acceptable results. In order to obtain suitable
hardwood cuttings, parent plants must have been sub
jected to aggressive pruning to achieve rejuvenation.
Crobeddu and Pignati (2005) obtained 76% rooting
success using semihardwood cuttings harvested in July
from rejuvenated parent plants and maintained in a
mist environment at a basal temperature above 20 ºC.
It is also possible to propagate this species with some
■
Collecting time
winter
summer
winter
Auxin
concentration
1%
0.5 %
none
success from root cuttings. Göttsche (1978) obtained a
30% survival rate with this method, planting cuttings
vertically in a mixture of peat and sand (1:1), that is to
say, leaving part of the root cutting above the sub
strate.
In vitro propagation of Crataegus monogyna is possible
and offers better results than conventional vegetative
propagation. Wawrosch et al. (2007) used axillary buds
collected in winter as cuttings.
General references
Amaral Franco J do (1968) Crataegus L. In: Tutin TG et al. (eds).
Flora Europaea. Vol 2. Cambridge University Press,
Cambridge
Fineschi S, Salvini D, Turchini D, Pastorelli R, Vendramin GG
(2005) Crataegus monogyna Jacq. and C. laevigata (Poir.) DC.
(Rosaceae, Maloideae) display low level of genetic diversity
assessed by chloroplast markers. Plants Systematic and Evo
lution 250:187196
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Göttsche D (1978) Vermehrung einheimischer Straucharten
durch Wurzelschnittlinge. Forstarchiv 49:3336
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns Ge
hölzvermehrung. Parey Buchverlag, Berlin
Guitián J, Sánchez JM, Guitián P (1992) Niveles de fructifica
ción en Crataegus monogyna Jacq., Prunus mahaleb L. y Pru
nus spinosa L. (Rosaceae). Anales del Jardín Botánico de
Madrid 50:239245
Muñoz Garmendia F, Navarro C, Aedo C (1998) Crataegus L. In:
Muñoz Garmendia F, Navarro C (eds). Flora Ibérica. Vol 6. CSIC,
Madrid
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea. ANPA, Roma
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Specific references
Crobeddu S, Pignatti G (2005) Propagazione per talea di specie
mediterranee prove di substrato. Sherwood Foreste ed Alberi
Oggi 114:2731
Jones AT, Hayes MJ, Sackville Hamilton NR (2001) The effect
of provenance on the performance of Crataegus monogyna in
hedges. Journal of Applied Ecology 38:952–962
Talent N, Dickinson TA (2005) Polyploidy in Crataegus and
Mespilus (Rosaceae, Maloideae): evolutionary inferences from
flow cytometry of nuclear DNA amounts. Canadian Journal of
Botany 83:12681304
Wawrosch C, Prinz S, Soleiman Y, Kopp B (2007) Clonal prop
agation of Crataegus monogyna Jacq. (Lindm.). Planta Medica
73:1013
49 Crataegus monogyna
References
Dorycnium
rectum (L.) Ser.
Distribution and Ecology
General distribution: Southwestern and Southeastern
Europe, Western Asia, Northern Africa
Mediterranean region: Portugal, Spain (incl. Baleares),
France (incl. Corse), Italy (incl. Sardegna and Sicilia),
Albania, Greece (incl. Kriti), Turkey, Syria, Lebanon,
Israel, Tunisia, Algeria, Morocco
Diagnostic traits
This is a herbaceous perennial plant, sometimes
woody at the base, not spiny, that can reach a height
of 30 to 200 cm. Leaves are composed of five leaflets,
the basal two are ovate, acute and the other three are
obovatespatulate, mucronate. Other species of the
genus that are widely distributed in the Mediter
Reproductive biology
Sex expression
■
hermaphroditism
Flowering
whitepink flowers
clustered in
glomerules, 18 to 40
flowers in each
■ from May
to September
■
Leguminosae
EN: greater badassi
EL: μελιγκάρια
ES: unciana
FR: dorycnie dressée
IT: trifoglino palustre
PT: ervamatapulgas
Pollination
■
Dorycnium rectum grows in grasslands and in rush
beds on the banks of watercourses in the Mediterran
ian region. It prefers alkaline reaction soils. It fixes
atmospheric nitrogen.
ranean region (D. pentaphyllum, D. hirsutum and D.
gracile) are found in inland or coastal bush and pas
ture land. Morphologically, D. rectum can be identified
because its leaves have a rachis which is at least 3.5
mm long, while in the three other species the rachis
is smaller or absent.
entomophilous
Fruiting
Ripening
■
■
purple or brownish
purple legume, more
or less cylindrical
■ 1020 mm long
from July to
September
■ dispersal by
explosion
Dorycnium rectum 50
Variation and Hybridisation
There is no information on intraspecific variation and
hybridisation for this taxon.
Seed propagation
No information about seed production has been found
for this species. As a general guideline, data relating to
Dorycnium hirsutum are provided; the seeds of the lat
ter are larger in size, so the seeds of D. rectum should
be somewhat lighter.
Seed handling and storage
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
from July to August
gathering by hand
sequence for dehis
cent fruits
■ seed weight / kg
fruit: 163 445 g
(D. hirsutum)
■ purity: 8599 %
46 g (D. hirsutum)
Germination under controlled conditions
Pregermination treatments
■
scarification with concentrated
sulphuric acid (1520 minutes)
Nursery production
Sowing season
■
spring, with treatment
Vegetative propagation
Type of cutting
■
semihardwood
Position along
the shoot
terminal
Conditions
■
Germination Viability
20 ºC
■
Nursery practice
■
forestpot 300 cm3: 1/0 or 2/0
Number of
internodes Size
10 cm
Results obtained by Frangi and Nicola (2004) using Do
rycnium hirsutum cuttings suggest that the best time
of the year for collecting material is the spring, in the
month of April. Alegre et al. (1998) achieved their best
results with D. pentaphyllum and D hirsutum cuttings
T: 4 ºC
MC: 48 %
■ airtight container
■
8098 % (D. hirsutum)
Emergence
■
the same spring
Collecting time
april
Auxin
concentration
0.5 1 %
taken from the apex of stems and treated with hor
mones. These authors recommend protecting the ma
terial from the low night temperatures that can occur
in the spring.
51 Dorycnium rectum
Tolerance to desiccation: ORTHODOX
References
General references
Ball PW (1968) Dorycnium Miller. In: Tutin TG et al. (eds). Flora
Europaea. Vol 2. Cambridge University Press, Cambridge
Díaz Lifante Z (2000) Dorycnium Mill. In: Talavera S et al. (eds).
Flora Ibérica. Vol 7(II). CISC, Madrid
Specific references
Alegre J, Toledo JL, Martinez A, Mora O, Andres EF (1998)
Rooting ability of Dorycnium spp. under different conditions.
Scientia Horticulturae 76:123129
Frangi P, Nicola S (2004) Studio della propagazione per talea
di specie mediterranee di interesse ornamentale. Italus Hortus
11:191193
Dorycnium rectum 52
Distribution and Ecology
General distribution: Southwestern Europe
The tamujo growths in watercourses and dry ravines,
frequently associated with Nerium oleander. This
species prefers welldrained acid soils.
Mediterranean region: Portugal, Spain
Diagnostic traits
Flueggea tinctoria is a spiny deciduous shrub that
grows to 2 m, with many branches rising from the base;
the young branches are dark reddish. Leaves are alter
Reproductive biology
Sex expression
■
dioecy
Flueggea tinctoria
(L.) G.L. Webster
EN: EL: ES: tamujo
FR: IT: PT: tamuxo
Euphorbiaceae
Flowering
greenish, solitary or
fasciculated flowers;
male patent/erect;
female pendulous
■ from January to
April
■
Pollination
■
nate, simple, obovate and glabrous, with a blunt or
emarginate apex.
anemophilous
Fruiting
Ripening
■
■
■
threelobed capsule
3.54 mm
■
from May to June
dispersal by
gravity
Variation and Hybridisation
53 Flueggea tinctoria
There is no information on intraspecific variation and
hybridisation for this taxon.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
from May to June
gathering by hand
sequence for fleshy
fruits
■ seed weight / kg
fruit: 54 g
■ purity: 98 %
■
Germination under controlled conditions
Pregermination treatments
■
no treatments required
Nursery production
Sowing season
■
autumn or spring
■
■
hardwood
Position along
the shoot
indifferent
Germination Viability
20 ºC
■
Nursery practice
■
Vegetative propagation
Type of cutting
Conditions
■
Number of
internodes Size
References
General references
Benedí C (1997) Flueggea Willd. In: Castroviejo S et al. (eds).
Flora Ibérica. Vol 8. CSIC, Madrid
95 %
Emergence
forestpot 300 cm3: 1/0
container 3.5 l: 1/1
20 cm
T: 4 ºC
MC: 48 %
■ airtight container
4g
■
2 to 3 weeks after sowing
Collecting time
winter
Auxin
concentration
none
Gálvez A, Navarro RM (2001) Manual para la identificación y
reproducción de semillas de especies vegetales autóctonas de
Andalucía. Vol II. Consejería de Medio Ambiente, Junta de
Andalucía, Sevilla
Flueggea tinctoria 54
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus, West
ern and Middle Asia, Siberia, China, Northern Africa
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy, Croatia, BosniaHerzegovina, Montenegro,
Albania, Greece, Turkey, Tunisia, Algeria, Morocco
Diagnostic traits
Frangula alnus is a deciduous shrub or small tree with
ovaloblong, pointed leaves. Its height of 4 to 5 m dis
tinguishes it from Frangula rupestris (Scop.) Schur., an
Reproductive biology
■
hermaphroditism
Flowering
whitegreenish
flowers, clustered in
cymes
■ from March to July
■
Pollination
■
■
This species requires cool and damp soils, preferably
acidic, although it can also grow on calcareous sub
strates. It is found scattered through humid forests,
along riversides and in shady ravines in Eurosiberian
environments and reaches the Mediterranean region in
riparian areas where conditions are moist enough.
endemic bush of the eastern Mediterranean region that
grows to only 80 cm in height.
entomophilous
selfincompatible
Although frugivorous birds are the principal seed dis
persers for Alnus glutinosa, downstream dispersion by
water becomes predominant in the Mediterranean Re
Fruiting
Ripening
■
■
dark red globose
drupe
■ about 7 mm
from June to
October
■ dispersal by
frugivorous
vertebrates
gion, where riparian zones offer a suitable habitat
(Hampe, 2004).
55 Frangula alnus
Distribution and Ecology
Sex expression
Frangula
alnus Mill.
EN: alder buckthorn
EL: βουρβουλιά
ES: arraclán
FR: bourdaine
IT: frangola
PT: sanguinhodaágua
Rhamnaceae
Variation and Hybridisation
Of the several subspecies of Frangula alnus that have
been described, two are located in Mediterranean
countries. F. alnus subsp. baetica (Reverchon & Wilk)
Rivas Goday ex Devesa, which is limited to the south of
Spain and the north of Morocco, is bigger than the type
subspecies and has large leaves (514 x 25.5 cm). The
other, F. alnus subsp. pontica (Boiss.) Davis & Yalt., is a
shrub or small tree that is endemic to Anatolia and has
oblonglanceolate leaves and glabrous twigs instead of
the obovateelliptic leaves and pubescent twigs of the
type subspecies.
Studies using molecular techniques (Hampe et al.,
2003) reveal extensive genetic differentiation through
out the distribution range of the species, which in
cludes three lineages – Iberia, Anatolia and temperate
Europe – as a result of its evolutionary history. In mar
ginal Mediterranean populations, a high degree
of interpopulation genetic differentiation has been es
timated due to limited gene flow between populations
in these areas, even when close to each other, although
intrapopulation variation is low. This pattern of genetic
variation makes extreme caution advisable when mov
ing reproductive materials of this species; whenever
possible, local populations should be used, especially if
restoration is to take place in areas where endemic
subspecies grow.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from August to
November
■ gathering by hand or
using longhandled
tools
sequence for fleshy
fruits
■ seed weight / kg
fruit: 90150 g
■ purity: 98100 %
In the Mediterranean region, where only small popula
tions or more or less isolated individuals are to be
found, fruit production is scarce due both to limited
pollen (Medan, 1994; Hampe 2005) and to climatic
factors – especially drought – which lead to a signifi
cant degree of interannual crop variation (Hampe
2005). These aspects should be borne in mind when
Germination under controlled conditions
Pregermination treatments
■
prechilling (8 weeks)
Conditions
■
■
30 / 20 ºC
light
1627 g
T: 4 ºC
MC: 48 %
■ airtight container
■
planning to collect and produce reproductive material
of this species.
Fruits should be collected about two weeks before they
have ripened completely, in order to avoid predation by
birds.
Germination Viability
■
7094 %
Frangula alnus 56
Sowing season
■
autumn, without treatment,
or spring, with treatment
Nursery practice
bareroot: 50 g/m2; circumfe
rence up to 46 cm or total
height up to 80100 cm
3
■ forestpot 300 cm : 1/0
■ container 3.5 l: 1/1
■
According to Gálvez and Navarro (2001), the seeds of
F. alnus subsp. baetica do not require cold stratifica
Vegetative propagation
Type of cutting
■
semihardwood
Position along
the shoot
basal or middle
2 3 / 5 10 cm
References
■
the first spring
tion; they can be sowed directly and germinate in the
same spring they are seeded.
Number of
internodes Size
Cuttings may be obtained all along the set if young
mother plants are used, although those from the basal
and middle parts form stronger roots. When using cut
tings collected from adult material, the regenerative
capacity of terminal cuttings is considerably reduced
(Graves, 2002). Treatment with 0.3 0.8% indolbutyric
acid powder noticeably improves the results. Rooting
in vermiculite in a mist environment is recommended
(Sharma and Graves, 2005). Some references describe
Emergence
Collecting time
summer
Auxin
concentration
0.5 %
propagation using hardwood cuttings in other species
of the Rhamnaceae family, using material collected in
winter from the basal or middle part of the stem and
treated with hormones (Bañón et al., 2003; Dirr and
Heuser, 2006).
Frangula alnus has been successfully reproduced in
vitro, using axillary buds (Bignami 1983) and excised
embryos (Kovacevic and Grubisic, 2005).
General references
Specific references
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Bañón S, Martínez JJ, Fernández JA, Ochoa J, González A
(2003) Effect of indolebutyric acid and paclobutrazol on the
rooting of Rhamnus alaternus stem cuttings. Acta Horticul
turae 614:263267
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns Ge
hölzvermehrung. Parey Buchverlag, Berlin
Muñoz JM (1987) Frangula. In: Valdés B, Talavera S, Fernán
dezGaliano E (eds). Flora Vascular de Andalucía Occidental.
Vol 2. Ketres Editora SA, Barcelona.
Tutin TG (1968) Frangula Miller. In: Tutin TG et al. (eds). Flora
Europaea. Vol 2. Cambridge University Press, Cambridge
Yaltirik F (1967) Frangula Miller. In: Davis PH (ed). Flora of Turkey
and the Eastern Aegean Islands. Vol 2. University Press, Edinburg
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Bignami C (1983) In vitro propagation of Rhamnus frangula L..
Gartenbauwissenschaft 48:272274
Dirr MA, Heuser CW (2006) The Reference Manual of Woody
Plant Propagation: From Seed to Tissue Culture. A Practical
Working Guide to the Propagation of over 1100 Species, 2nd
ed. Varsity Pr. Inc, Athens
Gálvez A, Navarro RM (2001) Manual para la identificación y
reproducción de semillas de especies vegetales autóctonas de
Andalucía. Vol II. Consejería de Medio Ambiente, Junta de An
dalucía, Sevilla
57 Frangula alnus
Nursery production
Graves WR (2002) IBA, juvenility, and position on ortets in
fluence propagation of Carolina buckthorn from softwood
cuttings. Journal of Environmental Horticulture 20:5761
Hampe A (2004) Extensive hydrochory uncouples spatiotem
poral patterns of seedfall and seedling recruitmente in a “bird
dispersed” riparian tree. Journal of Ecology 92:797807
Hampe A (2005) Fecundity limits in Frangula alnus (Rham
naceae) relict populations at the species’ southern range mar
gin. Oecologia 143:377386
Hampe A, Arroyo P, Jordano P, Petit RJ (2003) Rangewide phy
logegraphy of a birddispersed Eurasian shrub: contrasting
Mediterranean and temperate glacial refugia. Molecular
Ecology 12:34153426
Kovacevic N, Grubisic D (2005) In vitro cultures of plants from
the Rhamnaceae: shoot propagation and anthraquinones pro
duction. Pharmaceutical Biology 43:420424
Medan D (1994) Reproductive biology of Frangula alnus
(Rhamnaceae) in southern Spain. Plant Systematics and Evo
lution 193:173186
Sharma J, Graves WR (2005) Propagation of Rhamnus alnifo
lia and Rhamnus lanceolata by seeds and cuttings. Journal of
Environmental Horticulture 23:8690
Frangula alnus 58
EN: narrowleaved ash
EL: νερόφραξος
ES: fresno de hoja estrecha
FR: frêne oxyphylle
IT: frassino meridionale
PT: freixodefolhasestreitas
Distribution and Ecology
General distribution: Southwestern, Southeastern,
Middle and Eastern Europe, Caucasus, Western Asia,
Northern Africa
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia,
BosniaHerzegovina, Montenegro, Albania, Greece,
Turkey, Tunisia, Algeria, Morocco
In the Mediterranean region, Fraxinus angustifolia
Diagnostic traits
Fraxinus angustifolia grows to a height of 15 to 20 m.
The leaves are deciduous and composed of (3)5 to
13(15) lanceolate, serrate leaflets. Its brownish buds
make it possible to differentiate this ash from Fraxinus
excelsior (common ash), a species with black terminal
buds. Another diagnostic trait that clearly differentiates
these species is the inflorescence, which is a raceme in
F. angustifolia and a panicle in F. excelsior. F. angusti
folia usually exhibits fewer, and smaller, leaflets per leaf.
In addition, in F. angustifolia the number of leaflet teeth
is equal to or lower than the number of lateral nerves
and the teeth arch outwards, whereas in F. excelsior
Fraxinus
angustifolia Vahl.
grows in riparian forests, normally in the high areas of
banks that are occasionally flooded for short periods
of time, in contact with zonal vegetation. It is some
times found in valley bottoms with a high groundwa
ter table or in cool, shady forests. It grows in pure
stands or mixed with other tree species. In some areas
of its eastern distribution range it also grows in
sweetwater swamps. Although somewhat indifferent
to substrate, this species prefers decarbonated, sandy
textured soils.
there are more teeth than lateral nerves and they point
towards the apex of the leaflet. Another distinctive fea
ture is that F. angustifolia usually has fewer flowers (or
fruits) per inflorescence than F. excelsior (15 to 20 vs. 50
to 150). F. angustifolia tolerates swampy soils better
than F. excelsior and likes higher temperatures.
F. angustifolia is easily distinguished from Fraxinus
ornus, since the latter’s flowers have white petals that
cluster in striking terminal inflorescences. F. ornus does
not usually mix with riparian vegetation, but grows on
sunny slopes in coniferous, leafy or mixed forests.
59 Fraxinus angustifolia
Oleaceae
Reproductive biology
Sex expression
■
andromonoecy
Flowering
inconspicuous
flowers, clustered
in racemes
■ from February to
May, before leaf
development
■
Pollination
■
anemophilous
Variation and Hybridisation
Three subspecies have been described on the basis of
samara shapes and leaflet numbers. Each subspecies is
found in a rather welldefined geographical area: spp.
angustifolia in the western Mediterranean region; spp.
oxycarpa (Bieb. ex Willd.) Franco & Rocha Alfonso in
east central Europe and southern Europe, from NE
Spain eastwards; and spp. syriaca (Boiss.) Yalt. in Turkey
and eastwards to Iran. This determination based on ge
ographical structure is also supported by phylogeo
graphical studies using molecular techniques (Heuertz
et al., 2006). However, it must be remembered that
there are intermediate forms between the above taxa
Fruiting
Ripening
■
■
distally winged
samara
■ 2040 mm long
from September
to October
■ dispersal by wind
(Fraxigen, 2005), as well as hybrids of F. angustifolia
and F. excelsior in areas where the two species are in
contact (FernándezManjares et al., 2006; Gerard et al,
2006).
Genetic studies of F. angustifolia (Fraxigen, 2005) esti
mate high levels of gene flow, transported between
stands by pollen, and high variation between individu
als within the same stand, due to the pollination pat
tern of this species. These results suggest that the
collection unit for a single lot may cover a more or less
large area that includes several stands.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
from October
gathering by hand, using
longhandled tools or
beating branches
sequence for fruits
that can be sown
directly
■ purity: 9099 %
This species exhibits interannual variation in fruit pro
duction and in some years there is almost no fruit at all.
The proportion of fruit with empty seeds is also high,
due to predation and seed abortion.
Samaras are collected in autumn, when they are brown.
High temperatures should be avoided in the period be
40100 g (samaras)
T: 4 ºC
MC: 48 %
■ airtight container
■
tween harvesting and fruit processing, since fermen
tation may occur (Piotto and Piccini, 2000). Treated
samaras that are ready to germinate can be preserved
for a year at a temperature of –3 ºC. The method in
cludes hot stratification for 15 days followed by cold
stratification for an additional 15 days and drying to a
9.5% moisture content (Piotto, 1997).
Fraxinus angustifolia 60
Germination under controlled conditions
Pregermination treatments
■
■
prechilling (616 weeks)
preheating (4 weeks) + prechilling
(48 weeks)
Conditions
■
25 / 4 ºC
The seeds of Fraxinus angustifolia are subject to phys
iological dormancy and require temperature fluctua
tions to germinate. If samaras are exposed to constant
Nursery production
Sowing season
■
autumn, without treatment, or early
spring, with treatment
Germination Viability
■
5080 %
or only slightly fluctuating temperatures, a secondary
dormancy may be induced (Piotto, 1994).
Nursery practice
bareroot: 200250 g/m2;
circumference up to 68 cm or
total height up to 100150 cm
3
■ forestpot 300 cm : 1/0
■ container 3.5 l: 1/1
■
Emergence
■
the first spring; 23 weeks after
spring sowing
Vegetative propagation
References
However, in vitro propagation is possible (PerezParrón
et al., 1994; Tonon et al., 2001a; Tonon et al., 2001b).
General references
Specific references
Amaral Franco J do, Rocha Alfonso ML da (1972) Fraxinus L.
In: Tutin TG et al. (eds). Flora Europaea. Vol 3. Cambridge Uni
versity Press, Cambridge
FernándezManjarres JF, Gerard PR, Dufour J, Raquin C, Fras
cariaLacoste N (2006) Differential patterns of morphological
and molecular hybridization between Fraxinus excelsior L. and
Fraxinus angustifolia Vahl (Oleaceae) in eastern and western
France. Molecular Ecology 15:3245–3257
Catalán G (1991) Semillas de árboles y arbustos forestales.
Ministerio de Agricultura Pesca y Alimentación. ICONA,
Madrid
GarcíaFayos P (coord) (2001) Bases ecológicas para la reco
lección, almacenamiento y germinación de semillas de espe
cies de uso forestal en la Comunidad Valenciana. Banc de
Llavors Forestals, Generalitat Valenciana, Valencia
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Nicolás JL, Iglesias S, Alía R (2001) Fichas descriptivas de es
pecies. In: García del Barrio JM et al. (coord). Regiones de
identificación y utilización de material forestal de reproduc
ción. Ministerio de Medio Ambiente, Madrid
Fraxigen (2005) Ash species in Europe. Biological characteris
tics and practical guidelines for sustainable use. Oxford
Forestry Institute, University of Oxford, UK
Gerard PR, FernándezManjarrés JF, FrascariaLacoste N
(2006) Temporal cline in a hybrid zone population between
Fraxinus excelsior L. and Fraxinus angustifolia Vahl. Molecular
Ecology 15:3655–3667
Heuertz M, Carnevale S, Fineschi S, Sebastiani F, Hausman JF,
Paule L, Vendramin GG (2006) Chloroplast DNA phylogeogra
phy of European ashes, Fraxinus sp. (Oleaceae): roles of hy
bridization and life history traits. Molecular Ecology
15:21312140
61 Fraxinus angustifolia
Fraxinus angustifolia does not form adventitious roots,
so vegetative propagation using cuttings is not viable.
PérezParrón MA, GonzálezBenito ME, Pérez C (1994) Mi
cropropagation of Fraxinus angustifolia from mature and ju
venile plant material. Plant Cell Tissue and Organ Culture
37:297302
Piotto B (1994) Effects of temperature on germination of
stratified seeds of three ash species. Seed Science and Tech
nology, 22:519529
Piotto B (1997) Storage of nondormant seeds of Fraxinus an
gustifolia Vahl. New Forest 14:157166
Piotto B, Piccini C (2000) Dormenza, germinazione e conser
vazione dei semi dei frassini spontanei in Italia. Sherwood
52:1923
Tonon G, Capuana M, Di Marco A (2001a) Plant regeneration
of Fraxinus angustifolia by in vitro shoot organogenesis. Sci
entia Horticulturae 87:291301
Tonon G, Kevers C, Thomas G (2001b) Changes in polyamines,
auxins and peroxidase activity during in vitro rooting of Frax
inus angustifolia shoots: an auxinindependent rooting model.
Tree Physiology 21:655663
Fraxinus angustifolia 62
EN: common ivy
EL: κισσός
ES: hiedra
FR: lierre
IT: edera
PT: hera
Distribution and Ecology
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus, West
ern Asia, Northern Africa, Macaronesia
Mediterranean region: Portugal, Spain (incl. Baleares),
France (incl. Corse), Italy (incl. Sardegna and Sicilia),
Croatia, BosniaHerzegovina, Montenegro, Albania,
Greece (incl. Kriti), Cyprus, Turkey, Syria, Lebanon, Is
rael, Egypt, Libya, Tunisia, Algeria, Morocco
Diagnostic traits
This species is a perennial climbing plant that can
reach 30 m in height. Its leaves are alternate and poly
morphic: leaves of the sterile branches are usually cor
date or palmate with 3 to 5 lobes; leaves of the fertile
branches are usually entire or subentire, elliptic, ovate
or rhombic. In general, the name ‘ivy’ includes not only
H. helix L., but also H. hibernica (G. Kirchn.) Bean., H.
maderensis K. Koch ex A. Rutherf. and other species
with more or less welldefined areas of distribution.
Hedera
helix L.
Common ivy is indifferent to the mineral nature of the
substrate and can grow on swampy or very dry land;
however, it has a preference for moist, fertile soils. It is
found in forests, in moist thickets, on rocky walls and
in shady ravines and is very common in Mediterranean
riparian forests.
These taxa are not easily differentiated and individu
als with intermediate characteristics are found. Diag
nostic traits are mostly related to leaf trichome
characteristics.
Hedera helix is a diploid species and H. hibernica is
tetraploid (Vargas et al., 1999).
63 Hedera helix
Araliaceae
Reproductive biology
Sex expression
■
hermaphroditism
Flowering
yellowishgreen
flowers, clustered in
umbels
■ from July
to December
■
Pollination
■
■
entomophilous
selfincompatible
Although seeds are dispersed from November to June,
peak berry consumption occurs from April to May, or
Variation and Hybridisation
The interspecific taxonomic complexity also applies in
traspecifically and several subspecies have been de
scribed as occurring in different geographic ranges, in
varying numbers depending on the author (Metcalfe,
2005; Valcárcel et al., 2003; Webb, 1968). Genetic
studies using molecular markers have identified the
western Mediterranean region as the area of greatest
genetic diversity in Hedera species, with geographical
Seed propagation
Not much information is available on the sexual prop
agation of ivy, given the enormous ease with which the
Fruiting
Ripening
■
■
black subglobose
berry
■ 79 mm
from November
to June
■ dispersal by
frugivorous
vertebrates
before January/February in a hard winter (Metcalfe,
2005).
variation patterns for the different types found (Grivet
and Petit, 2002). It is therefore advisable to be cautious
when moving reproductive materials from one place to
another, at least at a regional scale. This precaution is
of benefit in terms of promoting the use of the au
tochtonous taxon of each different territory, given that
identification is quite a problem for nonspecialists.
plant propagates vegetatively.
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
from autumn to spring
gathering by hand or
using longhandled tools
sequence for fleshy
fruits
■ seed weight / kg
fruit: (data not
found)
■ purity: 8090 %
H. helix is a shady or semishady species, but it is usu
ally the individuals in the sun that flower and bear
fruit. It is advisable to remove the pulp, which contains
germination inhibitors, although fruit sowing is a com
1935 g
T: 01 ºC
MC: 5560 % for
fruits
■ airtight container
■
mon practice in nurseries. Care must be taken to pre
vent water loss from fruits during storage. Fruit lots
may be stored for 3 to 4 months in a damp place.
Hedera helix 64
Germination under controlled conditions
■
prechilling (4 weeks)
Nursery production
Sowing season
■
autumn, without treatment,
or spring, with treatment
Conditions
■
29 / 6 ºC
Type of cutting
■
semihardwood
Position along
the shoot
basal or middle
■
Nursery practice
■
Vegetative propagation is the method commonly used
to produce ivy plants. Multiplication is relatively sim
ple, although it is necessary to work with young mate
rial because adult tissues do not form roots and do not
respond to auxin treatments (Geneve, 1991; Geneve et
al., 1988). Since cuttings with longer internodes form
more and better roots, it is advisable to collect cuttings
from mother plants located in shady areas, which tend
to form shoots that have longer internodes and are less
lignified than those from plants which receive direct
sunlight (Mortensen and Larsen, 1989). The best cut
References
■
the first spring; 14 weeks after
spring sowing
full sunlight reduce growth (Mortensen and Larsen,
1989).
Number of
internodes Size
12
6570 %
Emergence
forestpot 300 cm3:
1/0 or 2/0
Cultivation in partial shade with temperatures around
20 ºC is recommended, since high temperatures and
Vegetative propagation
Germination Viability
Collecting time
summer
Auxin
concentration
none or < 0.5 %
tings are obtained from the internodes located in the
middle and lower thirds of the stems (Poulsen and An
dersen, 1980). Vegetative propagation is also possible
using petioles from which the leaf blade has been re
moved (Geneve et al., 1988).
In vitro propagation of ivy is possible using nonligni
fied segments of stems. It is possible to obtain more
microcuttings per explant if the apical bud is removed
from the stems (Aljuboory et al., 1991; Auderset et al.,
1996; Awad and Banks, 1981; Banks 1979).
General references
Specific references
Flynn S, Turner RM, Stuppy WH (2006) Seed Information
Database (release 7.0, Oct. 2006) (online URL
http://www.kew.org/data/sid)
Aljuboory KH, Williams DJ, Skirvin RM (1991) Growthregula
tors influence root and shoot development of micropropa
gated algerian ivy. HortScience 26:10791080
Valcárcel V, McAllister HA, Rutherford A, Mill RR (2003)
Hedera L. In: Nieto Feliner G et al., (eds). Flora Ibérica. Vol 10.
CSIC, Madrid
Auderset G, Moncousin C, Rourke J, Morre DJ (1996) Stimu
lation of root formation by thiol compounds. HortScience
31:240243
Webb DA (1968). Hedera L. In: Tutin TG et al. (eds). Flora
Europaea. Vol 2. Cambridge University Press, Cambridge
Awad AEE, Banks MS (1981) Callus initiation and development
of Hedera helix L. as affected by auxin and cytokinin in the
media. Gartenbauwissenschaft 46:116119
65 Hedera helix
Pregermination treatments
Banks MS (1979) Plant regeneration from callus from two
growth phases of English ivy, Hedera helix L. Zeitschrift fur
Pflanzenphysiologie 92:349353
Geneve RL (1991) Patterns of adventitious root formation in
English ivy. Journal of Plant Growth Regulation 10:215220
Geneve RL, Hackett WP, Swanson BT (1988) Adventious root
initiation in debladed petioles from the juvenile and mature
phase of English ivy. Journal of the American Society for Hor
ticultural Science 113:630635
Grivet D, Petit RJ (2002) Phylogeography of the common ivy
(Hedera sp.) in Europe: genetic differentiation through space
and time. Molecular Ecology 11:1351–1362
Metcalfe D (2005) Hedera helix L. Journal of Ecology
93:632648
Mortensen LM, Larsen G (1989) Effects of temperature on
growth of six foliage plants. Scientia Horticulturae 39:149159
Poulsen A, Andersen AS (1980) Propagation of Hedera helix:
Influence of irradiance to stock plants, length of internode
and topophysis of cutting. Physiologia Plantarum 49:359365
Vargas P, McAllister HA, Morton C, Jury SL, Wilkinson MJ
(1999) Polyploid speciation in Hedera (Araliaceae): phyloge
netic and biogeographic insights based on chromosome
counts and ITS sequences. Plant Systematics and Evolution
219:165179
Hedera helix 66
Distribución general: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus, West
ern, Middle and Eastern Asia, Siberia, China, Eastern
and Western Canada, Northeastern, NorthCentral,
Northwestern, Southeastern, SouthCentral and South
western U.S.A., Mexico
Mediterranean region: Portugal, Spain, France (incl.
Diagnostic traits
Humulus lupulus is the only hop species found in Eu
rope. It is a rhizomatous plant with an annual lianoid
aerial stem, equipped with trichomes that it uses for
Reproductive biology
■
dioecy
Flowering
yellowishgreen
flowers, male inflores
cences clustered in
panicles, female
inflorescences, similar
to cones, solitary or
clustered in cymes
■ from May to August
■
Pollination
■
Corse), Italy (incl. Sardegna and Sicilia), Croatia,
BosniaHerzegovina, Montenegro, Albania, Greece,
Turkey
Common hop is found in cool, humid environments in
temperate and cold areas and in riparian forests in the
Mediterranean region. It grows on moist, occasionally
swampy, alkaline to slightly acid soils.
support, and may grow to a height of 5 to 10 m. Its
leaves are generally opposed, widely ovatecordate,
with 3 to 5 coarsely dentate lobes.
anemophilous
Fruiting
Ripening
■
■
globose achene,
covered by yellow
glands
■ about 3 x 2.5 mm
from September
to October
■ dispersal by wind
67 Humulus lupulus
Distribution and Ecology
Sex expression
Humulus
lupulus L.
EN: common hop
EL: λυκίσκος
ES: lúpulo
FR: houblon
IT: luppolo
PT: engatadeira
Cannabaceae
Variation and Hybridisation
Small (1978) identified several hop varieties on the
basis of quantitative and qualitative morphological
leaftraits and geographical distribution. European
populations would mostly be included under H. lupulus
var. lupulus. Later phylogenetic studies using DNA have
suggested the existence of two main types, European
Seed propagation
The hop is not usually sexually propagated, due to the
difficulties of collecting reasonable amounts of seed
and Asian/North American, while China is possibly the
origin of the genus (Murakami et al., 2006). The Euro
pean type seems to exhibit a low level of genetic vari
ation as compared to the North American type,
probably due to rapid, recent expansion.
and to its low viability.
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from September
to October
■ gathering by hand or
using longhandled tools
sequence for
dehiscent fruits
■ seed weight / kg
fruit: (data not
found)
■ purity: 95 %
Germination under controlled conditions
Pregermination treatments
■
prechilling (510 weeks)
Nursery production
Sowing season
■
autumn, without treatment, or
spring, with treatment
Conditions
■
25 / 15 ºC (8 / 16 h)
Nursery practice
■
forestpot 300 cm3:
1/0 or 2/0
The seeds should be covered thinly after sowing.
Seedlings are very delicate and are susceptible to frost
or cold injury. It is possible to sow in small pots (volume
T: 4 ºC
MC: 48 %
■ airtight container
2.8 3.5 g
■
Germination Viability
■
95 %
Emergence
■
spring; 3 to 4 weeks after
sowing
up to 7075 cm3). Once the seedlings have become es
tablished in these smaller cells, they can be removed
and transplanted into the growth containers.
Humulus lupulus 68
Type of cutting
■
■
softwood
rhizome
Position along
the shoot
indifferent
Number of
internodes Size
2
10 cm
Hops are normally propagated by means of rhizomes
and also by softwood cuttings (Buzi, 2000). This species
produces very long rhizomes from which a great quan
tity of propagation material may be obtained. The roots
are cut into pieces at the end of the growth period and
buried horizontally in sand. Once the cuttings have
sprouted, they are transferred to a container. The re
generative capacity of aerial cuttings also makes these
an easy propagation method, although strongly de
pendent on the clone. Hormone treatment accelerates
rooting and helps to make it more uniform (Howard,
1967). Summer cuttings, using material from the mid
dle part of the sets, should be held in a mist environ
ment. Howard (1965) recommends using cuttings with
References
General references
Catalán P (1993) Humulus L. In: Castroviejo S et al. (eds). Flora
Ibérica. Vol 3. CSIC, Madrid
Ellis RH, Hong TD, Roberts EH (1985) Handbook of Seed Tech
nology for Genebanks Volume II. Compendium of Specific
Germination Information and Test Recommendations Hand
books for Genebanks: No. 3. IPGRI, Rome
Flynn S, Turner RM, Stuppy WH (2006) Seed Information Da
tabase (release 7.0, Oct. 2006) (online URL
http://www.kew.org/data/sid)
Collecting time
springsummer
winter
Auxin
concentration
none or < 0.5 %
none
two internodes and leaving the leaves on the upper in
tenode to maintain the photosynthetic activity of the
cutting and to promote the accretion of carbohydrates
to the basal area. Long days (16 hours of light) and
good illumination help to produce more and better
roots (Howard and Sykes, 1966).
In vitro propagation is possible, although there are
marked differences in the capacity of different individ
uals to respond to this method. Since this plant is of
great agricultural interest, several protocols for mass
micropropagation exist (Fortes and Pais, 2000; Gurri
arán et al., 1999; Roy et al., 2001; Smykalova et al.,
2001).
regeneration in cultures of Humulus lupulus L. (hop) cvs. Bre
wers Gold and Nugget. Plant Cell Reports 18:10071011
Howard BH (1965) Regeneration of the hop plant (Humulus
lupulus L.) from softwood cuttings. I. The cutting and its roo
ting environment. Journal of Horticultural Science 40:181
191
Howard BH (1967) Regeneration of the hop plant (Humulus
lupulus L.) from softwood cuttings. III. Trating with rootpro
moting substances. Journal of Horticultural Science 42:105
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Howard BH, Sykes JT (1966) Regeneration of the hop plant
(Humulus lupulus L.) from softwood cuttings. II. Modification
of carbohydrate ressources within cutting. Journal of Horti
cultural Science 41:155
Tutin TG, Edmonson JR (1993) Humulus L. In: Tutin TG et al.
(eds). Flora Europaea. Vol 1. 2nd edn. Cambridge University
Press, Cambridge
Murakami A, Darby P, Javornik B, Pais MSS, Seigner E, Lutz A,
Svoboda P (2006) Molecular phylogeny of wild Hops, Humu
lus lupulus L. Heredity 97:6674
Specific references
Buzi A (2000) Il luppolo come pianta ornamentale. Colture
Protette 29:6370
Fortes AM, Pais MS (2000) Organogenesis from internodede
rived nodules of Humulus lupulus var. Nugget (Cannabina
ceae). American Journal of Botany 87:971979
Gurriaran MJ, Revilla MA, Tames RS (1999) Adventitious shoot
Roy AT, Leggett G, Koutoulis A (2001) Development of a shoot
multiplication system for hop (Humulus lupulus L.). In Vitro
Cellular and Developmental Biology Plant 37:7983
Small E (1978) A numerical and nomenclatural analysis of
morphogeographic taxa of Humulus. Systematic Botany
3:3776
Smykalova I, Ortova M, Lipavska H, Patzak J (2001) Efficient in
vitro micropropagation and regeneration of Humulus lupulus on
low sugar, starchGelrite media. Biologia Plantarum 44:712
69 Humulus lupulus
Vegetative propagation
Laurus
nobilis L.
Lauraceae
EN: bay tree
EL: δάφνη
ES: laurel
FR: laurier sauce
IT: alloro
PT: loureiro
Distribution and Ecology
General distribution: Southwestern and Southeastern
Europe, Western Asia, Northern Africa
of many bay laurel populations, since the tree has been
widely cultivated in the Mediterranean region.
It is difficult to determine the autochthonous nature
The bay laurel is sensitive to cold and grows in tem
perate and moderately humid regions, where it is usu
ally found near the coast. It grows scattered in damp
forests, shady ravines and valley bottoms and among
riparian vegetation. More rarely, it can grow in pure
stands or as the dominant species in particularly humid
coastal areas.
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia,
BosniaHerzegovina, Montenegro, Albania, Greece (incl.
Kriti), Cyprus, Turkey, Syria, Lebanon, Israel, Libya,
Tunisia, Algeria, Morocco
Diagnostic traits
L. nobilis is a small to mediumsized perennial tree that
reaches 5 to 10 m in height. Its young twigs are
glabrous and its leaves are oblonglanceolate, coria
ceous, glabrous and aromatic. It can be distinguished
Reproductive biology
Sex expression
■
dioecy
Flowering
Laurus nobilis 70
yellowishgreen or
whitish flowers,
clustered in umbels,
4 to 6 flowers in
each
■ from February to
May
■
Pollination
■
from L. azorica (Seub.) Franco, which is found in the
Macaronesian islands and the north of Africa, in that
the latter’s leaves are initially somewhat hairy on the
back and its young twigs are densely hairy.
entomophilous
Fruiting
Ripening
■
■
black, ovoidglobose
berry
■ 1015 mm
from September
to October
■ dispersal by
frugivorous
vertebrates
Variation and Hybridisation
There is no information on intraspecific variation and
hybridisation for this taxon.
Seed propagation
Seed handling and storage
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from September
to October
■ gathering by hand or
using longhandled tools
sequence for fleshy
fruits
■ seed weight / kg
fruit: (data not
found)
■ purity: 98 %
The seeds of the sweet bay can be preserved for 48
months at 0 ºC and must be treated as recalcitrant
(Konstantinidou et al., 2007); care must be taken to
prevent desiccation during cleaning and storage. It is
advisable to remove the pulp of the fruits because it
contains germination inhibitors (Takos, 2001; Tilki,
■
prechilling (4–12 weeks)
Nursery production
Sowing season
■
autumn, without treatment, or
spring, with treatment
Conditions
■
■
20 ºC
light
Nursery practice
■
■
■
2004; Sari et al., 2006). However, direct fruit sowing is
possible. This is a common practice in nurseries because
it reduces handling and allows 2 to 4 months storage
of lots provided that any loss of water content from
the fruits is prevented over this period.
Germination under controlled conditions
Pregermination treatments
T: 01 ºC
MC: 5560 %
■ open container
8301000 g
forestpot 300 cm3: 1/0
container 3.5 l: 1/1
Germination Viability
■
5070 %
Emergence
■
the first spring, complete within
23 months
71 Laurus nobilis
Tolerance to desiccation: RECALCITRANT
Vegetative propagation
Type of cutting
■
semihardwood
Position along
the shoot
basal or middle
Number of
internodes Size
2 / 5 10 cm
Propagation of bay laurel by cuttings is difficult (Raviv
et al., 1983; Viola et al., 2004). The capacity to produce
adventitious roots is highly variable between individu
als. The best results are obtained by collecting cuttings
in summer, from juvenile parent plants with profuse
leaves (Piccioni et al., 1996). Raviv and Putievsky (1984)
recommend rooting in hotbeds in a mist atmosphere,
using a mixture of peat and perlite (1:1) as substrate.
References
General references
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
GarcíaFayos P (coord) (2001) Bases ecológicas para la reco
lección, almacenamiento y germinación de semillas de espe
cies de uso forestal en la Comunidad Valenciana. Banc de
Llavors Forestals, Generalitat Valenciana, Valencia
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di
alberi e arbusti della flora mediterranea. ANPA, Roma
Collecting time
August to September
Auxin
concentration
0.5 %
Bay laurel can also be propagated by aerial layering,
although this method is not practical if many plants
are required.
In vitro propagation of bay laurel from axillary buds has
been successfully attempted (Souayah et al., 2002).
Raviv M, Putievsky E (1984) Rooting of stem cuttings of bay
laurel: rooting media and fungicidal treatments. Hassadeh
64:22472249
Raviv M, Putievsky E, Ravid U, Senderovitch D, Snir N, Roni R
(1983) Native bay laurel (Laurus nobilis L.) as an ornamental
plant. Acta Horticulturae 132:3542
Sari AO, Oguz B, Bilgic A (2006) Breaking seed dormancy of
laurel (Laurus nobilis L.). New Forests 31:403408
Tutin TG (1993) Laurus L. In: Tutin et al. (eds). Flora Europaea.
Vol 1. 2nd edn. Cambridge University Press, Cambridge
Souayah N, Khouja ML, Khaldi A, Rejeb MN, Bouzid S (2002)
Breeding improvement of Laurus nobilis L. by conventional and
in vitro propagation techniques. Journal of Herbs, Spices and
Medicinal Plants 9:101105
Villar L (1986) Laurus L. In: Castroviejo S et al. (eds). Flora Ibé
rica. Vol 1. CSIC, Madrid
Takos I (2001) Seed dormancy in bay laurel (Laurus nobilis L.).
New Forests 21:105–114
Specific references
Tilki F (2004) Influence of pretreatment and desiccation on
the germination of Laurus nobilis L. seeds. Journal of Environ
mental Biology 25:157161
Konstantinidou E, Takos I, Merou T (2007) Desiccation and sto
rage behavior of bay laurel (Laurus nobilis L.) seeds. European
Journal of Forest Research 127: 125131
Piccioni E, Longari F, Standardi A, Ciribuco S (1996) Propaga
zione per talea e allevamento in vaso di alcune specie arbus
tive. Informatore Agrario 52:8791
Viola F, Forleo LR, CocozzaTalia MA (2004) Propagazione aga
mica di alcune specie della macchia mediterranea. Italus Hor
tus 11:186190
Laurus nobilis 72
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus, West
ern Asia, Northern Africa
Mediterranean region: Portugal, Spain, France, Italy,
Croatia, BosniaHerzegovina, Montenegro, Albania,
Greece, Turkey, Morocco
Diagnostic traits
Ligustrum vulgare is a shrub of 1 to 5 m in height, nor
mally deciduous although it can keep its leaves in warm
climates. Its leaves are opposite, ellipticlanceolate to
oblanceolate. In Europe there are many nonnative or
Reproductive biology
■
hermaphroditism
Flowering
white flowers,
clustered in thyrses
■ from May to July
■
Pollination
■
The common privet withstands cold continental cli
mates, but with warm summers. It is found as scattered
individuals in thorny thickets, ravines, glades and river
side areas. It prefers limy soils with clayey or silty tex
tures and a certain amount of humidity.
namental taxa that must be avoided in restoration
projects, such as the perennial privets from Eastern
Asia: Ligustrum lucidum Aiton, a small tree, and Ligus
trum ovalifolium Hassk, a shrub.
entomophilous
Fruiting
Ripening
■
■
■
black globose berry
68 mm
from September
to October,
remaining on the
tree in winter
■ dispersal by
frugivorous
vertebrates
73 Ligustrum vulgare
Distribution and Ecology
Sex expression
Ligustrum
vulgare L.
EN: common privet
EL: αγριομυρτιά
ES: aligustre
FR: troène commun
IT: ligustro
PT: alfenheiro
Oleaceae
Variation and Hybridisation
Differences in the amount of fruit production, fruit size,
and number of seeds per fruit have been observed be
tween individuals of these species (Obeso and Grubb,
1993). These differences may affect the genetic varia
tion of collected seed lots; so similar amounts of fruit
and seeds should be collected from each individual.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from September
to December
■ gathering by hand or
using longhandled tools
sequence for fleshy
fruits
■ seed weight / kg
fruit: 66290 g
■ purity: 90100 %
■
Germination under controlled conditions
Pregermination treatments
■
prechilling (812 weeks)
Nursery production
Sowing season
■
autumn, without treatment, or early
spring, with treatment
Vegetative propagation
Type of cutting
■
■
hardwood
semihardwood
Position along
the shoot
indifferent
basal
Conditions
■
Germination Viability
20 / 10 ºC
■
Nursery practice
bareroot: 50 g/m2; circumfe
rence up to 46 cm or total
height up to 80100 cm
3
■ forestpot 300 cm : 1/0
■ container 3.5 l: 1/1
■
Number of
internodes Size
20 cm
10 cm
Ligustrum vulgare 74
Common privet is easily reproduced by vegetative prop
agation. It is advisable to plant the cutting directly in
the container (Mac Cárthaig and Spethmann, 2000) at
the end of winter to prevent frost damage. Hansen and
Kristiansen (2000) recommend collecting semihard
T: 4 ºC
MC: 48 %
■ airtight container
825 g
7496 %
Emergence
■
the first spring
Collecting time
winter
July to September
Auxin
concentration
none or 0.5 %
none or 0.5 %
wood cuttings at the end of summer, because the root
ing capacity of material collected in October or later
diminishes rapidly. It may be possible to propagate
privet using root cuttings, since it is a species that is
able to produce root suckers naturally.
References
General references
Specific references
Amaral Franco J do (1972) Ligustrum. In: Tutin TG et al. (eds).
Flora Europaea. Vol 3. Cambridge University Press, Cambridge
Hansen J, Kristiansen K (2000) Root formation, bud growth
and survival of ornamental shrubs propagated by cuttings on
different planting dates. Journal of Horticultural Science and
Biotechnology 75:568574
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Obeso JR, Grubb PJ (1993) Fruit maturation in the shrub Ligus
trum vulgare (Oleaceae): lack of defoliation effects. Oikos
68:309316
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di
alberi e arbusti della flora mediterranea. ANPA, Roma
75 Ligustrum vulgare
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Liquidambar
orientalis Mill.
Altiginaceae
EN: oriental sweet gum
EL: υγραδάμπαρη
ES: liquidámbar oriental
FR: liquidambar oriental, copalme d'Orient
IT: liquidambar orientale
PT: liquidâmbaroriental
Distribution and Ecology
General distribution: Western Asia
Oriental sweet gum grows on dry slopes and in humid
areas such as riverside zones, marshy places and the
bottom of valleys. Optimal growth is on deep, moist,
nutrientrich soils.
Mediterranean region: Greece (Rodhos), Turkey
Diagnostic traits
Liquidambar orientalis is a deciduous tree that grows to
a height of 30 to 35 m. Its palmatifid leaves have 5
lobes and finely sinuatedentate or serrate margins.
They are glabrous or, more rarely, exhibit sparse tufts of
hair beneath, at the base of the main veins. The prin
Reproductive biology
Sex expression
■
monoecy
Flowering
small flowers
clustered in globose
inflorescences, male
inflorescences in
terminal racemes,
solitary female
inflorescences
■ from March to May
■
Liquidambar orientalis 76
Variation and Hybridisation
Pollination
■
cipal lobes of the leaves usually have secondary lobes.
This trait makes it easy to distinguish this species from
L. stryraciflua, an American species widely used for or
namental purposes.
entomophilous
Two varieties are described: L. orientalis var. orientalis
and L. orientalis var. integriloba. In the latter the lobes
Fruiting
Ripening
■
■
capsules arranged
helically in woody
infructescence, 25
to 30 capsules in
each
■ infructescence
2.53 cm
from November
to December
■ dispersal by wind
of the leaves are not divided, while in the former they
are. In addition, two morphological types have been
distinguished in relation to the ability to produce bal
sam: oilproducing trees are smaller and have longer
branches and larger barkscales than nonproducers
(Alan and Kaya, 2003). It would seem there are certain
coldadaptation differences between populations lo
cated at altitudes of less than four hundred metres
(plain sweet gum) or more than four hundred meters
(mountain sweet gum) (Alan and Kaya, 2003).
Seed propagation
Liquidambar orientalis is usually produced from seed;
however, no data have been found regarding its pro
duction, possibly due to its restricted distribution and
more or less local use. As a general guideline, data for
Liquidambar styraciflua are supplied. Nevertheless, it
This species has a greater morphological and genetic
likeness to L. stryraciflua L. than to eastern Asian
sweet gum species (Hoey and Parks, 1991; Ickert
Bond et al., 2005; IckertBond and Wen, 2006). It may
hybridise with the American taxon and produce indi
viduals with a higher number of seeds per fruit (San
tamour, 1972).
should be borne in mind that although the number of
seeds per fruit is very similar in both plants, the seed
length is greater in L. orientalis (IckertBond et al.,
2005), so the weight per kilo of fruit and per 1,000
seeds would be higher than the values indicated.
Seed handling and storage
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from November to
December
■ climbing or using
longhandled tools
sequence for
dehiscent fruits,
natural drying
■ seed weight / kg fruit:
3090 g (L. styraciflua)
■ purity: 9095 %
Oriental sweet gum produces fruit every year. Abun
dant crops, however, are more common every three
years (Alan and Kaya, 2003). Infructescences are col
47 g (L. styraciflua)
lected when their green colour loses its brightness and
starts to turn yellowish.
Germination under controlled conditions
Pregermination treatments
■
prechilling (4–6 weeks)
Nursery production
Sowing season
■
autumn, without treatment, or
spring, with treatment
Conditions
■
30 / 20 ºC
Nursery practice
■
bareroot: 100 g/m2
Seeds are very sensitive to dehydration during germi
nation, so the substrate must be kept moist.
T: 4 ºC
MC: 1015 %
■ airtight container
(L. styraciflua)
■
Germination Viability
■
5070 % (L. styraciflua)
Emergence
■
the first spring
77 Liquidambar orientalis
Tolerance to desiccation: ORTHODOX
Vegetative propagation
Type of cutting
■
semihardwood
Position along
the shoot
basal or middle
Number of
internodes Size
23
No references to vegetative propagation of Liqui
dambar orientalis have been found, although there are
studies for L. styraciflua and L. formosana, which are
species of marketing interest. Both will reproduce by
vegetative propagation, although with difficulty, using
semihardwood cuttings: survival rates are always below
60%, even under optimal conditions (He et al., 2004;
Sutter and Barker, 1985).
References
General references
Catalán G (1991) Semillas de árboles y arbustos forestales.
Ministerio de Agricultura Pesca y Alimentación. ICONA,
Madrid
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns Ge
hölzvermehrung. Parey Buchverlag, Berlin
Peşmen H (1972) Liquidambar L. In: Davis PH (ed). Flora of
Turkey and East Aegean Islands. Vol 4. University Press,
Edinburgh
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Specific references
Alan M, Kaya Z (2003) EUFORGEN Technical Guidelines for ge
netic conservation and use for oriental sweet gum (Liq
uidambar orientalis). International Plant Genetic Resources
Institute, Rome
Brand MH (1990) Sweetgum tissue culture. Combined Proce
edings of the International Plant Propagators Society 40:586
590
Brand MH, Lineberger RD (1991) The effect of leaf source and
developmental stage on shoot organogenic potential of swe
etgum (Liquidambar styraciflua L.) leaf explants. Plant Cell Tis
sue and Organ Culture 24:17
Collecting time
summer
Auxin
concentration
1%
In vitro propagation seems a more effective alternative.
Erdag and Emek (2005) managed to regenerate adult
individuals of oriental Liquidambar from axillary buds.
There are several references to micropropagation of
the American and Asian species of Liquidambar (Brand,
1990; Brand and Lineberger, 1991; Durkovich et al.,
2005).
Durkovic J, Pichler V, Lux A (2005) Micropropagation with a
novel pattern of adventitious rooting in Formosan sweetgum.
Canadian Journal of Forest Research 35:27752780
Erdag B, Emek Y (2005) In vitro adventitious shoot regenera
tion of Liquidambar orientalis Miller. Journal of Biological
Sciences 5:805808
He GP, Chen YT, Luo WJ, Zhang JZ, Feng JM, Xu YQ (2004)
Study on the technical of cutting propagation of tender
branch for broadleaf tree species. Forest Research, Beijing
17:810814
Hoey MT, Parks CR (1991) Isozyme divergence between East
ern Asian, North American, and Turkish species of Liquidambar
(Hamamelidaceae). American Journal of Botany 78:938947
IckertBond SM, Pigg KB, Wen J (2005) Comparative in
fructescence morphology in Liquidambar (Altingiaceae) and
its evolutionary significance. American Journal of Botany
92:12341255
IckertBond SM, Wen J (2006) Phylogeny and biogeopraphy
of Altiginaceae: Evidence from combined analysis of five non
coding chloroplast regions. Molecular Phylogenetics and Evo
lution 39: 512528
Santamour FS (1972) Interspecific hybridization in Liqui
dambar. Forest Science 18:2326
Sutter EG, Barker PB (1985) In vitro propagation of mature Li
quidambar styraciflua. Plant Cell Tissue and Organ Culture
5:1321
Liquidambar orientalis 78
EN: honeysuckle
EL: αγιόκλημα
ES: madreselva
FR: chèvrefeuille
IT: caprifoglio
PT: madressilva
Lonicera etrusca G. Santi
Lonicera implexa Aiton
Lonicera implexa
Lonicera etrusca
Caprifoliaceae
General distribution: Southwestern and Southeastern
Europe, Western Asia, Northern Africa
Lonicera etrusca
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia,
BosniaHerzegovina, Montenegro, Albania, Greece (incl.
Kriti), Cyprus, Turkey, Syria, Lebanon, Israel, Tunisia, Al
geria. Morocco
Lonicera implexa
Mediterranean region: Portugal, Spain (incl. Baleares),
France (incl. Corse), Italy (incl. Sardegna, Sicilia), Cro
atia, BosniaHerzegovina, Montenegro, Albania, Gre
ece, Tunisia, Algeria, Morocco
Lonicera etrusca and L. implexa are thermophile hon
eysuckles with a wide distribution in the European
Mediterranean region. Other honeysuckles that require
cooler environments, like L. periclymenum, may form
part of the lianoid layer of riparian vegetation in
Mediterranean areas. All these species also grow in
hedges, scrub and open forests with a certain level of
humidity.
79 L. etrusca - L. implexa
Distribution and Ecology
Diagnostic traits
Both species are woody climbers with decussate leaves,
the first pair connate below the inflorescence, and
flowers arranged in terminal glomerules. L. implexa is
perennial, with coriaceous leaves and sessile inflores
cences, while L. etrusca is deciduous, with pedunculate
inflorescences that are sometimes accompanied by an
other two lateral glomerules. Both species are easily
differentiated from L. periclymenum because the top
leaves of the later have a short petiole. L. implexa, with
two to nine flowers per cluster, must not be confused
with L. splendida, a perennial Hispanic endemism that
Reproductive biology
Sex expression
■
hermaphroditism
Flowering
yellowishwhite
flowers, often
tinged with purple
■ from May to August
■
has flowers with longer styles and stamens. Another
climber that is similar, although found in cooler cli
mates, is L. caprifolium L., which has sessile inflores
cences with connate leaves below them; it is
deciduous, and thus easily identifiable in winter.
It is not advisable to employ L. japonica in restoration
projects. This is an Asian species that is very often used
in gardening and is considered invasive. Its flowers,
arranged in pairs on axillary peduncles, are initially white
and turn yellow when mature, and its fruits are blue.
Pollination
■
entomophilous
Fruiting
Ripening
■
■
■
reddish berry
48 mm
from September
to October,
sometimes later
■ dispersal by
frugivorous
vertebrates
Variation and Hybridisation
There is no information on intraspecific variation and
hybridisation for these taxa.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from September
to October
■ gathering by hand
sequence for fleshy
fruits
■ seed weight / kg
fruit: 3076 g (L.
etrusca); 118157 g
(L. implexa)
■ purity: 9599 %
L. etrusca - L. implexa 80
Honeysuckle crops can suffer great losses due to at
tacks by lice, which vary in severity from year to year
(Jordano, 1990). Honeysuckle fruits should be collected
711 g (L. etrusca)
1114 g (L. implexa)
T: 0 ºC a 4 ºC
CH: 48 %
■ envase hermético
■
as soon after ripening as possible to reduce losses to
birds.
Germination under controlled conditions
■
■
mechanical scarification
scarification with concentrated
sulphuric acid (1020 minutes)
Conditions
■
20 / 10 ºC; 20 ºC
Although prechilling (48 weeks) is recommended for
Lonicera species because they show some embryo dor
mancy, these two Mediterranean species appear to
Nursery production
Sowing season
■
autumn, without treatment, or
spring, with treatment
Vegetative propagation
Type of cutting
softwood
hardwood
■ root
■
■
Position along
the shoot
basal or middle
basal or middle
Germination Viability
■
have seedcoatconnected dormancy and scarification
is enough to germinate the seeds.
Nursery practice
■
Emergence
forestpot 300 cm3:
1/0 or 2/0
Number of
internodes Size
12
10 cm
10 cm
Abundant information on vegetative propagation of
honeysuckles exists, since the genus Lonicera includes
several species and hybrids of ornamental interest
which are produced almost exclusively by this method.
However, there is little specific information on L. im
plexa and L. etrusca. Honeysuckles are usually propa
gated from softwood cuttings collected in summer. The
best time to harvest is from June onwards (Cabot et al.,
2002), since spring cuttings do not give good results.
Mac Cárthaig and Espethmann (2000) recommend se
rial cutting propagation for rejuvenation of the mate
rial. It is advisable to collect material from the middle
References
General references
Bolòs O de, Vigo J (1995) Flora dels Països Catalans. Vol III.
Editorial Barcino, Barcelona
Browicz K (1976) Lonicera L. In: Tutin TG et al. (eds). Flora Eu
ropaea. Vol 4. Cambridge University Press, Cambridge
7097 %
■
the first spring; 58 weeks after
spring sowing
Collecting time
summer
winter
winter
Auxin
concentration
none or < 0.5 %
none or < 0.5 %
none or < 0.5 %
and basal parts of the stems or sets in a mist environ
ment (Podkopaev, 1987). It is also possible to use hard
wood (Albrecht and Schulze, 1980) and root cuttings
(Götsche, 1978), although these methods are used less
frequently.
There are several references to in vitro experiments to
propagate other species of the genus Lonicera (Kahru,
2003; Boonnour et al., 1988; Georges et al., 1993),
which may provide an approach for propagating the
species discussed in this guide.
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Gálvez A, Navarro RM (2001) Manual para la identificación y
reproducción de semillas de especies vegetales autóctonas de
Andalucía. Vol II. Consejería de Medio Ambiente, Junta de An
dalucía, Sevilla
81 L. etrusca - L. implexa
Pregermination treatments
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di
alberi e arbusti della flora mediterranea. ANPA, Roma
Ruiz Téllez T, Devesa JA (2007) Lonicera L. In: Castroviejo S
(coord). Flora Ibérica. Vol 15. CSIC, Madrid
Specific references
monales. In: Cermeño P (ed). I Jornadas Ibéricas de Plantas
Ornamentales. Junta de Andalucia. Consejeria de Agricultura
y Pesca, Sevilla
Georges D, Chenieux JC, Ochatt SJ (1993) Plant regeneration
from agedcallus of the woody ornamental species Lonicera
japonica cv. “Hall’s Prolific”. Plant Cell Reports 13:9194
Göttsche D (1978) Vermehrung einheimischer Straucharten
durch Wurzelschnittlinge. Forstarchiv 49:3336
Albrecht HJ, Schulze G (1980) Vermehrung von Ziergeholzen
durch Steckholz in Plastfolienzelten. Gartenbau 27:122124
Jordano P (1990) Biología de la reproducción de tres especies
del género Lonicera (Caprifoliaceae) en la Sierra de Cazorla.
Anales del Jardín Botánico de Madrid 48:3152
Boonnour K, Wainwright H, Hicks RGT (1988) The micropro
pagation of Lonicera periclymenum L. (honeysuckle). Acta Hor
ticulturae 226:183189
Karhu ST (2003) Performance of Lonicera microcuttings as af
fected by mineral nutrients and genotype. Acta Horticulturae
616:181184
Cabot P, Llauradó M, Busquets M (2002) Estudio del enraiza
miento de estaquillas de Teucrium polium spp. capitatum (L.)
Arc. y Lonicera implexa Ait. en diferentes concentraciones hor
Podkopaev AA (1987) Propagation of ornamental species of
Lonicera by green cuttings. Lesnoe Khozyaistvo 1:6566
L. etrusca - L. implexa 82
Myrtus
communis L.
EN: myrtle, common myrtle
EL: μυρτιά
ES: mirto
FR: myrte
IT: mirto
PT: murta
Myrtaceae
Distribution and Ecology
General distribution: Southwestern and Southeastern
Europe, Western Asia, Northern Africa, Macaronesia
Mediterranean region: Portugal, Spain (incl. Baleares),
France (incl. Corse), Italy (incl. Sicilia), Croatia, Bosnia
Herzegovina, Montenegro, Albania, Greece (incl. Kriti),
Cyprus, Turkey, Syria, Lebanon, Israel, Libya, Tunisia,
Algeria, Morocco
The common myrtle is a thermophile species that is
typical of the Mediterranean maquis in areas of coastal
influence. It prefers noncompact soil with a certain
level of humidity and thus can frequently be found next
to rivers, in the bottoms of valleys and in shady areas.
It grows more frequently on acidreaction soils. It re
sprouts vigorously from stumps after fire or cutting.
Diagnostic traits
Myrtus communis is a perennial, highly aromatic shrub
with opposite, decussate leaves, ovatelanceolate,
acute, attenuate at base.
Sex expression
■
hermaphroditism
Flowering
white flowers,
clustered in panicles
■ from May to August,
sometimes also in
autumn
■
Pollination
■
■
entomophilous
selfcompatible
Fruiting
Ripening
■
■
bluishblack, rarely
whitecreamy,
ellipsoidal to
subglobose berry
■ 610 mm
from October to
January
■ dispersal by
frugivorous
vertebrates
83 Myrtus communis
Reproductive biology
Variation and Hybridisation
Genetic studies performed with isoenzymes reveal a
high level of variation within populations, as well as
between populations that are distant from each other
(Messaoud et al., 2006).
Some authors have defined a tarentina subspecies,
whose leaves are described as smaller. This subspecies
may be a naturalised variety which was widely culti
vated in the past to make use of the plant’s many dif
ferent applications.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
late autumn
gathering by hand
sequence for fleshy
fruits
■ seed weight / kg
fruit: 30125 g
■ purity: 98100 %
There is great individual variation in the production of
fruit and significant interannual fluctuations also occur
■
prechilling (38 weeks)
Conditions
■
Nursery production
Sowing season
■
autumn, without treatment, or
spring, with or without treatment
Myrtle seedlings are very sensitive to cold.
Germination Viability
20 ºC
Myrtle seeds do not need treatments, but prechilling
could be useful to accelerate germination and make it
more homogeneous. There are no significant differen
■
■
8098 %
ces in seed germination capacity between individuals
with bluish or white fruits (Traveset et al., 2001).
Nursery practice
■
■
(Cani, 1996; Traveset et al., 2001; Mulas and Fadda,
2004).
Germination under controlled conditions
Pregermination treatments
T: 4 ºC
MC: 48 %
■ airtight container
27 g
forestpot 300 cm3: 1/0
container 3.5 l: 1/1
Emergence
■
the first spring; complete within
3 months
Myrtus communis 84
Type of cutting
■
■
semihardwood
hardwood
Position along
the shoot
basal or middle
basal or middle
Number of
internodes Size
23
15 cm
There is abundant information about myrtle propaga
tion because of the growing interest in its essential oils.
Very good results have been obtained using softwood
cuttings harvested from rejuvenated parent plants in
July or August, achieving 90% rooting success (Pignati
and Crobeddu, 2005). Cutting in summer requires a
misted environment (Cervelli, 2005; Scortichini, 1986).
Klein et al. (2000) recommend collecting cuttings in
November or December and planting at an ambient
temperature of 20 °C with bed heating, since these
conditions produce the best results (70%). In this
species, substrate mixtures of straw, peat and coconut
fibre (Crobeddu and Pignati, 2005) or peat and perlite
(1:1) (De Vita and Lauro, 2004) have been used. Myrtle
References
General references
Campbell MS (1968) Myrtus L. In: Tutin TG et al. (eds). Flora
Europaea. Vol 2. Cambridge University Press, Cambridge
Catalán G (1991) Semillas de árboles y arbustos forestales.
Ministerio de Agricultura Pesca y Alimentación. ICONA,
Madrid
GarcíaFayos P (coord) (2001) Bases ecológicas para la reco
lección, almacenamiento y germinación de semillas de espe
cies de uso forestal en la Comunidad Valenciana. Banc de
Llavors Forestals, Generalitat Valenciana, Valencia
Paiva J (1997) Myrtus L. In: Castroviejo S et al. (eds). Flora
Ibérica. Vol 8. CSIC, Madrid
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea. ANPA, Roma
Specific references
Cani MR (1996) Osservazioni sulla biologia e valutazione della
biodiversitá naturale per la domesticazione del Myrtus com
munis. Tesis de licenciatura. Facoltá di Agraria. Universitá degli
Srudi di Sassari, Sassari
Cervelli C (2001) Una collezione di mirto per pensare al mer
cato. Colture Protette 30:5962
Collecting time
summer
winter
Auxin
concentration
0.5 %
0.5 %
shows a very high individual variation in the ability to
root (Cervelli, 2001; Mulas and Cani, 1996). Acclimati
zation of the rooted material is an extremely sensitive
stage in this species; the greatest losses occur during
this phase (Frau et al., 2001; Milia et al., 1996).
Myrtle is successfully regenerated from axillary buds
(KhoshKhui et al., 1984; Nobre, 1994; Ruffoni et al.,
2003) and meristems (Frau et al., 2001; Morini et al.,
2002). Also, the proportion of losses in the acclimati
zation phase has been successfully reduced to 3% in
plants obtained through in vitro culture (Hatzilazarou
et al., 2003).
Cervelli C (2005) La specie arbustive della macchia medite
rranea. Un patrimonio da valorizzare. Collana Sicilia Foreste
26:39154
Crobeddu S, Pignatti G (2005) Propagazione per talea di spe
cie mediterranee. Prove di substrato. Sherwood Foreste ed Al
beri Oggi 114:2731
De Vita M, Lauro P (2004) Influenza dei substrati sull’accres
cimento di genotipi di mirto coltivato in vaso. Atti VII Giornate
Scientifiche SOI, 46 maggio 2004, Napoli
Frau A, Cadinu M, Repetto A, Zedda A (2001) Micropropaga
zione di cinque cloni di mirto sardo. Informatore Agrario
57:6567
Hatzilazarou S, Grammatikos H, Economou AS, Rifaki N, Ralli
P (2003) Rooting in vitro and acclimatization of Myrtus com
munis microcuttings. Acta Horticulturae 616:259264
KhoshKhui M, Shekafandeh A, Azarakhsh H (1984) Micro
propagation of myrtle. Scientia Horticulturae 22:139146
Klein JD, Cohen S, Hebbe Y (2000) Seasonal variation in roo
ting ability of myrtle (Myrtus communis L.) cuttings. Scientia
Horticulturae 83:7176
Messaoud C, Khoudja ML, Boussaid M (2006) Genetic diversity
and structure of wild Tunisian Myrtus communis L. (Myrtaceae)
populations. Genetic Resources and Crop Evolution 53:407–417
85 Myrtus communis
Vegetative propagation
Milia M, Pinna ME, Satta M, Scarpa GM (1996) Propagazione
del mirto (Myrtus communis L.) mediante l’uso di tecniche di
verse. Rivista Italiana EPPOS 19:117123
Pignatti G, Crobeddu S (2005) Effects of rejuvenation on cut
ting propagation of Mediterranean shrub species. Foresta
2:290295 (online URL: http://www.sisef.it/)
Morini S, Frediani F, Onofrio CD (2002) Indagini sulla micro
propagazione del mirto. Italus Hortus 9:4148
Ruffoni B, Airo M, Fascella G, Mascarello C, Zizzo G, Cervelli
C (2003) Rooting and acclimatization of ornamental myrtle
genotypes. Acta Horticulturae 616:255258
Mulas M, Cani MR (1996) Variability of rooting ability of soft
wood cuttings in myrtle germplasm. Beitrage zur Zuchtungs
forschung Bundesanstalt fur Zuchtungsforschung an
Kulturpflanzen 2:191194
Mulas M, Fadda A (2004) First observations on biology and
organ morphology of myrtle (Myrtus communis L.) flower.
Agricoltura Mediterranea 134:223235
Nobre J (1994) In vitro shoot proliferation of Myrtus communis
L. from fieldgrown plants. Scientia Horticulturae 58:253258
Scortichini M (1986) Il mirto. Rivista di Frutticoltura e di Or
tofloricoltura 48:4753
Traveset A, Riera N, Mas RE (2001) Ecology of the fruitcolor
polymorphism in Myrtus communis and differential effect of
birds and mammals on seed germinatrion and seedling
growth. Journal of Ecology 89:749760
Myrtus communis 86
General distribution: Southwestern and Southeastern
Europa, Western Asia, Arabian Peninsula, Indian Sub
continent, China, Northern, West Tropical and North
east Tropical Africa
Mediterranean region: Portugal, Spain (incl. Balea
res), France (incl. Corse), Italy (incl. Sardegna and Si
cilia), Croatia, Albania, Greece (incl. Kriti), Cyprus,
Diagnostic traits
The oleander is a perennial bush that can reach 4 to 6
m in height. Its leaves are lanceolate, with a conspi
Reproductive biology
■
hermaphroditism
Flowering
pink flowers,
clustered in corymbs
■ from March
to October
■
Pollination
■
■
Turkey, Syria, Lebanon, Israel, Libya, Tunisia, Algeria,
Morocco
The oleander is a sunloving species that grows in tem
perate climates. It can withstand floods and prolonged
periods of drought. In the Mediterranean region it is
found next to temporary and permanent watercourses,
often dominating the landscape.
cuous whitish main vein, coriaceous and glabrous.
entomophilous
selfcompatible
Fruiting
Ripening
■
■
■
double follicle
816 cm long
from December
to March
■ dispersal by wind
87 Nerium oleander
Distribution and Ecology
Sex expression
Nerium
oleander L.
EN: oleander
EL: πικροδάφνη
ES: adelfa
FR: laurierrose
IT: oleandro
PT: loendro
Apocynaceae
Variation and Hybridisation
It is sometimes difficult to establish whether popula
tions are introductions or not, since this species has
been cultivated widely and naturalises easily. There are
many commercial varieties of oleander, with flowers of
different sizes, colours and shapes, which are propa
gated vegetatively for ornamental purposes. Pagen
(1988) suggests that individuals with very aromatic
double flowers are cultivated varieties from the eastern
distribution range of the species that were introduced
in the 17th century, since the Mediterranean pheno
types have simple, nonaromatic flowers. The use of
Asiatic material should be avoided in restoration proj
ects in Mediterranean environments, where materials
of local provenance must always be employed.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
from January to February
gathering by hand
sequence for
dehiscent fruits
■ seed weight / kg
fruit: 68121 g
■ purity: 9099 %
In spite of its pollination problems, this species pro
duces a large number of seeds per plant; the number of
flowers per individual and seeds per fruit is exceedingly
high. This allows a great many seeds to be collected
from each plant. There are interannual fluctuations,
■
without treatment
Nursery production
Sowing season
■
spring, without treatment
Vegetative propagation
Type of cutting
■
Nerium oleander 88
■
hardwood
semihardwood
Conditions
■
Position along
the shoot
basal or middle
terminal
20 ºC
Nursery practice
■
■
forestpot 300 cm3: 1/0
container 3.5 l: 1/1
Number of
internodes Size
1520 cm
12
■
however, and seed production also varies between in
dividuals (Herrera, 1991). It is therefore advisable to at
tempt to balance maternal contributions and collect
from many different individuals in order to increase the
genetic variation of the seed lot.
Germination under controlled conditions
Pregermination treatments
T: 4 ºC
MC: 48 %
■ airtight container
24 g
Germination Viability
■
8897 %
Emergence
■
710 days after sowing
Collecting time
DecemberFebruary
JulyAugust
Auxin
concentration
none or < 0.5 %
none or < 0.5 %
References
General references
Markgraf F (1972) Nerium L. In: Tutin TG et al. (eds). Flora
Europaea. Vol 3. Cambridge University Press, Cambridge
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di
alberi e arbusti della flora mediterranea. ANPA, Roma
Specific references
García España V (1998) Producción de Adelfa. In: Ballester
Olmos JF (ed). Producción de Plantas Ornamentales. Escuela
Universitaria de Ingeniería Técnica Agrícola de Valencia,
Valencia
Hatzilazarou S, Ttooulos C, Economou AS, Rifaki N, Ralli P
(2003) In vitro and ex vitro rooting and plantlet acclimatiza
tion in Nerium oleander. Acta Horticulturae 616:221225
require protected conditions and bed heating is critical
(Jiménez, 1978; Ochoa et al., 2004). When using semi
woody cuttings, two or three leaves are left on the upper
knot; these leaves may be cut in half in order to reduce
the transpiration surface (Standardi and Mariani, 1994).
Cuttings prepared in the summer should be set out under
mist in a substrate that allows good drainage (García Es
paña, 1998; Ochoa et al., 2003).
In vitro propagation from leaves has been carried out
successfully (Santos et al., 1994), with acclimatisation
rates of up to 90 % (Roncasaglia et al., 2002). Hatzi
lazarou et al. (2003) have performed ex vitro rooting
trials using microcuttings.
nesis in oleander cuttings. Proceedings of the International
Symposium On the Horizons of Using Organic Matter Subs
trates in Horticulture 608:101106
Ochoa J, Bañón S, Fernández JA, Franco JA, MartínezSánchez
JJ (2004) Rooting medium temperature and carbohydrates af
fected oleander rooting. Acta Horticulturae 659:239244
Pagen FJJ (1988) Oleandres. Nerium L. and the oleander cul
tivars. Series of revision of Apocynaceae. Part XX. Agricultural
University Wageningen Papers 872, The Netherland
Pal D, Gupta SK, Afroz N, Singh C (1988) Regeneration of stem
cuttings of Nerium oleander Linn. as influenced by indole ace
tic acid and planting posture. Advances in Plant Sciences
1:219222
Patil AA, Shirol AM (1991) Studies on rooting of oleander cut
tings. South Indian Horticulture 39:4853
Herrera J (1991) The reproductive biology of a riparian
Mediterranean shrub, Nerium oleander L. (Apocynaceae).
Botanical Journal of the Linnean Society 106:147172
Rocha SC, Quisen RC, Queiroz JA, Zufellato KC (2004) Propa
gaçao vegetativa de espirradeira pela tecnica da estaquia.
Scientia Agraria 5:7377
Jiménez R (1978) Ensayo de enraizamiento de esquejes de Ne
rium oleander variegatum. Informaciones de Floricultura y
Plantas Ornamentales 10:1922
Roncasaglia R, Dradi G, Baggio G (2002) Utilizzo della coltura
in vitro per l’ottenimento di piante di oleandro (Nerium ole
ander L.) ad elevato accestimento. Italus Hortus 9:7375
Kose H, Kostak S (2000) Panasal zakkumun (Nerium oleander
L. cv. Variegata) celikle cogaltlmas ve paclobutrazolun buyume
ve ciceklenmeye etkileri. Anadolu 10:3142
Ochoa J, Bañón S, Fernández JA, Franco JA, González A (2003)
Influence of cutting position and rooting media on rhizoge
Santos I, Guimaraes I, Salema R (1994) Somatic embryogene
sis and plant regeneration of Nerium oleander. Plant Cell, Tis
sue and Organ Culture 37:8386
Standardi A, Mariani A (1994) Indagine sulla propagazione per
talea dell’oleandro. Colture Protette 23:7983
89 Nerium oleander
Usually, for ornamental plants, oleander is propagated
by cuttings. Some authors achieve better results with
lignified material obtained in the winter than with ma
terial collected in the summer (Jiménez, 1978; Kose and
Kostak, 2000; Patil and Shirol, 1991). However, other
studies recommend setting out cuttings in the spring,
using new material, and advise against doing so in win
ter (García España, 1998; Ochoa et al., 2004; Standardi
and Mariani, 1994). Either method achieves results of
over 90%. Treatment of oleanders with auxins does not
improve rooting rates and may even be counterproduc
tive (Jiménez, 1978; Pal et al., 1988; Patil and Shirol,
1991; Rocha et al., 2004). Woody cuttings should be 1 to
2 cm in diameter and must be defoliated. These cuttings
Pistacia
lentiscus L.
Distribution and Ecology
General distribution: Southwestern and Southeast
ern Europe, Western Asia, Northern Africa,
Macaronesia
Mediterranean region: Portugal, Spain (incl. Baleares),
France (incl. Corse), Italy (incl. Sardegna and Sicilia),
Croatia, BosniaHerzegovina, Montenegro, Albania,
Diagnostic traits
Pistacia lentiscus is a perennial shrub, 1 to 3 m in
height, or sometimes a small tree of up to 6 m. Its
leaves are composed of 2 to 12 oblonglanceolate or
elliptic leaflets on a winged rachis. It is easily distin
guished from other species of the genus that are found
Reproductive biology
Sex expression
■
dioecy
Flowering
small reddish or
yellowish flowers,
clustered in racemes
■ from March to May
■
Anacardiaceae
EN: mastic tree
EL: σχίνος
ES: lentisco
FR: lentisque
IT: lentisco
PT: aroeira
Pollination
■
Greece (incl. Kriti), Cyprus, Turkey, Syria, Lebanon, Is
rael, Libya, Tunisia, Algeria, Morocco
The mastic tree grows on all types of soils. It is a ther
mophile species that is found in abundance in garrigues
and open forests, in areas that are not excessively dry.
It resprouts from stumps.
in the Mediterranean region and the Middle East (P. at
lantica, P. palaestina, P. terebinthus and P. khinjuk) in
that it is the only perennial, and it can be included in
a separate group on the basis of characterisation using
molecular techniques (GolaGoldhirsh et al., 2004).
anemophilous
Fruiting
Ripening
■
■
blackish globose
drupe
■ 47 mm
from October to
December
■ dispersal by
frugivorous
vertebrates
Pistacia lentiscus 90
Variation and Hybridisation
The mastic tree can be distinguished by molecular
techniques from the semideciduous hybrid it forms
with P. terebinthus (P. × saportae Burnat.). While this
hybrid exhibits larger fruits and a narrower leaf rachis
wing, the morphological differences are sometimes not
very marked, making identification difficult (Werner et
al., 2001). It would also seem that the hybrid tends to
produce very few fruits, and when it does they are vain
or have unviable seeds (Werner et al., 2001; Montser
ratMartí and PérezRontomé, 2002).
As a precautionary measure it is advisable to use local
material for habitat restoration, since genetic varia
tions between different provenances have been estab
lished using molecular techniques (Werner et al., 2002;
Barazani et al., 2003).
Seed propagation
Seed handling and storage
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from October
to November
■ gathering by hand
sequence for fleshy
fruits
■ seed weight / kg
fruit: 60325 g
■ purity: 98100%
This species produces a great number of flowers and
fruits, but the number of fruits with viable seeds is very
low, since many flowers do not produce fruit and a
large proportion of fruits do not contain seeds
(MartínezPalle and Aronne, 2000). Collection of fruits
that are white or reddish at maturity should be avoided,
as this is a clear indication of embryo abortion or
parthenocarpy (Jordano, 1988; 1989). Black drupes
should be harvested, since the proportion of seeds in
this type of fruit is always higher than in the former
type (Verdú and GarcíaFayos, 2002). Additionally, the
■
gentle mechanical scarification
Conditions
■
■
production of fruits with seeds is highly variable within
the plants of a single population (MartínezPalle and
Aronne, 2000; Verdú and GarcíaFayos, 2002), a bian
nual fluctuation in the production of fruits and in the
proportion of seedless fruits or fruits with unviable
seeds has also been estimated, and the crop may some
times be lost (GarcíaFayos, 2001). It has been deter
mined that a balanced proportion of male and female
plants and densities of over one hundred individuals
per hectare promote good yields (Verdú and García
Fayos, 1998).
Germination under controlled conditions
Pregermination treatments
20 ºC
Germination Viability
■
Mechanical scarification is not essential, but it speeds
up germination time and homogenises emergence.
Nursery production
Sowing season
■
autumn, without treatment, or
spring, with or without treatment
Nursery practice
forestpot 300 cm3: 1/0
■ container 3.5 l: 1/1
■
T: 4 ºC
MC: 48 %
■ airtight container
1025 g
7595 %
Emergence
■
in spring; 2 to 4 weeks
91 Pistacia lentiscus
Tolerance to desiccation: ORTHODOX
Vegetative propagation
Type of cutting
■
■
hardwood
semihardwood
Position along
the shoot
terminal
terminal
Number of
internodes Size
10 cm
10 cm
Mastic, like other species of the Pistacia genus, does
not propagate easily using cuttings (Joley and Opitz,
1971). However, if this method is to be employed, the
material should be collected from rejuvenated mother
plants (Isfendiyaroglu, 2000; Pignati and Crobeddu,
2005; Viola et al., 2004). Collecting time is a key fac
tor in the success of cuttings, but the optimum period
varies according to different authors. Thus Isfendi
yaroglu (2000) and Viola et al. (2004) recommend the
months of January and February, respectively, reporting
good results (over 75%). Pignati and Crobeddu (2005),
for their part, consider the month of July to be better
References
General references
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
GarcíaFayos P (coord) (2001) Bases ecológicas para la reco
lección, almacenamiento y germinación de semillas de espe
cies de uso forestal en la Comunidad Valenciana. Banc de
Llavors Forestals, Generalitat Valenciana, Valencia
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di
alberi e arbusti della flora mediterranea. ANPA, Roma
Tutin TG (1968) Pistacia L. In: Tutin TG et al. (eds). Flora
Europaea. Vol 2. Cambridge University Press, Cambridge
Specific references
Barazani O, Dudai N, GolanGoldhirsh A (2003) Comparison
of Mediterranean Pistacia lentiscus genotypes by random am
plified polymorphic DNA, chemical and morphological analy
ses. Journal of Chemical Ecology 29:19391952
Barghchi M, Alderson PG (1983) In vitro propagation of Pista
cia species. Acta Horticulturae 131:4960
Crobeddu S, Pignatti G (2005) Propagazione per talea di specie
mediterranee prove di substrato. Sherwood Foreste ed Alberi
Oggi 114:2731
Pistacia lentiscus 92
Fascella G, Airo M, Zizzo G, Ruffoni B (2004) Prime osser
vazioni sulla coltivazione in vitro di lentisco (Pistacia lentis
cus L.). Italus Hortus 11(4):141143
Collecting time
winter
summer
Auxin
concentration
1%
1%
than April, since successful results exceed 80% in the
summer. Rooting under mist is recommended, with
basal heating, using a peat and perlite substrate (1:1)
(Crobeddu y Pignati, 2005).
In vitro propagation of mastic and other Pistacia
species has been performed successfully (Barghchi and
Alderson, 1983; Fascella et al., 2004; Gatti et al., 2004;
Onay, 2000). If vegetative propagation is desired, this
method is an alternative to the difficulties of repro
duction from cuttings.
Gatti E, Predieri S, Govoni M (2004) Coltura in vitro di piante
mediterranee autoctone: cisto, elicriso, lentisco e rosmarino.
Italus Hortus 11:135137
GolanGoldhirsh A, Barazani O, Wang ZS, Khadka DK, Saun
ders JA, Kostiukovsky V, Rowland LJ (2004) Genetic rela
tionships among Mediterranean Pistacia species evaluated
by RAPD and AFLP markers. Plant Systematics and Evolution
246:918
Isfendiyaroglu M (2000) Cutting propagation of mastic tree
(Pistacia lentiscus var. Chia Duham.). NUCIS Newsletter
9:4244
Joley LE, Opitz KW (1971) Further experiences with propaga
tion of Pistacia. Combined Proceedings of the International
Plant Propagators Society 21:6776
Jordano P (1988) Polinización y variabilidad de la producción
de semillas en Pistacia lentiscus (L.) (Anacardiaceae). Anales
del Jardín Botánico de Madrid 45:213231
Jordano P (1989) Predispersal biology of Pistacia lentiscus L.
(Anacardiaceae): cumulative effects on seed removal by birds.
Oikos 55:375386
MartínezPalle E, Aronne G (2000) Reproductive cycle of
Pistacia lentiscus (Anacardiaceae) in Southern Italy. Plant
Biosystems 134:365371
MontserratMartí G, PérezRontomé C (2002) Fruit growth
dynamics and their effects on the phenological pattern of na
tive Pistacia populations in NE Spain. Flora 197:161174
Pignatti G, Crobeddu S (2005) Effects of rejuvenation on cut
ting propagation of Mediterranean shrub species. Foresta
2:290295 (online URL: http://www.sisef.it/)
Verdú M, GarcíaFayos P (1998) Ecological causes, function and
evolution of abortion and parthenocarpy in Pistacia lentiscus L.
(Anacardiaceae). Canadian Journal of Botany 76:134141
Verdú M, GarcíaFayos P (2002) Ecología reproductiva de Pis
tacia lentiscus L. (Anacardiaceae): un anacronismo evolutivo
en el matorral mediterráneo. Revista Chilena de Historia Na
tural 75:5765
Viola F, Forleo LR, Cocozza MA (2004) Propagazione agamica
di alcune specie della macchia mediterranea. Italus Hortus
11:186190
Werner O, SánchezGómez P, CarriónVilches MA, Guerra J
(2002) Evaluation of genetic diversity in Pistacia lentiscus L.
(Anacardiaceae) from the southern Iberian Peninsula and
North Africa using RAPD assay. Implications for reafforesta
tion policy. Israel Journal of Plant Science 51:1118
Werner O, SánchezGómez P, Guerra J, Martínez JF (2001)
Identification of Pistacia x saportae Burnat. (Anacardiaceae)
by RAPD analysis and morphological characters. Scientia Hor
ticulturae 91:179186
93 Pistacia lentiscus
Onay A (2000) Micropropagation of pistachio from mature
trees. Plant Cell Tissue and Organ Culture 60:159162
Platanus
orientalis L.
Distribution and Ecology
General distribution: Southeastern Europe, Western
Asia
Mediterranean region: Italy (incl. Sicilia), Montenegro,
Albania, Greece (incl. Kriti), Cyprus, Turkey, Syria,
Lebanon, Israel
Diagnostic traits
Platanus orientalis is a deciduous tree that reaches a
height of 30 m, with rhytidome that peels off in sheets.
Leaves exhibit 3 to 5 dentate lobes that are greater in
length than in width. Female flower heads are normally
Reproductive biology
Sex expression
■
monoecy
Flowering
small flowers,
clustered in heads
■ from March to May
■
Platanaceae
EN: oriental plane tree
EL: πλάτανος η ανατολική
ES: plátano oriental
FR: platane d’Orient
IT: platano orientale
PT: plátanooriental
Pollination
■
The oriental plane grows in humid forests, valley bot
toms and riparian areas.
arranged in groups of three to six on a long peduncle.
Fruits have a more or less pyramidal or wide wedge
shaped apex.
anemophilous
Fruiting
Ripening
■
■
clavate achenes
in globose
infructescence
from October to
November,
persisting on the
tree until the
following spring
■ dispersal by wind
Platanus orientalis 94
Variation and Hybridisation
Platanus acerifolia (Aiton) Willd. is very common in
Western Europe as an ornamental species. Popularly
known as London plane or hybrid plane, its leaf char
acteristics are halfway between those of P. orientalis
and P. occidentalis L., with deep lobes that are wider
and less deep than those of Platanus orientalis, and its
infructescences are arranged in groups of two. Tests
using molecular techniques have confirmed the hybrid
origin of this species, with P. orientalis possibly acting
as the female parental species (Besnard et al., 2002).
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
sequence for dehis
cent fruits
■ seed weight / kg
fruit: 500600 g
(P. occidentalis)
■ purity: 85 %
Germination under controlled conditions
Pregermination treatments
■
prechilling (612 weeks)
Conditions
■
20 ºC to 25 ºC
Pregermination treatments are not strictly required for
oriental plane seeds, but prechilling accelerates and
Nursery production
Sowing season
■
winter, without treatment, or
spring, with treatment
■
Germination Viability
■
3040 %
homogenises germination.
Nursery practice
■
T: 7 ºC a 4 ºC
MC: 48 %
■ airtight container
24 g
bareroot: circumference up to
810 cm
Emergence
■
in spring, complete in one
month
95 Platanus orientalis
from late autumn
to winter
■ gathering by hand, using
longhandled tools, or
collecting naturally fallen
material
Vegetative propagation
Type of cutting
■
■
hardwood
semihardwood
Position along
the shoot
basal
basal
Number of
internodes Size
25 cm
10 cm
Cuttings from the basal area should be used when the
ortet is an adult plant (Nahal and Rahme, 1990; Vla
chov, 1988), whereas if it is a young plant, the part of
the stem from which the cutting is obtained does not
influence rooting capacity as much. Rejuvenation prun
ing considerably improves the results (Vlachov, 1988).
To propagate adult individuals, Arene et al. (2001) rec
ommend using twoyearold material or hardwood
mallet cuttings, making the upper cut just above the
first bud. According to Vlachov (1998), better results
are obtained from hardwood cuttings in winter (80
100%) than from semihardwood material in summer
(3060%). Cutting propagation in winter requires pro
References
General references
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns Ge
hölzvermehrung. Parey Buchverlag, Berlin
Piotto B, Di Noi A (eds.) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea. ANPA, Roma
Tutin TG (1993) Platanus L. In: Tutin TG et al. (eds). Flora Eu
ropaea. Vol 1. 2nd edn. Cambridge University Press, Cambridge
Yaltirik F (1982) Platanus L. In: Davis PH (ed). Flora of Turkey
and East Aegean Islands. Vol 7. University Press, Edinburgh
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Specific references
Arene L, Cadic A, Djulbic M, Gros A, Renoux A (2001) Multi
plication du platane par bouturage hivernal sur couche et mi
crobouturage in vitro. PHM Revue Horticole 423:2326
Platanus orientalis 96
Besnard G, Tagmount A, Baradat P, Vigouroux A, Bervillé A
(2002) Molecular approach of genetic affinities between wild
and ornamental Platanus. Euphytica 126:401–412
Collecting time
DecemberJanuary
August
Auxin
concentration
none
none
tected conditions (Grolli et al., 2005; Nahal and Rahme,
1990; Vlachov, 1988). Treatment with auxin can have
negative effects, particularly if combined with bed
heating (Grolli et al., 2005; Panetsos et al., 1994; Vla
chov, 1988).
Micropropagation of P. orientalis appears to be possi
ble as P. acerifolia has been multiplied in vitro (Grolli
et al., 2004; Liu and Bao, 2003). Likewise, P. orientalis
has regenerated satisfactorily from latent buds (Arene
et al., 2001) and from calluses that have formed on leaf
explants (Qiang et al., 2003).
Grolli PR, Morini S, Loreti F (2004) The micropropagation of
Platanus acerifolia Willd. Advances in Horticultural Science
18:121126
Grolli PR, Morini S, Loreti F (2005) Propagation of Platanus
acerifolia Willd. by cutting. Journal of Horticultural Science
and Biotechnology 80:705710
Liu G, Bao M (2003) Adventitious shoot regeneration from in
vitro cultured leaves of London plane tree (Platanus acerifo
lia Willd.). Plant Cell Reports 21:640644
Nahal I, Rahme A (1990) Le platane d’Orient (Platanus orien
talis L.) dans la region du Proche Orient. Forêt Mediterrane
ènne 12:115124
Panetsos KP, Scaltsoyiannes AV, Alizoti PG (1994) Vegetative
propagation of Platanus orientalis x P. occidentalis F1 hybrids
by stem cuttings. Forest Genetics: International Journal of Fo
rest Genetics 1:125130
Qiang FG, Ping JJ, Qing JY, Feng L (2003) In vitro efficient plant
regeneration with Platanus orientalis L. leaves as explants.
Acta Horticulturae Sinica 30:236238
Vlachov DD (1988) Vegetative propagation of sp. Platanus L.
through rooting of cuttings. Acta Horticulturae 226:375378
Salicaceae
Populus
alba L.
EN: abele, white poplar
EL: λεύκη η λεύκη
ES: álamo blanco, chopo blanco
FR: peuplier blanc, peuplier de Hollande
IT: pioppo bianco, gattice
PT: álamobranco, choupobranco
Distribution and Ecology
General distribution: Southwestern, Southeastern,
Middle and Eastern Europe, Caucasus, Western and
Middle Asia, Siberia, China, Northern Africa
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia,
BosniaHerzegovina, Montenegro, Albania, Greece (incl.
Kriti), Turkey, Syria, Lebanon, Libya, Tunisia, Algeria,
Morocco
The white poplar is a sunloving species that grows on
neutral or alkaline substrata and can withstand heavy
soils and a certain level of salinity. It gathers in dense
stands or grows scattered in low, sometimes swampy
riverine areas. In cool environments it may grow in
mixed poplar groves with Populus nigra, while in
coastal or warmer areas it becomes dominant because
it is more thermophile.
Diagnostic traits
The traits by which P. alba, P. tremula and the spontaneous
hybrid of these two species (P. x canescens Sm.) can be
distinguished are detailed in a table in an Appendix.
Sex expression
■
dioecy
Flowering
■
■
inconspicuous
flowers, clustered in
pendulous catkins
from February to
April, before leaf
development
Pollination
■
anemophilous
Fruiting
Ripening
■
■
■
oblongconic
capsule
about 4 mm
■
from March
to June
dispersal by wind
97 Populus alba
Reproductive biology
Variation and Hybridisation
The white poplar is widely used for ornamental pur
poses because of its plasticity and its attractive white
bark. P. alba cv. ‘Roumi’, commonly known as “Bol
leana”, is used a great deal because of its uniform
arrangement of branches, growing from the very base
of the tree, and its pyramidal shape. Materials from
cultivated varieties such as this should be restricted to
urban gardens and must not be used for riparian
restorations.
Preliminary analysis of Spanish populations of Populus
alba by molecular techniques reveals a strong geo
graphic pattern of genetic variation by hydrological
basins (S. GonzálezMartínez, pers. com.). Conse
quently, it is advisable to limit the use of reproductive
materials to the river basin of origin and to avoid trans
ferring materials from one basin to another.
The enormous facility with which white poplars re
sprout from the roots could lead to a situation where
populations of these trees exhibit a low level of genetic
variation, as marker studies have shown in some Sar
dinian populations, which reveal few genotypes and a
tendency to spatial grouping (Zappelli et al., 2005); this
clonal gathering is also found in Spanish populations
(S. GonzálezMartínez, pers. com). Consequently, when
collecting vegetative material of this species it is ad
visable to leave a distance of several metres between
ortets or groups of suckers and collect relatively few
materials from each individual or group of individuals.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
■
■
■
■
from March to June
using longhandled tools
or collecting naturally
fallen material
sequence for
dehiscent fruits
■ purity: 4050 %
Collection must begin when the capsules start to open;
this requires careful field supervision, since seeds are
quickly scattered by the wind. Keeping the fruitpro
ducing catkins at room temperature for 3 to 5 days al
lows the capsules to open up fully and permits seed
dehiscence. It is not necessary to remove the tufts of
hair that cover the seeds, although doing so does make
handling easier. Separation can be performed using
finemesh sieves and compressed air. Cleaning and
preparation should be carried out in less than a week,
■
without treatment
Conditions
■
0.10.6 g
T: 18 ºC
MC: 68 %
■ airtight container
■
■
since the seeds lose their viability rapidly if kept in nor
mal temperature and moisture conditions. Seeds with
a moisture content of 5 % to 8 % may be preserved for
a couple of years in airtight containers at 4 ºC to 5 ºC;
for longer periods, temperatures below 0 ºC are
recommended. Seeds that have been preserved for a
long time must be rehydrated gradually (using moist
air, for instance), as they can be damaged by excessi
vely rapid moisture uptake.
Germination under controlled conditions
Pregermination treatments
Storage
20 ºC to 25 ºC
Germination Viability
■
8595 %
Populus alba 98
Sowing season
■
spring, without treatment,
immediately after collecting
Nursery practice
bareroot: about 3,000
seeds/m2; circumference up to
68 cm or total height up to
100150 cm
3
■ forestpot 300 cm : 1/0
■ container 3.5 l: 1/1
■
The seeds should not be covered or pressed into the
substrate after sowing. The seedlings are very delicate
and are susceptible to drought during the first month.
When plants are produced in containers, it is possible
Vegetative propagation
Type of cutting
■
■
hardwood
root
Position along
the shoot
basal
The possibility of successfully reproducing white poplar
using cuttings largely depends on the ortet. Therefore,
if working with a large number of clones, a relatively
high percentage of losses should be expected (Sekawin,
1975). If working with material from individuals which
are difficult to root, results may be improved by using
any of the following methods: cut the material just
below a knot; submerge the basal end of the cuttings
in water, maintaining a temperature of 16 °C and total
darkness until the roots begin to emerge (Phipps et al.,
1983); plant cuttings with bed heating at 20 °C; or col
lect material from parent stocks established for this
purpose. Whichever the method, it requires vigorous
material from the bottom of the crown or from new
References
General references
Amaral Franco J do (1993) Populus L. In: Tutin TG et al. (eds).
Flora Europaea. Vol 1. 2nd edn. Cambridge University Press,
Cambridge
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns Ge
hölzvermehrung. Parey Buchverlag, Berlin
■
1224 h after sowing
to sow in small pots (volume up to 7075 cm3). Once
the seedlings have become established in these smaller
cells, they can be removed and transplanted into the
growth containers.
Number of
internodes Size
10 15 cm
5 10 cm
Emergence
Collecting time
FebruaryMarch
early spring
Auxin
concentration
none or 0.5 %
none
shoots. The cuttings should be between 8 and 20 mm
in diameter (Phipps and Netzer, 1981; Sabatti et al.,
2001).
Another possible propagation method is derived from
the natural ability of this species to sprout from the
root. Root segments may be used for rooting and direct
plant production or as a source of green shoots for use
as cuttings.
In vitro propagation is possible and has been carried
out successfully using catkins (Good et al., 1992, 2001),
stem segments (Sellman et al., 1989) and axillary leaf
buds (Bagnaresi and Minotto, 1982).
Nicolás JL, Iglesias S, Alía R (2001) Fichas descriptivas de es
pecies. In: García del Barrio JM et al. (coord). Regiones de
identificación y utilización de material forestal de reproduc
ción. Ministerio de Medio Ambiente, Madrid
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di
alberi e arbusti della flora mediterranea. ANPA, Roma
Soriano C (1993) Populus L. In: Castroviejo S et al. (eds). Flora
Ibérica. Vol 3. CSIC, Madrid
99 Populus alba
Nursery production
Specific references
Bagnaresi U, Minotta G (1982) Ricerche sulla micropropaga
zione di pioppi della Sez. Leuce. Annali, Accademia Italiana di
Scienze Forestali 31:239254
Bueno MA, Astorga R, Manzanera JA (1992) Micropropagación
de Populus alba ‘Siberia Extremeña’ a partir de amentos. Inves
tigación Agraria, Sistemas y Recursos Forestales 1:163171
Bueno MA, Manzanera JA, Grau JM, Sánchez N, Gómez A (2001)
Propagación in vitro de Populus tremula L. y Populus alba L. In:
Actas del I Simposio del Chopo, 911 mayo 2001, Zamora
Phipps HM, Hansen EA, Fege AS (1983) Preplant soaking of dor
mant Populus hardwood cuttings. Research Paper NC241. USDA
Forest Service, North Central Forest Experiment Station, St Paul
Phipps HM, Netzer DA (1981) The influence of collection time
and storage temperature on Populus hardwood cutting deve
lopment. Tree Planters’ Notes 32:3336
Sabatti M, Nardin F, Olivero M, Alasia F, ScarasciaMugnozza
G (2001) Propagazione vegetativa del pioppo bianco (Populus
alba) mediante talee legnose: variabilità genetica e modalità
di trattamento del materiale. Alberi e Foreste per il Nuovo Mi
llennio. In: Atti del III Congresso Nazional S.I.S.E.F, 1518
Ottobre 2001, Viterbo
Sekawin M (1975) La Génétique du Populus alba L. Annales
Forestales 6:157189
Sellmer JC, McCown BH, Haissi BE (1989) Shoot culture dyna
mics of six Populus clones. Tree Physiology 5:219227
Zapelli I, Fossati T, Patrignani G, Brundu G, Camarda I, Sala F,
Castiglione S (2005) AFLPs to assess the controversial status
of Populus alba L. of Sardinia. International workshop: The role
of biotechnology for the characterisation and conservation of
crop, forestry, animal and fishery genetic resources, 57
March, Turin, Italy (online URL http://www.fao.org/biotech/to
rino05.htm)
Populus alba 100
Distribución general: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus, West
ern and Middle Asia, Siberia, Northern Africa
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia,
BosniaHerzegovina, Montenegro, Albania, Greece,
Turkey, Tunisia, Algeria, Morocco
Diagnostic traits
P. nigra may easily be confused with genotypes origi
nated by its hybridisation with P. deltoides, which have
been widely cultivated. Traits that allow these two
Reproductive biology
■
dioecy
Flowering
■
■
inconspicuous flo
wers, clustered in
pendulous catkins
from February to
April, before leaf
development
Pollination
■
The black poplar is a sunloving species that pioneers
riverside forests and prefers alkaline to neutral reac
tion soils. It cannot withstand swampy ground for pro
longed periods of time. It grows individually or in the
form of small stands along the banks of rivers.
species and their hybrid (P. x canadensis) to be distin
guished are shown in a table in an Appendix.
anemophilous
Fruiting
Ripening
■
■
■
ellipsoidal capsule
79 mm
■
from March to
June
dispersal by wind
101 Populus nigra
Distribution and Ecology
Sex expression
Populus
nigra L.
EN: black poplar
EL: λεύκη η μαύρη
ES: chopo
FR: peuplier noir
IT: pioppo nero
PT: choupo negro
Salicaceae
Variation and Hybridisation
Three subspecies are described: P. nigra subsp. nigra,
which is found in central and eastern Europe and is
characterised by having glabrous twigs and leaves; P.
nigra subsp. betulifolia (Pursh), from western Europe,
with pubescent twigs and young leaves and noncau
date leaves on short shoots; and P. nigra subsp. caud
ina, found in the Mediterranean region, which has
hispid twigs and young leaves and caudate leaves on
short shoots.
Several varieties of poplar have been defined. Some of
them come from cultivated clones; such as the Lom
bardy poplar (P. nigra var. italica Münchh), a fastigiate
variety that may result from one or more male geno
types and has been spread widely through Europe since
the 18th century. Use of this material of uncertain ori
gin must be avoided in favour of autochthonous sub
species and genotypes.
Although the risk of backcrossing of Populus nigra with
commercial clones of P. × canadensis seems quite un
likely in some populations, due to differences in phe
nology (Gebhardt et al., 2001; Fossati et al., 2003),
overlapping floral phenology has been observed in oth
ers (Vander Broeck et al., 2003). The possibility of in
trogression of EuroAmerican hybrids with the
autochthonous species has been confirmed in differ
ent populations using molecular techniques (Vanden
Broeck et al., 2004; Pospíšková and Šálková, 2006);
these studies have shown that small isolated popula
tions of P. nigra surrounded by more or less extensive
commercial plantations are especially vulnerable (Van
den Broeck et al., 2005). Introgression of P. trichocarpa
genes (Lefèvre et al., 2002) is also possible. Collection
of seeds in this type of situation should be avoided in
favour of vegetative multiplication, in such a way as to
ensure the taxonomical identity of the propagated ma
terial and favour the use and preservation of the au
tochthonous species.
The study of spatial variations of genetic diversity
seems to show that there are no great differences be
tween regions but there are differences between pop
ulations within the same region (Legionnet and Lefèvre,
1996); these differences between populations that are
near to each other would seem to indicate limited ge
netic flow (Legionnet and Lefèvre, 1996; Imbert and
Lefèvre, 2003; Pospíšková and Šálková, 2006), in spite
of the fact that the pollen and seeds of this species are
dispersed by the wind. However, the highest levels of
diversity seem to appear at population level, even in
not very extensive populations, as seed propagation
predominates over vegetative propagation (Arens et al.,
1998; Pospíšková and Šálková, 2006). Considering the
results of genetic studies, a seedcollection unit might
cover scattered populations and individuals which are
not divided by topographical barriers that reduce pollen
flow.
It should be borne in mind that this species also repro
duces naturally by means of vegetative propagation
and that there may therefore be ramets of the same
clone with varying degrees of spatial aggregation
within a given stand or even at a distance of several
kilometres. The incidence of such natural vegetative re
production seems to vary a great deal (Legionnet et al.
1997; Arens et al., 1998; Barsoum et al., 2004), de
pending on the past history of smallscale spatial dis
turbances and on available resources in terms of the
number and age of trees. It is thus difficult to establish
simple recommendations to ensure the collection of
different clones. In these cases, molecularmarker char
acterisation should be used to differentiate genotypes
and guarantee a certain level of genetic variation using
a wellbalanced mixture of clones.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
Populus nigra 102
from March to June
using longhandled tools
or naturally fallen
material
sequence for
dehiscent fruits
■ purity: 4050 %
0.91 g
T: 18 ºC
MC: 68 %
■ airtight container
■
since the seeds lose their viability rapidly if kept in nor
mal temperature and moisture conditions. Seeds with
a moisture content of 5 % to 8 % may be preserved for
a couple of years in airtight containers at 4 ºC to 5 ºC;
for longer periods, temperatures below 0 ºC are rec
ommended. Seeds that have been preserved for a long
time must be rehydrated gradually (using damp air, for
instance), as they can be damaged by excessively rapid
moisture uptake.
Germination under controlled conditions
Pregermination treatments
■
without treatment
■
Nursery production
Sowing season
■
Conditions
spring, without treatment,
immediately after collecting
20 ºC to 25 ºC
Nursery practice
Type of cutting
■
hardwood
Position along
the shoot
indifferent
■
bareroot: circumference up to
68 cm or total height up to
100150 cm
3
■ forestpot 300 cm : 1/0
■ container 3.5 l: 1/1
Populus nigra is very easily propagated by cuttings (Da
genbach, 1997). Material collected at practically any
time of the year will grow roots, although hardwood
cuttings are the best material to use (Blake and Atkin
son, 1986; Gunes, 2000). It is advisable to use material
taken from sets or oneyearold shoots with a diame
ter of 12 to 20 mm (Holzberg, 1999).
■
1224 h after sowing
to sow in small pots (volume up to 7075 cm3). Once
the seedlings have become established in these smaller
cells, they can be removed and transplanted into the
growth containers.
Number of
internodes Size
20 30 cm
8595 %
Emergence
■
The seeds should not be covered or pressed into the
substrate after sowing. The seedlings are very delicate,
and are susceptible to drought during the first month.
When plants are produced in containers, it is possible
Vegetative propagation
Germination Viability
Collecting time
February
Auxin
concentration
none
In vitro regeneration of this species is also possible; it
has been successfully achieved with axillary and apical
buds and with stem segments (Kapusta and Skibinska,
1985; Naujoks and Wuhlisch, 2004; Noël et al., 2002).
103 Populus nigra
Collection must begin when the capsules start to open;
this requires careful field supervision, since seeds are
quickly scattered by the wind. Keeping the fruitpro
ducing catkins at room temperature for 3 to 5 days al
lows capsules to open up fully and permits seed
dehiscence. It is not necessary to remove the tufts of
hair that cover the seeds, although doing so does make
handling easier. Separation can be performed using
finemesh sieves and compressed air. Cleaning and
preparation should be carried out in less than a week,
References
General references
Amaral Franco J do (1993) Populus L. In: Tutin TG et al. (eds).
Flora Europaea. Vol 1. 2nd edn. Cambridge University Press,
Cambridge
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Nicolás JL, Iglesias S, Alía R (2001) Fichas descriptivas de es
pecies. In: García del Barrio JM et al. (coord). Regiones de
identificación y utilización de material forestal de reproduc
ción. Ministerio de Medio Ambiente, Madrid
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea. ANPA, Roma
Soriano C (1993) Populus L. In: Castroviejo S et al. (eds). Flora
Ibérica. Vol 3. CSIC, Madrid
Specific references
Arens P, Coops H, Vosman B (1998) Molecular genetic analy
sis of black poplar (Populus nigra L.) along Ditch rivers. Mo
lecular Ecology 7:1118
Barsoum N, Muller E, Skot L (2004) Variations in levels of clon
ality among Populus nigra L. stands of different ages. Evolu
tionary Ecology 18:601624
Blake TJ, Atkinson SM (1986) The physiological role of absicic
acid in the rooting of poplar and aspen stump sprouts. Physio
logia Plantarum 67:638643
Dagenbach H (1997) Praktische Vorschlage zur Nachzucht der
einheimischen Schwarzpappel (Populus nigra L.). Holzzucht
51:2326
Fossati T, Grassi F, Sala F, Castiglione S (2003) Molecular
analysis of natural populations of Populus nigra L. intermin
gled with cultivated hybrids. Molecular Ecology 12:2033
Gebhardt K, Pohl A, Vornam B (2001) Genetic inventory of
black poplar populations in the Upper Rhine floodplains: con
clusions for conservation of an endangered plant species. In:
van Dam BC and Bordács S (eds). Proceedings of the Interna
tional Symposium: Genetic diversity in river populations of
European Black Poplar. Szeksárd, Hungary, 1620 May 2001.
Verlag C. Nyomda, Budapest
Gunes T (2000) Peroxidase and IAAoxidase activities during
rooting in cuttings of three poplar species. Turkish Journal of
Botany 24:97101
Populus nigra 104
Holzberg H (1999) Propagation strategies of Populus nigra
under natural environmental conditions and artificial propa
gation in the nursery. Holzzucht 52:1416
Imbert E, Lefèvre F (2003) Dispersal and gene flow of Populus
nigra (Salicaceae) along a dynamic river system. Journal of
Ecology 91:447456
Kapusta J, Skibinska A (1985) Induction of morphogenesis and
regeneration in the callus of Populus alba L. and P. nigra L.
Journal of Tree Sciences 4:3438
Lefèvre F, Bordács S, Cottrell J, Gebhardt K, Smudlers MJM,
Vanden Broeck A, Vornam B, Van Dam BC (2002) Recommen
dations for riparia ecosystem management based on the gen
eral frame defined in EUFORGEN and results from EUROPOP
(2002). In: van Dam B, Bordács S (eds). Genetic diversity in
river populations of European Black Poplar. Csiszár Nyomda,
Budapest
Legionnet A, FaivreRampant P, Villar M, Lefèvre F (1997) Sex
ual and asexual reproduction in natural stands of Populus
nigra. Botanical Acta 110:257263
Legionnet A, Lefèvre F (1996) Genetic variation of the ripar
ian pioneer tree species Pinus nigra L. I. Study of population
structure based on isozymes. Heredity 77:629637
Naujoks G, Wühlisch G von (2004) Micropropagation of Po
pulus nigra L.: a potential contribution to gene conservation
and tree improvement. In: Koskela J, de Vries SMG, Kajba D
and von Wühlisch G (comp). Populus nigra Network, Report of
seventh (25–27 October 2001, Osijek, Croatia) and eighth me
etings (22–24 May 2003, Treppeln, Germany). International
Plant Genetic Resources Institute, Rome
Noël N, Leplé JC, Pilate G (2002) Optimization of in vitro mi
cropropagation and regeneration for Populus x interamericana
and Populus x euramericana hybrids (P. deltoides, P. tricho
carpa, and P. nigra). Plant Cell Reports 20:11501155
Pospíšková M, Šálková I (2006) Population structure and
parentage analysis of black poplar along the Moravia River.
Canadian Journal of Forest Research 36:10671076
Vanden Broeck A, Cox K, Quataert P, Van Bockstaele E, Van
Slycken J (2003) Flowering phenology of Populus nigra L., P.
nigra cv. italica and P. x canadensis Moench. and the poten
tial for natural hybridisation in Belgium. Silvae Genetica
52:280283
Vanden Broeck A, Storme V, Cottrell JE, Bockstaele E, Quataert
P, Van Slycken J (2004) Gene flow between cultivated poplars
and native black poplar (Populus nigra L.): a case study along
the river Meuse on the DutchBelgian border. Forest Ecology
and Management 197:307310
Vanden Broeck A, Villa M, Van Bockstaele E, Van Slycken J
(2005) Natural hybridization between cultivated poplars
and their wild relatives: evidence and consequences for
native poplar populations. Annals of Forest Science
62:601613
Salicaceae
Populus
tremula L.
EN: European aspen
EL: λεύκη η τρέμουσα
ES: álamo temblón
FR: peuplier tremble
IT: pioppo tremolo
PT: choupotremedor
Distribution and Ecology
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus, West
ern and Middle Asia, Siberia, Mongolia, China, North
ern Africa
Mediterranean region: Spain, France (incl. Corse), Italy
(incl. Sardegna), Croatia, BosniaHerzegovina, Montene
gro, Albania, Greece, Turkey, Syria, Lebanon, Algeria
European aspen is a pioneer species that grows in a
wide range of climatic conditions on fertile, preferably
neutral soils, although it can withstand alkaline reac
tion substrates. Its superficial root system allows it to
grow on shallow but moist soils. P. tremula does not
usually occur in very large populations. It may be found
as a pioneer in open, treeless areas or in coniferous,
leafy or mixed forests. In the Mediterranean region it is
associated with soils with a phreatic water supply and
grows on slopes, valley bottoms and the banks of wa
tercourses, where it may mix with other riparian ele
ments. This taxon is useful in hydrological restoration
projects in moist Mediterranean mountain areas.
Diagnostic traits
The diagnostic traits that make it possible to distin
guish this species from Populus alba and their hybrid,
P. x canescens, are shown in an Appendix.
Sex expression
■
dioecy
Flowering
inconspicuous
flowers, clustered in
pendulous catkins
■ from February to
April, before leaf
development
■
Pollination
■
anemophilous
Fruiting
Ripening
■
■
ovate capsule,
granular
■ 34 mm
from April to
June
■ dispersal by wind
105 Populus tremula
Reproductive biology
Variation and Hybridisation
P. tremula has a strong ability to sprout from the root,
so within a single stand some individuals may belong to
the same genotype, as different genetic studies have
confirmed. In addition, it would seem that there is a
tendency towards aggregation in the spatial distribu
tion of clones (Suvanto and LatvaKarjanma, 2005;
LatvaKarjanma, 2006). However, a great deal of ge
netic variation between relatively close populations has
been observed (Sierra de Grado et al., 2003). In order to
increase the genetic variation within batches of repro
ductive material, it is advisable to collect such material
from several populations that are more or less near to
each other; within the populations themselves, mate
rial should be collected from individuals that are dis
tant from each other or have differently shaped leaves,
since this trait seems to be a good criterion for dis
crimination between genotypes (López de Heredia et
al., 2004). If material is collected for vegetative prop
agation, the sex of the ortet must be considered, both
male and female material should be produced and the
new plantation should be managed in such a way as
to promote regeneration through seeds.
P. tremula hybridises naturally with P. alba, producing
P x canescens (Aiton) Sm.; introgression in this case is
unidirectional and the aspen acts as a male parental
(Lexer et al., 2005). P. x canescens grows on river banks
and in ravines, where it shares its habitat with P. alba,
or in somewhat cooler environments at higher alti
tudes. Its morphology varies greatly, depending on the
degree of backcrossing, and individuals exhibit a gra
dation of traits that may be closer to either P. alba or
P. tremula. Classification difficulties may be solved
using molecular markers (Fossati et al., 2004). P. x
canescens is a dioecious species and its seeds have a
very low level of viability.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
from April to June
using longhandled tools
or naturally fallen
material
sequence for
dehiscent fruits
■ purity: 4050 %
The most common pattern of P. tremula is an imbal
ance of the maletofemale ratio within stands (Wor
rell, 1995; Worrell et al., 1999), which can lead to
limited seed production.
Collection must begin when the capsules start to open;
this requires careful field supervision, since seeds are
quickly scattered by the wind. Keeping fruitproducing
catkins at room temperature for 3 to 5 days allows cap
sules to open up fully and permits seed dehiscence. It
is not necessary to remove the tufts of hair that cover
the seeds, although doing so does make handling eas
0.060.16 g
T: 18 ºC
HC: 68 %
■ airtight container
■
ier. Separation can be performed using finemesh sieves
and compressed air. Cleaning and preparation should
be carried out in less than a week, since the seeds lose
their viability rapidly if kept in normal temperature and
moisture conditions. Seeds with a moisture content of
5 % to 8 % may be preserved for a couple of years in
airtight containers at 4 ºC to 5 ºC; for longer periods, it
is advisable to store them at 20 ºC (Simak, 1982).
Seeds that have been preserved for a long time must be
rehydrated gradually (using moist air, for instance), as
they can be damaged by excessively rapid moisture
uptake.
Populus tremula 106
Germination under controlled conditions
■
without treatment
■
Nursery production
Sowing season
■
Conditions
spring, without treatment,
immediately after collecting
20 ºC to 25 ºC
Nursery practice
Type of cutting
■
■
root
softwood
Position along
the shoot
■
9095 %
Emergence
bareroot: 1020 g/m2; cir ■ 1224 h after sowing
cumference up to 68 cm or
total height up to 100150 cm
3
■ forestpot 300 cm : 1/0
■ container 3.5 l: 1/1
■
The seeds should not be covered or pressed into the
substrate after sowing. Seedlings are very delicate and
are susceptible to drought during the first month.
When plants are produced in containers, it is possible
Vegetative propagation
Germination Viability
to sow in small pots (volume up to 7075 cm3). Once
the seedlings have become established in these smaller
cells, they can be removed and transplanted into the
growth containers.
Number of
internodes Size
40 cm
2
European aspen and its hybrids are difficult to repro
duce using cuttings from branches, since the aerial or
gans do not induce root primordia (Blake and Atkinson,
1986; Wyckoff and Zasada, 2003).
The more common method of achieving vegetative
propagation in European aspen is by means of root cut
tings. During the winter, root cuttings are buried hori
zontally in boxes with moist sand. Once the shoots are
about 5 cm in length, they are cut, treated with indol
butyric acid in powder form, planted in a substrate of
peat and vermiculite (1:1) and held in a tunnel in a mist
atmosphere. Once roots are formed, the plants are
gradually hardened off. Later, they are transferred to
cultivation beds or, preferably, to containers (Moon
Collecting time
winter
spring or summer
Auxin
concentration
none
0.5 %
2003; Trees for life, 2004). Haapala et al. (2004) sug
gest propagating from softwood cuttings approxi
mately 23 mm in diameter, obtained from parent
plants rejuvenated by means of in vitro cultivation and
rooted in forest pots under conditions of high environ
mental humidity.
Ahuja (1983) describes a method for fast microprop
agation of Populus tremula. Cultivation is started with
apical or axillary buds in a medium with cytoquinines.
Later, the microstems are rooted in a medium with
auxins or directly in a substrate under controlled cli
mate conditions. This second option reduces the cost
considerably.
107 Populus tremula
Pregermination treatments
References
General references
Amaral Franco J do (1993) Populus L. In: Tutin TG et al. (eds).
Flora Europaea. Vol 1. 2nd edn. Cambridge University Press,
Cambridge
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Nicolás JL, Iglesias S, Alía R (2001) Fichas descriptivas de es
pecies. In: García del Barrio JM et al. (coord). Regiones de
identificación y utilización de material forestal de reproduc
ción. Ministerio de Medio Ambiente, Madrid
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di
alberi e arbusti della flora mediterranea. ANPA, Roma
Soriano C (1993) Populus L. In: Castroviejo S et al. (eds). Flora
Ibérica. Vol 3. CSIC, Madrid
Specific references
Ahuja MR (1983) Somatic cell differentiation and rapid clonal
propagation of aspen. Silvae Genetica 32:131135
Blake TJ, Atkinson SM (1986) The physiological role of absicic
acid in the rooting of poplar and aspen stump sprouts. Physio
logia Plantarum 67:638643
Fossati T, Patrignani G, Zapelli I, Sabatti M, Sala F, Castiglione
S (2004) Development of molecular markers to assess the level
of introgression of Populus tremula into P. alba natural popu
lations. Plant Breeding 123:382385
Haapala T, Pakkanen A, Pulkkinen P (2004) Variation in survi
val and growth of cuttings in two clonal propagation methods
for hybrid aspen (Populus tremula x P. tremuloides). Forest
Ecology and Management 193:345354
LatvaKarjanmaa T (2006) Reproduction and population struc
ture in European aspen. Ph.D Thesis. University of Helsinki,
Finland
Lexer C, Fay MF, Joseph JA, Nica MS, Heinze B (2005) Barrier
to gene flow between two ecologically divergent Populus
species, P. alba (white poplar) and P. tremula (European aspen):
the role of ecology and life history in gene introgression. Mo
lecular Ecology 14:1045–1057
López de Heredia U, Sierra de Grado R, Cristóbal MD, Martínez
Zurimendi P, Pando V, Martín MT (2004) A comparison of
isozyme and morphological markers to assess the within pop
ulation variation in small populations of European aspen (Pop
ulus tremula L.) in Spain. Silvae Genetica 53:227233
Luna T (2003) Propagation protocol for aspen using root cut
tings. Native Plants Journal 4:129131
Sierra de Grado R, Martínez Zurimendi P, López de Heredia La
rrea U (2003) Reproducción sexual y diversidad genética de
Populus tremula. In: Sierra de Grado R (coord). El álamo tem
blón (Populus tremula L.). Bases para su cultivo, gestión y con
servación. Ediciones MundiPrensa, Madrid
Simak M (1982) Germination and storage of Salix caprea L. and
Populus tremula L. seeds. In: Wang BSP, Pitel JA (eds). Proceed
ings of the international symposium on forest tree seed storage.
IUFRO Canadian Forestry Service, Chalk River: 142160
Suvanto LI, LatvaKarjanmaa TB (2005) Clone identification
and clonal structure of the European aspen (Populus tremula).
Molecular Ecology 14:28512860
Trees for life (2004) The propagation of aspen from root cut
tings. (online URL
http://www.treesforlife.org.uk/tfl.aspen_propagation.html)
Worrell R (1995) European aspen (Populus tremula L.): a review
with particular reference to Scotland I. Distribution, ecology
and genetic variation. Forestry 68:93105
Worrell R, Gordon AG, Lee RS, McInroy A (1999) Flowering and
seed production of aspen in Scotland during a heavy seed year.
Forestry 72:2734
Wyckoff GW, Zasada JC (2003) Populus L. In: Bonner FT (ed).
Woody Plant Seed Manual. USDA Forest Service. Reforesta
tion, Nurseries and Genetics Resources, Connecticut. (online
URL http://www.nsl.fs.fed.us/wpsm/Populus.pdf)
Populus tremula 108
Prunus
mahaleb L.
EN: mahaleb cherry, Saint Lucie cherry
EL: αγριοκέρασο αχαλέμπιο; κεράσι
μικρόκαρπο
ES: cerezo de Santa Lucía, cerecino
FR: bois de SainteLucie, cerisier de
SainteLucie
IT: ciliegio di Santa Lucia, ciliegio canino
PT: cerejeiradeSanta Lúcia, cerejeira
mahaleb
Distribution and Ecology
General distribution: Southwestern, Southeastern,
Middle and Eastern Europe, Caucasus, Western and
Middle Asia, Northern Africa
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sicilia), Croatia, BosniaHerzegovina,
Montenegro, Albania, Greece, Turkey, Lebanon, Morocco
Diagnostic traits
Prunus mahaleb is a deciduous shrub that usually
reaches a height of 2.5 m, although it can sometimes
grow higher. Its leaves are broadly ovate or subcordate,
sometimes suborbicular, up to 7 cm long, glabrous
Reproductive biology
Sex expression
■
gynodioecy
Flowering
white flowers,
clustered in short
corymbiform,
racemiform cymes
■ from May to June
■
Pollination
■
The St. Lucie cherry prefers calcareous, alkaline to neu
tral soils and cool, moist environments. It is found as
scattered individuals or in small groups in thorny thick
ets, humid forest glades, next to streams and ravines,
and on cliffs and rocky, shady slopes.
above and glabrous o slightly pubescent beneath. Its
flowers have glabrous ovaries and are arranged in
groups of three to eleven in short corymbiform, racemi
form cymes.
entomophilous
Fruiting
Ripening
■
■
■
glossy black drupe
610 mm
from June to
September
■ dispersal by
frugivorous
vertebrates
109 Prunus mahaleb
Rosaceae
Variation and Hybridisation
Functionally female individuals seem to have greater
advantages than hermaphrodites as regards traits as
sociated with fertility and tend to produce more fruit
and heavier seeds, possibly due to higher levels of
crossfertilisation, especially in good fruitproducing
years (Jordano, 1993). Their progeny also have a higher
degree of genetic variation due to the absence of self
fertilisation and the contribution of a greater number
of parents to pollination, particularly in more or less
isolated individuals and lowdensity populations (Gar
cía et al., 2005). The number of fruits produced varies
depending on genotype and plant size.
Genetic studies show that the seeds of this species are
efficiently spread by frugivores, which helps to preserve
high levels of genetic diversity within populations (Jor
dano and Godoy, 2000). However, it would seem that
the nonrandom behaviour of pollinators and dispersers
might give the gene flow a certain spatialdistribution
pattern, thus producing genetic structuring within a
single population or between geographically close pop
ulations (García et al., 2007). In particular, seedmigra
tion distances vary depending on the frugivorous
species that acts as a carrier (Jordano et al., 2007), but
dispersal within short distances from the mother tree
seems to be the most frequent pattern (Godoy and
Jordano, 2001).
All these factors make it advisable for fruit collection
not to concentrate solely on individuals of greater size
but to cover a high number of individuals, which should
be somewhat distant from each other, and to include a
high proportion of functionally female trees. In addi
tion, in order to ensure that batches will have a rea
sonable level of genetic variation, it is advisable to
collect material from different more or less neighbour
ing populations within a region of provenance.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
■
■
■
■
from July to August
gatherIng by hand or
beating branches
sequence for fleshy
fruits
■ seed weight / kg fruit:
130265 g
■ purity: 100 %
Fruits must be fully mature when collected, but to pre
vent losses by birds, harvesting should not be delayed.
4893 g
Storage
T: 5 ºC to 4 ºC
MC: 48 %
■ airtight container
■
■
Prunus mahaleb 110
Germination under controlled conditions
■
prechilling (816 weeks)
Conditions
■
■
21 / 16 ºC
light
Like those of other Prunus species, the seeds of the St.
Lucie cherry have dormant embryos and thus require
prolonged cold stratification. They may enter a second
period of dormancy (secondary dormancy) if this cold
stage is interrupted by drying at room temperature
(Baskin and Baskin, 1998) or by a rise in temperature.
However, induction of secondary dormancy by raising
the temperature is used in Prunus avium to achieve
better germination percentages. Suszka et al. (1994)
recommend applying a stratification system that in
Nursery production
Sowing season
■
late winter or early spring,
with treatment
Vegetative propagation
Type of cutting
■
semihardwood
Position along
the shoot
terminal or
subterminal
Germination Viability
■
volves successive hot and cold phases, as happens in
the wild. Optimal treatments are lengthy, lasting for
periods of 24 to 28 weeks, and shorter periods of treat
ment are suggested (2 weeks at 25 ºC + 2 weeks at 3
ºC + 2 weeks at 25 ºC + 12 to 16 weeks at 3 ºC), al
though they may not be suitable for all seed lots. See
ley and Damavandy (1985) estimate that cold
stratification at 4 ºC for 100 days is the optimal treat
ment for ending dormancy.
Nursery practice
■
■
forestpot 300 cm3: 1/0
container 3.5 l: 1/1
Number of
internodes Size
12
Although it is hard to propagate the Saint Lucie cherry
from cuttings, some selected genotypes are multiplied
regularly for use as rootstocks for Prunus avium vari
eties. The cuttings should be buried 10 cm deep in a
perlite substrate or a mix of peat and perlite under a
mist system (Bush, 1978; Vlasic, 1972) and hormones,
which are indispensable for root formation, should be
applied (Lipecki and Selwa, 1978). Ford et al. (2002)
4089 %
Emergence
■
late spring, may be completed
in the second spring
Collecting time
JulyAugust
Auxin
concentration
0.5 1 %
recommend treating the parent plants with gibberellin
to obtain rejuvenated material.
In vitro propagation is the method used commercially
for the production of Prunus mahaleb for use as a root
stock. To initiate the culture, shoots obtained at the
start of budding are used as explants (Dradi et al., 1996;
Saponari et al., 1999).
111 Prunus mahaleb
Pregermination treatments
References
General references
Blanca G, Díaz de la Guardia C (1998) Prunus L. In: Muñoz
Garmendia F, Navarro C (eds). Flora Ibérica. Vol 6. CSIC,
Madrid
Catalán G (1991) Semillas de árboles y arbustos forestales.
Ministerio de Agricultura Pesca y Alimentación. ICONA,
Madrid
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns Ge
hölzvermehrung. Parey Buchverlag, Berlin
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Specific references
Baskin CC, Baskin JM (1998) Seeds. Ecology, biogeography,
and evolution of dormancy and germination. Academic Press,
San Diego
Bush R (1978) Summer rooting of stone fruit understock cut
tings. Combined Proceedings of the International Plant Pro
pagators Society 28:6364
Dradi G, Vito G, Standardi A (1996) In vitro mass propagation
of eleven Prunus mahaleb ecotypes. Acta Horticulturae
410:477483
Ford YY, Taylor JM, Blake PS, Marks TR (2002) Gibberellin A3
stimulates adventitious rooting of cuttings from cherry (Pru
nus avium). Plant Growth Regulation 37:127133
García C, Arroyo JM, Godoy JA, Jordano P (2005) Mating pat
terns, pollen dispersal, and the ecological maternal nei
ghbourhood in a Prunus mahaleb L. population. Molecular
Ecology 14:1821–1830
García C, Jordano P, Godoy JA (2007) Contemporary pollen and
seed dispersal in a Prunus mahaleb population: patterns in dis
tance and direction. Molecular Ecology 16:19471955
Godoy JA, Jordano P (2001) Seed dispersal by animals: exact
identification of source trees with endocarp DNA microsatel
lites. Molecular Ecology 10:2275–2283
Jordano P (1993) Pollination biology of Prunus mahaleb L.: de
ferred consequences of gender variation for fecundity and
seed size. Biological Journal of the Linnean Society 50:6584
Jordano P, García C, Godoy JA, GarcíaCastaño, JL (2007) Dif
ferential contibution of frugivores to complex seed dispersal
patterns. Proceedings of the National Academy of Sciences of
the USA 104:32783282
Jordano P, Godoy JA (2000) RAPD variation and population
genetic structure in Prunus mahaleb (Rosaceae), an animal
dispersed tree. Molecular Ecology 9:12931305
Lipecki J, Selwa J (1978) The effect of coumarin and some re
lated compounds on the rooting of softwood cuttings of Pru
nus mahaleb. Acta Horticulturae 80:7981
Saponari M, Bottalico G, Savino V (1999) In vitro propagation
of Prunus mahaleb and its sanitation from prune dwarf virus.
Advances in Horticultural Science 13:5660
Seeley SD, Damavandy H (1985) Response of seed of seven
deciduous fruits to stratification temperatures and implica
tions for modelling. Journal of The American Society for Hor
ticultural Science 110:726729
Suszka B, Muller C, BonnetMasimbert M (1994) Graines des
feuillus forestiers, de la récolte au semis. INRA, Paris
Vlasic A (1972) Mahaleb propagation by cuttings. Jugoslo
vensko Vocarstvo 6:693698
Prunus mahaleb 112
Prunus
spinosa L.
EN: blackthorn, sloe
EL: τσαπουρνιά, προύμνη η
ακανθώδης
ES: endrino, espino negro
FR: prunellier, épine noire
IT: prugnolo, pruno selvatico
PT: abrunheirobravo, ameixeira
Distribution and Ecology
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus, West
ern Asia, Northern Africa
Mediterranean region:
Portugal, Spain (incl. Baleares), France (incl. Corse), Italy
(incl. Sardegna and Sicilia), Croatia, BosniaHerzegovina,
Diagnostic traits
P. spinosa is a shrub, usually 1 to 3 m in height al
though it can sometimes reach 6 m. It is a spiny de
ciduous plant with blackish bark and many twisting
branches. Its leaves are obovate, oblanceolate or al
most elliptic, up to 4 cm long, glabrescent or pubes
Reproductive biology
Sex expression
■
hermaphroditism
Flowering
white flowers,
solitary or in
fascicles of two
or three
■ from (January)
February to May,
before or at the
same time as leaves
■
Pollination
■
■
Montenegro, Albania, Greece, Turkey, Tunisia, Algeria,
Morocco
The blackthorn is found in thorny thickets, glades and
forest fringes, on riversides and bordering paths. It
grows on different types of soil but prefers alkaline to
slightly acid substrates.
cent above and more or less pubescent beneath. Flow
ers are mostly solitary, sometimes arranged in clusters
of two to three, with white petals that are occasionally
veined with red. Pedicels are glabrous or puberule and
are shorter than the ripe fruit.
entomophilous
selfincompatible
Fruiting
Ripening
■
■
dark blue or blackish
violet, subglobose
or ovoid drupe,
pruinose
■ 720 mm
from September
to December
■ dispersal by
frugivorous
vertebrates
113 Prunus spinosa
Rosaceae
Variation and Hybridisation
Prunus spinosa is an allotetraploid species that may
have derived from a cross between Prunus cerasifera
and some other unknown species (ReyndersAloisi and
Grellet 1994). It has a high degree of morphological
and genetic variation (Mohanty et al., 2000). This vari
ation could be due to the fact that P. spinosa does not
seem to exhibit apomixis, and its reproductive system
would therefore only be associated with pollinators
(Guitian et al, 1993). The species also seems to be self
incompatible and thus requires crosspollination in
order to bear fruit (Yeboah Gyan and Woodell, 1987). It
intercrosses with P. insititia, producing hybrids (P ×
fruticans Weihe) that are difficult to recognise.
Studies using molecular techniques have shown that
interpopulation variation is relatively low in the black
thorn when compared to other woody species and that
it does not exhibit any kind of spatial structuring pat
tern, perhaps because of the rapid and easy dispersal of
the species by animals. However, it would seem that
southern populations – possible refuges during glacia
tions – tend to have higher levels of diversity, and in
clude specific genetic variants (Mohanty et al., 2002).
This makes it advisable to be cautious when moving re
productive materials between areas; using local mate
rial is recommended.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from late summer
to autumn
■ gathering by hand
sequence for fleshy
fruits
■ seed weight / kg fruit:
84160 g
■ purity: 100 %
Fruit abortion during the initial stage of development
seems to be frequent in the blackthorn, to a degree that
varies from one plant to another (Guitián et al., 1992).
■
■
prechilling (1224 weeks)
preheating (24 weeks) + prechilling
(4–18 weeks)
Conditions
■
18 ºC to 22 ºC
Prunus spinosa 114
Blackthorn seeds, like those of other Prunus species,
have dormant embryos, and thus require prolonged
cold stratification. They may enter a second period of
dormancy (secondary dormancy) if this cold stage is in
terrupted by drying at room temperature (Baskin and
Baskin, 1998) or by a rise in temperature. However, in
duction of secondary dormancy by raising the temper
ature is used in Prunus avium to achieve better
germination percentages. Suszka et al. (1994) recom
mend applying a stratification system that involves
T: 5 ºC to 4 ºC
MC: 48 %
■ airtight container
■
Collection of material should not concentrate only on
the most productive individuals.
Germination under controlled conditions
Pregermination treatments
89250 g
Germination Viability
■
7090 %
successive hot and cold phases, as happens in the wild.
Optimal treatments are lengthy, lasting for periods of
24 to 28 weeks, and shorter periods of treatment are
suggested (2 weeks at 25 ºC + 2 weeks at 3 ºC + 2
weeks at 25 ºC + 12 to 16 weeks at 3 ºC), although they
may not be suitable for all seed lots. Seeley and Dama
vandy (1985) estimate that cold stratification at 4 ºC
for 100 days is the optimal treatment for ending
dormancy.
Sowing season
■
late winter or early spring,
with treatment
Vegetative propagation
Type of cutting
■
hardwood
Position along
the shoot
basal or middle
Nursery practice
■
■
forestpot 300 cm3: 1/0
container 3.5 l: 1/1
Number of
internodes Size
10 15 cm
The blackthorn is normally propagated from seed al
though occasionally it is multiplied using hardwood
cuttings taken from parent plants grown for that spe
cific purpose. Results obtained using material from wild
individuals display great variability (Ruiz, 1989).
References
General references
Blanca G, Díaz de la Guardia C (1998) Prunus L. In: Muñoz Gar
mendia F, Navarro C (eds). Flora Ibérica. Vol 6. CSIC, Madrid
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea. ANPA, Roma
Webb DA (1968) Prunus L. In: Tutin TG et al. (eds). Flora Eu
ropaea. Vol 2. Cambridge University Press, Cambridge
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Specific references
Baskin CC, Baskin JM (1998) Seeds. Ecology, biogeography, and
evolution of dormancy and germination. Academic Press, San Diego
Battistini A, Paoli G (2002) Large scale micropropagation of
several peach rootstocks. Acta Horticulturae 592:2933
Guitián J, Guitián P, Sánchez JM (1993) Reproductive biology
of two Prunus species (Rosaceae) in the Northwest Iberian
Peninsula. Plant Systematics and Evolution 185:153165
Guitián J, Sánchez JM, Guitián P (1992) Niveles de fructifi
cación en Crataegus monogyna Jacq., Prunus mahaleb L. y
Prunus spinosa L. (Rosaceae). Anales del Jardín Botánico de
Madrid 50:239245
Emergence
■
late spring, may be completed in
the second spring
Collecting time
winter
Auxin
concentration
0.5 1 %
Trials of in vitro propagation of hybrids between this
species and other Prunus, used as rootstock for plum
and peach varieties, have been conducted (Battistini
and Paoli, 2002; Krizan et al., 2007).
Krizan B, Ondrusikova E, Trckova K, Benedikova D (2007) Ef
fects of paclobutrazol and indole3butyric acid on in vitro
rooting and growth of some rootstocks of the genus Prunus L.
Europena Journal of Horticultural Science 72:198201
Mohanty A, Martín JP, Aguinagalde I (2000) Chloroplast DNA
diversity within and among populations of the allotetraploid
Prunus spinosa L. Theoretical and Applied Genetics
100:1304–1310
Mohanty A, Martín JP, Aguinagalde I (2002) Population genetic
analysis of European Prunus spinosa (Rosaceae) using chloro
plast DNA markers. American Journal of Botany 89:12231228
ReyndersAloisi S, Grellet F (1994) Characterisation of the ri
bosomal DNA units in two related Prunus species (P. cerasifera
and P. spinosa). Plant Cell Reports 13:641646
Ruiz J (1989) Cultivo del endrino (Prunus spinosa L.) en Nava
rra. Navarra Agraria 44:58 (online URL
http://www.grn.es/fl/public/a10.htm)
Seeley SD, Damavandy H (1985) Response of seed of seven
deciduous fruits to stratification temperatures and implica
tions for modelling. Journal of The American Society for Hor
ticultural Science 110:726729
Suszka B, Muller C, BonnetMasimbert M (1994) Graines des
feuillus forestiers, de la recolte au semis. INRA, Paris
Yeboah Gyan K, Woodell SRJ (1987) Flowering phenology,
flower colour and mode of reproduction of Prunus spinosa L.
(Blackthorn), Crataegus monogyna Jacq. (Hawthorn), Rosa
canina L. (Dog Rose) and Rubus fruticosus L. (Bramble) in Ox
fordshire, England. Functional Ecology 1:261268
115 Prunus spinosa
Nursery production
Rubus
ulmifolius Schott
Distribution and Ecology
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Western Asia,
Northern Africa, Macaronesia
Mediterranean region: Portugal, Spain (incl. Baleares),
France (incl. Corse), Italy (incl. Sardegna and Sicilia),
Croatia, BosniaHerzegovina, Montenegro, Albania,
Diagnostic traits
Rubus ulmifolius is a spiny, semideciduous bush. It is
one of the few European species of the genus that re
produces sexually, and is diploid. Its diagnostic traits
include dark mauve, pruinose turions; alternate leaves,
with five leaflets, white tomentose beneath (with stel
late hairs); stipules linear, with the upper side of the
Reproductive biology
Sex expression
■
hermaphroditism
Flowering
■
■
pink, sometimes
white flowers,
clustered in cymes
from May to August
Rosaceae
EN: elmleaf blackberry
EL: βάτος
ES: zarzamora
FR: ronce (commune)
IT: rovo
PT: silva
Pollination
■
Greece (incl. Kriti), Turkey, Tunisia, Algeria, Morocco
The elmleaf blackberry grows in glades and forest
fringes, humid thickets, hedges bordering roads and
cultivated land, ravines and riverside areas. It is indif
ferent to the mineral content of the substrate, and
prefers warm temperate climates.
petioles sulcate on the lower half only; smooth, pink or
sometimes white petals. It produces abundant shiny
black drupes. It is, however, a very polymorphic species,
particularly as regards traits such as leaf shape, rami
fication of inflorescences or petal colour.
entomophilous
Fruiting
Ripening
■
■
■
shiny, black
polydrupe
about 10 mm
from August to
November
■ dispersal by
frugivorous
vertebrates
Rubus ulmifolius 116
Variation and Hybridisation
It is sometimes difficult to identify species of the Rubus
genus. Many taxa have been included under Rubus ul
mifolius because of its high level of polymorphism, the
result of its phenotypic plasticity and facility for pro
ducing lowstability hybrids.
Seed propagation
Seed handling and storage
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from late summer
to autumn
■ gathering by hand
sequence for fleshy
fruits
■ seed weight / kg fruit:
255 g
■ purity: 6098 %
Large intra and interindividual differences exist in fruit
characteristics of the species, which may exhibit large
23 g
fruits with few seeds or small fruits with many seeds
(Jordano, 1982).
Germination under controlled conditions
Pregermination treatments
■
■
prechilling (1216 weeks)
preheating (812 weeks) + prechilling
(812 weeks)
Conditions
■
■
30 / 15 ºC; 25 / 10 ºC
light (12 h/day)
The seeds of the elmleaf blackberry germinate with
difficulty, due to their hard coat. Scarification followed
by cold stratification has achieved good results in other
Rubus species. Moore et al. (1994) and Peacock and
Nursery production
Sowing season
■
spring, with treatment
■
Germination Viability
■
65 %
Hummer (1996) used concentrated sulphuric acid.
However, Campbell et al. (1988) obtained the best re
sults by scarifying manually or treating the seeds with
a 15% sodium hypochlorite solution for 18 hours.
Nursery practice
■
T: 4 ºC
MC: 48 %
■ airtight container
■
forestpot 300 cm3: 1/0 or 2/0
container 3.5 l: 1/1
Emergence
■
1 to 3 months
117 Rubus ulmifolius
Tolerance to desiccation: ORTHODOX
Vegetative propagation
Type of cutting
■
■
root
semihardwood
Position along
the shoot
basal or middle
Number of
internodes Size
8 10 cm
12
The most common way of propagating most species of
the genus Rubus is to use root cuttings. Semihardwood
cuttings can be taken from parent plants or from shoots
obtained from root cuttings. If mother plants are used,
the cuttings should come from short shoots as they are
more vigorous and form better roots than those obtained
from very long shoots (Busby and Himelrick, 1999). Cut
tings should be placed under mist in an aerated sub
References
General references
HeslopHarrison Y (1968) Rubus L. In: Tutin TG et al. (eds).
Flora Europaea. Vol 2. Cambridge University Press, Cambridge
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns Ge
hölzvermehrung. Parey Buchverlag, Berlin
MonasterioHuelin E (1998) Rubus L. In: Muñoz Garmendia F,
Navarro C (eds). Flora Ibérica. Vol 6. CSIC, Madrid
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea. ANPA, Roma
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Specific references
Broome OC, Zimmerman RH (1978) In vitro propagation of
blackberry. HortScience 13:151153
Busby AL, Himelrick DG (1999) Propagation of blackberries
(Rubus spp.) by stem cuttings using various IBA formulations.
Proceedings of the Seventh International Rubus Ribes Sympo
sium 505:327332
Campbell PT, Erasmus DJ, van Staden J (1988) Enhancing seed
germination of sand blackberry. HortScience 23:560561
Collecting time
spring
JuneAugust
Auxin
concentration
none
0.5 %
strate. It is also possible, but not very common, to use
hardwood cuttings (Zimmerman et al., 1980).
A variety of trials of in vitro propagation of the genus
Rubus have been carried out. Regeneration is per
formed using meristems (Bromme and Zimmerman,
1978; Ferradini et al,. 1997) or leaf explants (Graham et
al, 1997; Jun et al., 2006).
Ferradini N, Effati M, Standardi A (1997) Propagazione in vitro
di alcuni genotipi di Rubus. Italus Hortus 4:38
Graham J, Iasi L, Millam S (1997) Genotypespecific regene
ration from a number of Rubus cultivars. Plant Cell Tissue and
Organ Culture 48:167173
Jordano P (1982) Migrant birds are the main seed dispersers
of blackberries in southern Spain. Oikos 38:183193
Jun WY, Ming X, Hua JG, Bo SC, Qin ZH, Le HP (2006) In
vitro organogenesis and plant regeneration from leaves of
blackberry (Rubus occidentalis). Journal of Fruit Science
23:468470
Moore JN, Brown GJ, Lundergan C (1974) Effect of duration of
acid scarification on endocarp thickness and seedling emer
gence of blackberries. HortScience 9:204205
Peacock DN, Hummer KE (1996) Pregermination studies with
liquid nitrogen and sulphuric acid on several Rubus species.
HortScience 31:238239
Zimmerman RH, Galletta GJ, Broome OC (1980) Propagation
of thornless blackberries by onenode cuttings. Journal of the
American Society for Horticultural Science 105:405407
Rubus ulmifolius 118
EN: willow, osier
EL: ιτιά
ES: sauce, mimbre
FR: saule, osier
IT: salice
PT: salgueiro, borrazeira
Salix
spp.
Salix alba L.
Salicaeae
The most common species of willow in the Mediter
ranean region are found in riparian areas and wetlands,
on many different types of soil. They can easily with
stand waterlevel fluctuations and usually remain on
river banks permanently, behaving like pioneers due to
their facility for vegetative propagation and their abil
ity to take root again after periodic floods. Among the
species included in this guide, S. fragilis and S. triandra
are present in the Mediterranean region but are more
common in other, cooler areas of their distribution
range. S. atrocinerea and S. eleagnos are also species
that tend to grow in cooler environments than S. pur
Diagnostic traits
Willows are trees or shrubs with alternate, rarely op
posed, deciduous leaves with short petioles and winter
buds coated with a single scale. Willows exhibit a wide
range of morphological variations at intraspecific level,
especially as regards vegetative structures, and deter
purea, S. salviifolia and S. pedicellata. Of these three,
the latter two are the most thermophilous and are only
found in the Mediterranean region. Willows grow on
alkaline to neutral soils but some species have clear
preferences. This is the case of the calcicole S. eleagnos,
or of S. salviifolia, which thrives on acid substrates,
while S. fragilis and S. purpurea prefer neutral soils; S.
atrocinerea avoids salty ground, but S. alba can with
stand a certain level of salinity.
The distribution of the Salix species that occur in the Eu
ropean Mediterranean Region is given in an Appendix.
mination can thus be somewhat complex in young in
dividuals or outside the flowering season. Traits that
make it possible to distinguish the riparian species
found in the Mediterranean Region can be found in an
Appendix.
119 Salix
Distribution and Ecology
Reproductive biology
Sex expression
■
dioecy
Flowering
flowers clustered
in catkins
■ from January to
March, from February
to April, from March
to May (from more to
less thermophile)
■
Pollination
■
entomophilous and
anemophilous
Although willowflowers produce nectar, it has been
shown that they may also be pollinated by the wind.
The effectiveness of anemophile pollination seems to
Variation and Hybridisation
Within the Salix genus, dioecy, intraspecific morpho
logical variation and the relatively frequent occurrence
of natural hybridisation, which may even produce fer
tile descendants, often make it difficult to determine
taxonomical limits and assign individuals to a particu
lar species. Thus, S. amplexicaulis is very close to S. pur
purea, differing only in the morphological traits of its
leaves. S. atrocinerea exhibits a high level of morpho
logical variation and may hybridise with willows of
cooler climates such as S. aurita or S. caprea or, in areas
of contact, with its vicariant S. cinerea L., from which
it differs in the reddish colouring of the hairs on its
leaves. S. purpurea frequently hybridises with S. vimi
nalis, while S. salviifolia might hybridise with S. pedi
cellata in some areas of its distribution range.
Another complex situation is that of S. × rubens, a
species that comprises the hybrids of Salix alba and
Salix fragilis; it seems to have two genetically differ
entiated groups, each similar to one of the parent
species, but this division may or may not match the at
tributions based on morphological traits (Triest et al.,
2000; De Cock et al., 2003). Notwithstanding the pos
sibility of hybridisation, the genetic similarity to one or
the other parent species would be the consequence of
a low level of genomic affinity between these, which
would in turn result in low recombination potential,
thereby preventing introgression (Barcaccia et al.,
2003).
Another example of variation in willows is the recog
nition of subspecies in Salix alba, S. triandra and S.
purpurea.
Fruiting
Ripening
■
■
■
capsule
24 mm
from March to
June, one to two
months after
flowering
■ dispersal by wind
vary a lot between species, possibly in relation to the
morphological characteristics of female aments and
flowers (Karrenberg et al, 2002).
Some of the species have in the past been widely dis
seminated by man, given their use in basketry, and may
have become naturalised in many areas of Europe; this
could be the case of some populations of Salix fragilis
and S. triandra. There are many ornamental varieties
and there are even hybrids resulting from artificial
crossing of different species (Newsholme, 1992). Other
taxa have also been introduced into the Mediterranean
region, either for ornamental purposes (most com
monly, S. babylonica) or for use in basketry (S. viminalis
L., S. eriocephala Michx.). The use of these types of ma
terial in revegetation and restoration projects is not
recommended.
Although willows reproduce easily by means of vege
tative propagation, they are also scattered by the wind
and, secondarily, by water during spring floods, so a
certain degree of intrapopulational variation and gene
flow between populations is to be expected. Thus, fruits
or cuttings collected from populations that are close
to each other need not be identified as different lots.
However, it should be remembered that populations
may be less variable in river basins with alterations in
the natural flowregime, as these promote vegetative
regeneration of populations at times of unseasonal
floods (Basroum, 2002), or increase genetic isolation
by reducing the level of gene flow between populations
(Lascoux et al., 1996).
Salix 120
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
sequence for
dehiscent fruits
■ purity: 6070 %
Collecting must be carried out when the capsules ripen,
taking on a yellowishbrown colour, and the hair tufts
of the seeds begin to emerge. This requires close con
trol in the field, as seeds are soon scattered by the
wind. Once the fruits have been harvested, they are left
to dry at room temperature for one or two days until
they open. Although it is not strictly necessary, the
tufts of hairs can be removed by straining the seeds
through a fine mesh with the aid of compressed air.
0.060.08 g (S. alba),
0.14 g (S. fragilis)
Cleaning and preparation should be carried out in less
than a week, since the seeds soon lose their viability if
kept in normal temperature and moisture conditions.
If they are not to be used immediately, the seeds can be
kept in airtight containers at 4 ºC for a month; storage
for longer periods (3 to 5 years) will require moisture
control and storage in airtight containers at below 0
ºC (Maroder et al., 2000).
Germination under controlled conditions
Pregermination treatments
■
without treatment
Nursery production
Sowing season
■
spring, without treatment,
immediately after collecting
Conditions
■
20 ºC to 25 ºC
Nursery practice
bareroot: circumference
up to 68 cm or total height
up to 100150 cm
3
■ forestpot 300 cm : 1/0
■ container 3.5 l: 1/1
■
The seeds should not be covered or pressed into the
substrate after sowing. Seedlings are very delicate and
are susceptible to drought during the first month.
When plants are produced in containers, it is possible
T: 18 ºC
MC: 68 %
■ airtight container
■
Germination Viability
■
9095 %
Emergence
■
1224 h after sowing
to sow in small pots (volume up to 7075 cm3). Once
the seedlings have become established in these smaller
cells, they can be removed and transplanted into the
growth containers.
121 Salix
from March to June,
variable by species
and by site
■ gathering by hand or
using longhandled tools
Vegetative propagation
Type of cutting
■
hardwood
Position along
the shoot
indifferent
Number of
internodes Size
30 50 cm
Vegetative propagation of willows growing in riparian
areas is generally easy. Chmelar (1974) establishes two
groups of willows according to the manner of rooting:
the most common group that roots extensively along
virtually the entire stem, as in S. alba, S. purpurea or S.
eleagnos, and the group in which roots are formed only
at the base, as in S. atrocinerea (Vieitez and Peña,
1968). In willows it is possible to use material of over
a year in age with a diameter larger than 20 mm. Har
vested material may be stored at 4 ºC for two months
without problems (Chmelar, 1974; Volk et al., 2004). In
References
General references
Blanco P (1993) Salix L. In: Castroviejo S et al. (eds). Flora
ibérica. Vol. 3. CSIC, Madrid
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Christensen K.E (1997) Salix L. In: Strid A, Tan K (eds). Flora
Hellenica. Vol 1. Koeltz Scientific Books, Königstein
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Nicolás JL, Iglesias S, Alía R (2001) Fichas descriptivas de es
pecies. In: García del Barrio JM et al. (coord). Regiones de
identificación y utilización de material forestal de reproduc
ción. Ministerio de Medio Ambiente, Madrid
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea. ANPA, Roma
Rechinger KH, Akeroyd JR (1964) Salix L. In: Tutin TG et al.
(eds). Flora Europaea. Vol 1. 2nd edn. Cambridge University
Press, Cambridge
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Specific references
Barcaccia G, Meneghetti S, Albertini E, Triest L, Lucchin M
(2003) Linkage mapping in tetraploid willows: segregation of
molecular markers and estimation of linkage phases support
Collecting time
December to March
Auxin
concentration
none
species which root easily, it is possible to use minicut
tings (813 mm thick and 8 cm long) and root them di
rectly in forest pots (Dumroese et al., 2003; Mathers,
2003). It is also feasible to use softwood cuttings in
the summer under conditions of high relative humidity
(Newsholme, 1992).
There are many studies of micropropagation of the
genus Salix. Regeneration is possible from meristems
(Chung and Carrasco, 2001) and from axillary buds
(Bergmann et al., 1985).
an allotetraploid structure for Salix alba x Salis fragilis inter
specific hybrids. Heredity 90:169180.
Basroum N (2002) Relative contribution of sexual and asex
ual regeneration strategies in Populus nigra and Salix alba
during the first years of establishment on a braided gravel bed
river. Evolutionary Ecology 15:255279
Bergman L, Von Arnold S, Eriksson T (1985) Effects of N6
benzyladenine on shoots of five willow clones (Salix spp.) cul
tured in vitro. Plant Cell Tissue and Organ Culture 4:135144
Chmelar J (1974) Propagation of willows by cuttings. New Ze
aland Journal of Forestry Science 4:185190
Chung P, Carrasco B (2001) Micropropagation of Salix spp. for
foliate meristems. Report of the 21st session of the Interna
tional Poplar Commission and 40th session of its Executive
Committee, 2428 September 2000, Portland
De Cock K, Lybeer B, Vander Mijnsbrugge K, Zwaenepoel A,
Van Peteghem P, Quataert P, Breyne P, Goetghebeur P, Van
Slycken J (2003) Diversity of the willow complex Salix alba S. x rubens S. fragilis. Silvae Genetica 52:148153
Dumroese RK, Wenny DL, Morrison SJ (2003) Propagation pro
tocol for container willows and poplars using minicuttings.
Native Plants Journal 4:137139
Karrenberg S, Kollmann J, Edwards PJ (2002) Pollen vectors
and inflorescence morphology in four species of Salix. Plant
Systematics and Evolution 235:181188
Lascoux M, Thorsén J, Gullberg U (1996) Population structure
of a riparian willow species, Salix viminalis L. Genetical Re
search, Cambridge 68:4554
Salix 122
Mathers T (2003) Propagation protocol for bareroot willows in
Ontario using hardwood cuttings. Native Plants Journal
5:134136
Newsholme C (1992) Willows. The genus Salix. Timber Press,
Portland
Triest L, De Greef B, De Bondt R, Van Slycken J (2000) RAPD of
controlled crosses and clones from the field suggests that hy
brids are rare in the Salix albaSalix fragilis complex. Hered
ity 84:555563
Vieitez E, Peña J (1968) Seasonal Rhythm of Rooting of Salix
atrocinerea cuttings. Physiologia Plantarum 21:544555
Volk TA, Ballard B, Robison DJ, Abrahamson LP (2004) Effect
of cutting storage conditions during planting operations on
the survival and biomass production of four willow (Salix L.)
clones. New Forests: International Journal on the biology, bio
technology and management of afforestation and reforesta
tion 28:6378
123 Salix
Maroder HL, Prego IA, Facciuto GR, Maldonado SB (2000) Sto
rage behaviour of Salix alba and Salix matsudana seeds. An
nals of Botany 86:10171021
Sambucus
nigra L.
Distribution and Ecology
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus, West
ern Asia, Northern Africa, Macaronesia
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia,
BosniaHerzegovina, Montenegro, Albania, Greece,
Turkey, Tunisia, Algeria, Morocco
Sambucus nigra is found as scattered trees on the edge
of humid forests, among spiny deciduous bushes and,
Diagnostic traits
The black elder is a deciduous shrub or small tree that
can reach a height of 10 m. The pith in stems and
branches is large, white and spongy. Its leaves are op
posite, composed of 3 to 9 elliptic, ovate or ovate
lanceolate leaflets, usually asymmetric at the base,
Reproductive biology
Sex expression
■
hermaphroditism
Flowering
small white or
creamy flowers,
clustered in corymbs
■ from April to July
■
Caprifoliaceae
EN: common elder
EL: κουφοξυλιά
ES: saúco
FR: sureau noir
IT: sambuco
PT: sabugueiro
Pollination
■
in Mediterranean environments, in valleys and thal
wegs outside riparian areas or associated with perma
nent watercourses. It is indifferent to the mineral
nature of the substrate but requires loose, humid soils.
It grows well on eutrophic and disturbed areas and re
sprouts from the stump.
The natural distribution range of the species is difficult
to establish accurately, since the black elder has been
widely cultivated for its fruit.
with serrate margins. It is easily distinguished from
Sambucus racemosa in that the latter has red fruits, its
racemose inflorescences have smaller, greenish
coloured flowers and it requires cooler, more humid
climates.
entomophilous
Fruiting
Ripening
■
■
■
black, globose drupe
68 mm
Sambucus nigra 124
from August to
September
■ dispersal by
frugivorous
vertebrates
Variation and Hybridisation
There is no information on intraspecific variation and
hybridisation for this taxon.
Seed propagation
Seed handling and storage
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
from August to October
gathering by hand or
using longhandled tools
sequence for fleshy
fruits
■ seed weight / kg fruit:
35110 g
■ purity: 9899 %
Common elder usually bears abundant fruit every year;
however, fruit production can be low in very adverse
climates or in individuals that grow in the shade. The
23 g
proportion of fruits with viable seeds varies between
individuals, as parthenocarpy and embryo abortion can
both occur (Bolli, 1994; Atkinson and Atkinson, 2002).
Germination under controlled conditions
Pregermination treatments
preheating (612 weeks)
+ prechilling (12 weeks)
■ prechilling (12 weeks)
+ freezing (1 day)
■
Conditions
■
■
30 / 15 ºC; 20 / 10 ºC
light (14 h/day)
Common elder seeds experience morphophysiological
dormancy; at the time of fruit ripening, the embryo is
not fully developed. It requires a period of hot stratifi
cation to elongate, followed by a period of cold
Nursery production
Sowing season
■
early autumn, without treatment,
or spring, with treatment
Germination Viability
■
4585 %
stratification to elicit germination. The hot stratifica
tion phase may be replaced by the use of gibberellic
acid solution at a concentration of 1,000 mg/l (Hiday
ati et al., 2000).
Nursery practice
forestpot 300 cm3: 1/0
■ container 3.5 l: 1/1
■
T: 4 ºC
MC: 48 %
■ airtight container
■
Emergence
■
the first spring, may be
completed in the second spring
125 Sambucus nigra
Tolerance to desiccation: ORTHODOX
Vegetative propagation
Type of cutting
■
■
semihardwood
hardwood,
heeled
Position along
the shoot
indifferent
basal
Number of
internodes Size
10 cm
15 cm
The common elder propagates very easily from cuttings,
even when adult individuals are used (Good and Bellis,
1978; Legind and Kaak, 2002). Semihardwood cuttings
can be obtained from almost the whole plant, but
should be taken before they become very lignified and
the spongy pith begins to form (Sandrap, 2000). The
leaves on the upper internode should be left on, as this
improves the quality of the plant obtained (Ventrella
et al., 1998). It is recommended that cuttings be kept
References
General references
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Ferguson IK (1976) Sambucus L. In: Tutin TG et al. (eds). Flora
Europaea. Vol 4. Cambridge University Press, Cambridge
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Ruiz Téllez T, Devesa JA (2007) Sambucus L. In: Castroviejo S
(coord). Flora Ibérica. Vol 15. CSIC, Madrid
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Specific references
Atkinson MD, Atkinson E (2002) Sambucus nigra L. Journal of
Ecology 90:895923
Bolli R (1994) Revision of the genus Sambucus. Dissertationes
Botanicae 223:1227
Brassard N, Richer C, Charlebois D (2004) Micropropagation of
elderberry (Sambucus nigra ssp. canadensis). Agriculture and
AgriFood Canada/ Horticulture Research and Development
Collecting time
AprilSeptember
winter
Auxin
concentration
none
none
under mist (Gupta, 1994). In the event of using hard
wood material, the cuttings should be cut with a heel
so that the spongy pith is not exposed at the base
(Legind and Kaak, 2002).
In vitro regeneration of the common elder is possible
and is carried out using internodal segments (Brassard
et al., 2004).
Centre. (online URL http://www.cshs.ca/brassard/200407Mi
cropropagationelderberryv3files/slide0001.htm)
Good JE, Bellis JA, Munro RC (1978) Clonal variation in roo
ting of softwood cuttings of woody perennials occurring na
turally on derelict land. International Plant Propagators
Society Combined proceedings 28:192201
Gupta VN (1994) Effect of growth regulators on rooting in
Sambucus nigra L. semihardwood cuttings under intermit
tent mist. Horticultural Journal 7:145149
Hidayati SN, Baskin JM, Baskin CC (2000) Morphophysiologi
cal dormancy in seeds of two North American and one
Eurasian species of Sambucus (Caprifoliaceae) with underde
veloped spatulate embryos. American Journal of Botany 87:
1669–1678
Legind E, Kaack K (2002) Propagation of elder. Gron Viden,
Havebrug 143
Sandrap A (2000) La culture des sureaux (Elder). Fruit Belge
68:130132
Ventrella MC, Alves LOLR, Garcia VB, Amaral JP, Buim ARG
(1998) Efeito das folhas e do tipo de estaca no desenvolvi
mento do sistema radicular em estacas de sabugueiro (Sam
bucus nigra L.). UNIMAR Ciencias 7:8185
Sambucus nigra 126
EN: saltcedar, tamarisk
EL: αρμυρίκι
ES: taray
FR: tamaris
IT: tamerice
PT: tamargueira
Tamarix
spp.
Tamarix gallica L.
Tamaricaceae
Tamarisks are species that grow in arid and semiarid
climates but require temporary edaphic humidity from
surface or ground water. They are found in riverine
areas with water regimes ranging from permanent wa
tercourses to ephemeral streams, in humid depressions
and in sandy shoreline areas, as scattered individuals or
in continuous formations depending on water avail
ability. Tamarix boveana, T. canariensis and T. dalmat
ica withstand salinity very well and may be found next
to swamps and saltmarshes in inland and coastal
Diagnostic traits
Tamarisks are freelybranching shrubs with small leaves
that look like scales and glands that secrete salt. The
taxonomy of the genus is quite complex, since its
members exhibit very few external traits that are dis
tinctive and easy to see. Diagnostic traits are usually
related to the morphology of the small flowers, espe
cially the androecium and the bracts that subtend the
flowers (see the table of diagnostic traits in the Ap
pendix), so it is difficult to classify individuals as be
longing to a certain species unless they are flowering.
Tamarix inflorescences can form on currentyear
branches (aestival inflorescences) or on the previous
year’s (vernal inflorescences). However, this trait can
not be used for diagnosis since it is heavily influenced
by local and annual weather conditions. Flowers may
be tetramerous or pentamerous, but the number of flo
ral parts is of no use as a diagnostic characteristic on
its own, as it may be inconsistent.
areas. T. canariensis, T. africana, T. tetranda and T. smyr
nensis can withstand the cold relatively well and occur
in more continental environments.
The distribution of the Tamarix species that occur in
the European Mediterranean Region is given in an ap
pendix. Maps of the Eastern Mediterranean tamarisks
should be taken as only approximate because of the
lack of precise information about their distribution.
The difficulty of assigning individuals to certain species
on the basis of morphological traits, as in the case of
Tamarix gallica and T. canariensis, is corroborated by
molecular techniques. Thus Gaskin and Schaal (2003)
were unable to distinguish between these two species
by means of these techniques, perhaps because the
species turned out to be the same taxon or because
they were intercrossing. T. gallica and T. canariensis are
also difficult to distinguish morphologically from T.
africana in its summer forms (Baum, 1978).
Some Asiatic species of the genus are cultivated for or
namental purposes. These include T. ramosissima, which
can be confused with T. smyrnensis (Baum, 1978). They
should not be employed in restoration projects, since
they may hybridise with native species. Indeed, evi
dence has been found of hybridisation between T.
ramosissima and T. canariensis or T. gallica (Gaskin and
Schaal, 2003).
127 Tamarix
Distribution and Ecology
Reproductive biology
Sex expression
■
hermaphroditism
Flowering
Pollination
white or pink
flowers, clustered
in racemes
■ from March to June,
some species
sometimes also
in autumn
■
■
entomophilous
Fruiting
Ripening
■
■
■
capsule
from 2 to 8 mm
from May to
August, some
species
sometimes also
in autumn
■ dispersal by wind
Variation and Hybridisation
There is no information on intraspecific variation and
hybridisation for natural populations of the Tamarix
species in question.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
from May to August,
sometimes in autumn
■ gathering by hand
sequence for
dehiscent fruits
Sexual propagation of tamarisks is not commonly used
in plant production, since vegetative propagation of the
species is easily achieved. Collection should begin when
capsules ripen; the catkins are briefly left to dry, to
allow for full opening of the fruits. The small seeds are
■
without treatment
Conditions
■
■
■
T: 18 ºC
airtight container
difficult to handle and it is therefore not necessary to
separate them from their open capsules. These seeds
rapidly lose their viability at room temperature, but
may be preserved for 1 to 2 years if kept at tempera
tures below 0 ºC.
Germination under controlled conditions
Pregermination treatments
0.031 g (T. gallica)
20 ºC to 25 ºC
light
Germination Viability
■
8090 %
Tamarix 128
Nursery production
Sowing season
■
spring, without treatment,
immediately after collecting
Nursery practice
bareroot: circumference
up to 46 cm or total height
up to 100150 cm
3
■ forestpot 300 cm : 1/0
■ container 3.5 l: 1/1
■
Early seedling growth is very slow. The substrate must
be kept continually moist during seedling development.
Once the seedlings become established they can with
stand severe drought. When plants are produced in
Vegetative propagation
Type of cutting
■
hardwood
Position along
the shoot
indifferent
■
24 h after sowing
containers, it is possible to sow in small pots (volume
up to 7075 cm3). Once the seedlings have become es
tablished in these smaller cells, they can be removed
and transplanted into the growth containers.
Number of
internodes Size
20 30 cm
Emergence
Collecting time
winter
Auxin
concentration
none
Planting minicuttings (8–13 mm diameter, 8 cm long)
directly in forestpot is also possible.
General references
Baum BR (1968) Tamarix L. In: Tutin, TG et al. (eds). Flora Eu
ropaea. Vol 2. Cambridge University Press, Cambridge
Cirujano S (1993) Tamarix L. In: Castroviejo S et al. (eds). Flora
Ibérica. Vol 3. CSIC, Madrid
Nicolás JL, Iglesias S, Alía R (2001) Fichas descriptivas de es
pecies. In: García del Barrio JM et al. (coord). Regiones de
identificación y utilización de material forestal de reproduc
ción. Ministerio de Medio Ambiente, Madrid
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea. ANPA, Roma
Specific references
Baum BR (1978) The genus Tamarix. The Israel Academy of
Sciences and Humanities. Central Press, Jerusalem
Gaskin JF, Schaal BA (2003) Molecular phylogenetic investi
gation of U.S. Invasive Tamarix. Systematic Botany 28:8695
129 Tamarix
References
Ulmus
minor Mill.
Distribution and Ecology
Distribución general: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus, West
ern Asia, Northern Africa
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia,
BosniaHerzegovina, Montenegro, Albania, Greece (incl.
Kriti), Turkey, Cyprus, Libya, Tunisia, Algeria, Morocco
Diagnostic traits
Ulmus minor is a deciduous tree that can reach a great
height (25 to 30 m). Its leaves range from ovallance
olate to suborbicular, acute at apex, very asymmetrical
at base, with the basal lobe shorter than the petiole
and the margin irregularly dentate. The seeds are con
Reproductive biology
Sex expression
■
androdioecy
Flowering
inconspicuous flowers,
clustered in glomerules
■ from February to April,
before leaf development
■
Ulmaceae
EN: common elm, field elm
EL: φτελιά, καραγάτσι
ES: olmo común, álamo negro
FR: orme champêtre, ormeau
IT: olmo campestre, olmo comune
PT: negrilho, ulmeiro
The elm is a typical species of the Mediterranean ri
parian forest, where it grows in stands or as more or
less scattered individuals in areas where the phreatic
water levels are deepest or the water supply is lowest
in the summer, in contact with the zonal vegetation. It
prefers cool, deep, baserich soils. Cultivated elms are
commonly seen next to roads and ditches, or near rural
buildings.
tained in the upper third of the samara. It should not
be confused with the Asiatic Ulmus pumila, which is in
widespread gardening use because of its resistance to
Dutch elm disease. The latter’s leaves are barely asym
metrical and their margins have simple teeth.
Pollination
Fruiting
Ripening
■
■
■
■
anemophilous
selfincompatible
Ulmus minor reproduces naturally through seeds and
by resprouting from the root; genotypes with sterile fe
male flowers tend to produce a large number of empty
■
samara
up to 20 x 17 mm
from March to
April
■ dispersal by wind
fruits due to lack of pollination and to seed abortion
(LópezAlmansa and Gil, 2003).
Ulmus minor 130
Ulmus minor Miller (emend. Richens sensu latissimo) is
a complex with great morphological variation (Richens,
1983) in which some authors have distinguished sev
eral taxa (Melville, 1975). Among these, U. minor var.
vulgaris (Aiton) Richens, accepted by some authors as
Ulmus procera Salisb., can be distinguished from Ulmus
minor var. minor in that its leaves are rough on their
upper side, with wide teeth and a short, hairy petiole,
while the type variety has smooth, lustrous leaves with
a long, glabrescent petiole. Another similar taxon is
Ulmus canescens Melville (=Ulmus minor subsp.
canescens (Melville) Bowicz & Ziel.), which is found in
the central and eastern Mediterranean region. Its
young twigs and the reverse of its leaves are densely
pubescent. There are numerous intermediate forms
among varieties, as well as hybrids with Ulmus glabra
(U. x holandica Mill.) or Ulmus pumila, often including
strong introgression and backcrosses, making identifi
cation difficult.
The species has been widely disseminated in the past
and it is often difficult to determine whether a given
population is indigenous or has become naturalised.
Some authors doubt that the tree is native in certain
northern areas of its current range of distribution
(Richens and Jeffers, 1985). On the other hand, studies
performed using molecular techniques seem to confirm
the massive propagation in Spain and Great Britain of
a sterile clone of the vulgaris variety which grows very
fast but is particularly susceptible to Dutch elm dis
ease. This expansion could be due to the Romans’ use
of trees of this clone for vinetraining (Gil et al., 2004).
There is also evidence that both Ulmus plotii Druce (=
U. minor Mill. var. lockii (Druce) Richens) and Ulmus an
gustifolia (Weston) Weston (= U. minor Mill. subsp. an
gustifolia (Weston) Stace) are varieties that have been
disseminated in Great Britain because of the unique
ness of some of their morphological traits and their
ease of vegetative propagation (Coleman et al., 2000;
Hollingsworth and Armstrong, unpublished data – cited
in Coleman et al., 2000).
In spite of its intensive use by humans, a geographic
pattern of genetic variation has been assessed in the
common elm, according to the results of molecular
studies. This is possibly due to events in natural history,
having to do with different refuges during glaciations,
migration pathways and genetic isolation (Collin et al.,
2002). The existence of geographic differences of ori
gin is also reflected in the dissimilar behaviour of
clones from disparate regions as regards traits of adap
tive significance such as phenology (Ghelardini et al.,
2006). Analyses using molecular techniques have also
been performed in apparently natural Spanish popula
tions, not made up of specimens of the abovemen
tioned nonautochthonous clone of the vulgaris variety.
These studies reveal a certain genetic variation and
thus the existence of sexual propagation mechanisms
in the dynamics of such populations (FuentesUtrilla,
pers. com.).
Dutch elm disease, a malady transmitted by several
scolytids, has wiped out whole populations and nu
merous individuals throughout the range of distribu
tion of the species, leading to the disappearance of
most adult elms. Populations have remained in the
form of young plants that resprout from the roots of
the tree but die once they have reached a certain size.
To this should be added the ageold destruction of elms
in fertile waterside habitats cleared for farming. The
preservation of the species is thus considered a prior
ity in several European countries.
Several hybrids of Ulmus minor with Asiatic and Amer
ican species that are resistant to the disease exist on
the market, as well as some genotypes of the species
that have shown their resistance in several selection
programmes. Genetic transformation has been
achieved in genotypes of the vulgaris variety and their
resistance to Dutch elm disease is being evaluated
(Gartland et al., 2005).
The use of elms in restoration and reforestation proj
ects can be controversial and some precautions must
be taken. The first is to make sure that the material of
choice is free from disease; the second, to ensure that
new plantations do not jeopardise the survival and
characteristics of natural populations that may be lo
cated in neighbouring areas. In accordance with cur
rent knowledge, with a view to complying with a series
of basic principles that are essential for the preserva
tion of the species, it is advisable to bear in mind the
following recommendations:
whenever possible, only use materials from riparian
populations or individuals that exhibit signs of being
indigenous and show no symptoms of Dutch elm dis
ease. Even so, the use of materials that have not been
tested can never guarantee resistance;
avoid using the susceptible clone of the vulgaris vari
ety that has been massively disseminated in the past
in some countries, especially in grapegrowing areas.
Other varieties should be used instead. If genetic
131 Ulmus minor
Variation and Hybridisation
identification is not possible, seed production may be
used as a distinguishing trait to discriminate the un
desirable, diseaseprone genotype, which is sterile;
vegetative rather than seed propagation must be used
when specimens of U. pumila or its hybrids are pres
ent near the collection area, since U. minor can suf
fer introgression by the Asiatic species;
when there is no other option but to harvest from pop
ulations that include individuals with symptoms of
Dutch elm disease, only collect the seeds, as the veg
etative material might be infected;
genetic variation among lots should be encouraged by
collecting materials from few individuals per popu
lation in as many different populations as possible,
especially when phenotypic homogeneity gives rea
son to suspect that individuals belong to the same
genotype;
given the occurrence of varieties whose distribution is
limited to certain areas and the estimated existence
of a certain geographical pattern in the genetic vari
ation of this species, it is advisable not to transfer
materials over long distances;
the use of genetically modified materials should be
considered, with due regard to sterility or invasive
behaviour patterns;
finally, if healthy populations are found near the
restoration area, the risk of introducing new individ
uals which could act as a ‘bridge’ between diseased
and healthy specimens must be carefully assessed.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
from April to May
using longhandled tools
or beating branches
sequence for fruits
that can be sown
■ purity: 8598 %
Elm seeds quickly lose their viability at room tempera
ture and ambient moisture conditions. In small lots, full
Germination under controlled conditions
Pregermination treatments
■
without treatment
Conditions
■
20 ºC
Samara wings can be removed manually for germina
tion in controlled conditions.
68 g (samara)
T: 13 ºC to 0 ºC
MC: 27 %
■ airtight container
■
seeds can be separated from empty seeds by means of
visual inspection.
Germination Viability
■
1050 %
Ulmus minor 132
Nursery production
Sowing season
■
spring, without treatment,
immediately after collecting
Nursery practice
bareroot: 40 g/m2;
circumference up to 68 cm
or total height
up to 100150 cm
3
■ forestpot 300 cm : 1/0
■ container 3.5 l: 1/1
■
When plants are produced in containers, it is possible
to sow in small pots (volume up to 7075 cm3). Once
the seedlings have become established in these smaller
Vegetative propagation
Type of cutting
■
■
root
semihardwood
Position along
the shoot
basal or middle
The common elm is propagated very easily using root
segments and fairly easily when semihardwood cut
tings are used (Kobert, 1979). The root cuttings should
not be more than 15 mm in diameter. The semihard
wood cuttings should be put to root in an environment
with high relative humidity as elm leaves are particu
larly sensitive to drying (Mittempergher et al., 1992).
Hardwood material gathered in winter and put to root
with bed heating does form roots, but has very low sur
vival rates at the hardening off stage (Bartolini et al.,
1997; Griffin and Schroeder, 2004).
References
General references
■
1 to 2 weeks after sowing
cells, they can be removed and transplanted into the
growth containers.
Number of
internodes Size
5 8 cm
10 15 cm
Emergence
Collecting time
early spring
JuneJuly
Auxin
concentration
none
0.5 %
Owing to damage in natural populations caused by
Dutch elm disease, in vitro propagation is a promising
method for regenerating and conserving indigenous
specimens. In recent years, a variety of regeneration
trials have been conducted using cotyledons (Corre
doira et al., 2002), leaves (Conde et al., 2004; Dorion et
al., 2004) and segments of internodes (Diez and Gil,
2004; Dorion et al., 1993).
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Nicolás JL, Iglesias S, Alía R (2001) Fichas descriptivas de es
pecies. In: García del Barrio JM et al. (coord). Regiones de
identificación y utilización de material forestal de reproduc
ción. Ministerio de Medio Ambiente, Madrid
Christensen KI (1997) Ulmus L. In: Strid A, Tan K (eds). Flora
Hellenica. Vol 1. Koeltz Scientific Books, Königstein
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di
alberi e arbusti della flora mediterranea. ANPA, Roma
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Tutin TG (1993) Ulmus L. In: Tutin TG et al. (eds). Flora Euro
paea. Vol 1. 2nd edn. Cambridge University Press, Cambridge
133 Ulmus minor
Navarro C, Castroviejo S (1993) Ulmus. In: Castroviejo S et al.
(eds). Flora Ibérica. Vol 3. CSIC, Madrid
Specific references
Bartolini G, Fagnani A, Mittempergher L, Panicucci M (1997)
Propagazione del l’olmo (Ulmus spp.) per talea legnosa. Monti
e Boschi 48:4851
Coleman M, Hollingsworth ML, Hollingsworth PM (2000)
Application of RAPDs to the critical taxonomy of the English
endemic Ulmus plotii Druce. Botanical Journal of the Linnean
Society 133:241–262
Collin E, Santini A, Hollingsworth P (compilators) (2002)
Final report RES GEN CT9678 (online URL
http://www.cemagref.fr/Informations/Actualites/elm/final
rap.htm)
Gartland KMA, McHugh AT, Crow RM, Garg A, Gartland JS
(2005) Biotechnological Progress in dealing with Dutch Elm
Disease. In Vitro Cellular Development Biology Plant
41:364–367
Ghelardini L, Falusi M, Santini A (2006) Variation in timing of
budburst of Ulmus minor clones from different geographical
origins. Canadian Journal of Forest Research 36:19821991
Gil L, FuentesUtrilla P, Soto A, Cervera MT, Collada C (2004)
English elm is a 2,000yearold Roman clone. Nature 431:1053
Griffin JJ, Schroeder KR (2004) Propagation of Ulmus parvifo
lia `Emerald Prairie’ by Stem Cuttings. Journal of Environ
mental Horticulture 22:5557
Conde P, Loureiro J, Santos C (2004) Somatic embryogenesis
and plant regeneration from leaves of Ulmus minor Mill. Plant
Cell Reports 22:632639
Kobert H (1979) Vegetative Vermehrung von Waldbaumen
durch Triebstecklinge. Berichte der Eidgenossische Anstalt fur
das Forstliche Versuchswesen 201:8
Corredoira E, Vieitez AM, Ballester A (2002) Somatic embryo
genesis in elm. Annals of Botany 89:637644
LópezAlmansa JC, Gil L (2003) Empty samara and partheno
carpy in Ulmus minor s.l. in Spain. Silvae Genetica 52:241243
Diez J, Gil L (2004) Micropropagation of Ulmus minor and U.
minor x U. pumila from 4yearold ramets. Investigacion Agra
ria, Sistemas y Recursos Forestales 13:249254
Dorion N, BenJouira H, Jouanin L (2004) Optimization of elm
regeneration in vitro using leaf explants and evaluation of the
process in the transformation experiments. Investigacion
Agraria, Sistemas y Recursos Forestales 13:237247
Dorion N, Godin B, Bigot C (1993) Physiological state and clo
nal variability effects on low temperature storage of in vitro
shoot cultures of elms (Ulmus spp.). Scientia Horticulturae
56:5159
Melville R (1975) Ulmus. In: Stace CA (ed). Hybridisation and
the flora of the British Isles. Academic Press, London
Mittempergher L, Bartolini G, Ferrini F, Panicucci M (1992) As
pects of elm propagation by soft and hardwood cuttings. Suelo
y Planta 2:129137
Richens RH (1983) Elms. Cambridge University Press,
Cambridge
Richens RH, Jeffers JNR (1985) (publ. 1986) Numerical
taxonomy and ethnobotany of the elms of northern Spain.
Anales del Jardín Botánico de Madrid 42:325341
Ulmus minor 134
EN: laurustinus
EL: άγρια δάφνη
ES: durillo
FR: lauriertin, viornetin
IT: lentaggine
PT: folhado
Distribution and Ecology
General distribution: Southwestern and Southeastern
Europe, Western Asia, Northern Africa, Macaronesia
Mediterranean region: Portugal, Spain (incl. Baleares),
France (incl. Corse), Italy (incl. Sardegna and Sicilia),
Croatia, Albania, Greece, Turkey, Lebanon, Israel, Libya,
Tunisia, Algeria, Morocco
This thermophile shrub is typical of holmoak and
Mediterranean pine forests, but may also grow in
Diagnostic traits
Viburnum tinus is an evergreen shrub that does not ex
ceed 3 to 4 m in height. Its leaf traits make it easy to
distinguish from other species found in Europe and in
cluded in different sections of the genus. The leaves of
V. tinus are entire, shiny green and persistent, while
those of V. lantana and V. opulus, both of which are
deciduous, have dentate margins and are very hairy in
the former and palmatelobed in the latter. Moreover,
Viburnum
tinus L.
maquis on cool and rather moist soils and in more
humid Mediterranean forests. It is indifferent to
bedrock type, although it prefers loose, rich soils.
Laurustinus is not specifically a waterside species but
it is worth considering for this use because of its need
for some humidity as well as its interaction with ani
mals. It could be planted in areas of transition between
riparian and zonal vegetation.
V. opulus produces fruits that are bright red instead of
black when mature.
The differences between V. tinus, V. lantana and V. op
ulus have been also determined in phylogenetic stud
ies using molecular markers (Donoghue et al., 2004;
Winkworth and Donoghue, 2005). V. tinus is the most
thermophile of the three species.
135 Viburnum tinus
Caprifoliaceae
Reproductive biology
Sex expression
■
hermaphroditism
Flowering
■
■
Pollination
white flowers
clustered in cymes
anthesis from
November to June,
immature inflores
cences throughout
the year
■
■
entomophilous
selfcompatible
Variation and Hybridisation
Three subspecies have been described: V. tinus subsp.
rigidum (Vent.) P. Silva and Viburnum tinus subsp. sub
cordatum (Trel.) P. Silva, which are found in the Canary
Fruiting
Ripening
■
■
dark blue or black,
ovoid drupe
■ 58 mm length
from August to
September, may
remain on the
shrub until winter
■ dispersal by
frugivorous
vertebrates
Islands and the Azores, respectively, and Viburnum
tinus L. subsp. tinus L., which occurs in the rest of the
distribution range.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
autumn
gathering by hand
sequence for fleshy
fruits
■ seed weight / kg fruit:
355650
■ purity: 9598 %
V. tinus exhibits annual fruitproduction fluctuations
and bears a more abundant crop every three years
(Herrera, 1998).
Germination under controlled conditions
Pregermination treatments
■
preheating (812 weeks)
+ prechilling (8–12 weeks)
Conditions
■
■
30 / 20 ºC
light
Seeds of Viburnum tinus are very difficult to germinate.
They have a combination of weak morphophysiologi
cal dormancy and slow germination and require cyclic
temperatures, as in the wild (Karlsson et al., 2005). In
conventional hot and coldstratification treatment
procedures, prechilling can be replaced by the applica
3980 g
T: 0 ºC to 4 ºC
MC: 48 %
■ airtight container
■
Removal of pulp is necessary for germination of Vibur
num tinus seeds.
Germination Viability
■
4060 %
tion of alternating temperatures in an incubator, with
a lowtemperature peak of less than 15 ºC (García
Fayos, 2001). A daily temperature regime of 20 ºC / 10
ºC (light/dark) in an incubator for 5060 weeks could be
suitable for dormancy reduction and germination in
seeds without pretreatment (Karlsson et al., 2005).
Viburnum tinus 136
Sowing season
■
early autumn, without treatment,
or spring, with treatment
Vegetative propagation
Type of cutting
■
semihardwood
Position along
the shoot
terminal
Nursery practice
■
■
forestpot 300 cm3: 1/0
container 3.5 l: 1/1
Number of
internodes Size
34
Cuttings of laurustinus can be collected almost all the
year round, except when the plant is flowering (Cervelli,
2005; Lamb and Kelly, 1988). Application of a rejuve
nation treatment to the parent plants speeds up and
homogenises the rooting of cuttings (Pignatti and
Crobeddu, 2005). The rooting process works better
under mist; depending on weather conditions, bed
heating may be recommended (Giroux et al., 1999).
References
General references
Catalán G (1991) Semillas de árboles y arbustos forestales. Mi
nisterio de Agricultura Pesca y Alimentación. ICONA, Madrid
Ferguson IK (1976) Viburnum L. In: Tutin TG et al. (eds). Flora
Europaea. Vol 4. Cambridge University Press, Cambridge
Ruiz Téllez T, Devesa JA (2007) Viburnum L. In: Castroviejo S
(coord). Flora Ibérica. Vol 15. CSIC, Madrid
Specific references
Cervelli C (2005) La specie arbustive della macchia mediterranea.
Un patrimonio da valorizzare. Collana Sicilia Foreste 26:39154
Donoghue MJ, Baldwin BG, Winkworth RC (2004) Viburnum
phylogeny based on chloroplast trnK intron and nuclear ribo
somal ITS DNA sequences. Sysematic Botany 29:188198
GarcíaFayos P (coord) (2001) Bases ecológicas para la reco
lección, almacenamiento y germinación de semillas de espe
cies de uso forestal en la Comunidad Valenciana. Banc de
Llavors Forestals, Generalitat Valenciana, Valencia
Giroux GJ, Maynard BK, Johnson WA (1999) Comparison of
perlite and peat:perlite rooting media for rooting softwood
stem cuttings in a subirrigation system with minimal mist.
Journal of Environmental Horticulture 17:147151
Emergence
■
the first spring, slow and
irregular, and may be completed
in the second spring
Collecting time
May to September
Auxin
concentration
0.5 %
Rooting rates are usually over 80%, although high in
dividual variation should be expected if working with a
large number of clones (Cervelli, 2005; Piccioni et al.,
1996; Pignatti and Crobeddu, 2005).
Micropropagation of V. tinus has been carried out suc
cessfully by Nobre et al., (2000) using internodal seg
ments of young plants.
Herrera CM (1998) Longterm dynamics of Mediterranean fru
givorous birds and fleshy fruit: a 12year study. Ecological
Monographs 68:511538
Karlsson LM, Hidayatu SN, Walck JL, Milberg P (2005)
Complexe combination of seed dormancy and seedling devel
opment determine emergence of Viburnum tinus (Caprifoli
aceae). Annals of Botany 95: 323330
Lamb JG, Kelly JC (1988) Propagating viburnums. Plantsman
10:101103
Nobre J, Santos C, Romano A (2000) Micropropagation of the
Mediterranean species Viburnum tinus. Plant Cell Tissue and
Organ Culture 60:7578
Piccioni E, Longari F, Standardi A, Ciribuco S (1996) Propaga
zione per talea e allevamento in vaso di alcune specie arbus
tive. Informatore Agrario 52:8791
Pignatti G, Crobeddu S (2005) Effects of rejuvenation on cut
ting propagation of Mediterranean shrub species. Foresta
2:290295 (online URL http://www.sisef.it/)
Winkworth RC, Donoghue MJ (2005) Viburnum phylogeny
based on combined molecular data: implications for taxon
omy and biogeography. American Journal of Botany
92:653–666
137 Viburnum tinus
Nursery production
Vitex
agnuscastus L.
Distribution and Ecology
Distribución general: Southwestern, Southeastern and
Eastern Europe, Caucasus, Western and Middle Asia,
Northern Africa
Mediterranean region: Spain (incl. Baleares), France
(incl. Corse), Italy (incl. Sardegna and Sicilia), Croatia,
BosniaHerzegovina, Montenegro, Albania, Greece (incl.
Kriti), Cyprus, Turkey, Syria, Lebanon, Israel, Tunisia, Al
geria, Morocco
Diagnostic traits
The chaste tree is a deciduous shrub or small tree that
does not usually exceed 4 to 5 m in height. It is the
only species of its genus that grows naturally in Eu
Reproductive biology
Sex expression
■
hermaphroditism
Flowering
Vitex agnus-castus 138
bluish, rarely white,
flowers, clustered in
superimposed cymes
forming a long
panicle
■ from June
to September
■
Verbenaceae
EN: vitex, chastetree
EL: λυγαριά, λυγιά
ES: sauzgatillo, agnocasto
FR: gattilier, poivre des moines
IT: agnocasto, lagano, aino
PT: agnocasto, árvore da castidade
Pollination
■
This is a thermophile species that occurs in the form of
isolated specimens in willow and oleander formations
next to rivers and temporary watercourses, occasionally
giving rise to very dense, more or less pure stands. Al
though V. agnuscastus is a typical Mediterranean
species it does not grow in especially dry areas, as it
requires a certain level of environmental humidity. It
prefers well drained soils and cannot withstand high
salinity.
rope. Its leaves are opposite, petiolate and compound
palmate with five to seven linear lanceolate leaflets.
entomophilous
Fruiting
Ripening
■
■
blackish, globose
drupe
■ 34 mm
from October to
December
■ dispersal by
frugivorous
vertebrates
Variation and Hybridisation
There is no information on intraspecific variation and
hybridisation for this taxon.
Seed propagation
Seed handling and storage
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
autumn
gathering by hand
sequence for
indehiscent dry fruits
■ seed weight / kg fruit:
750 g
■ purity: 80 %
713 g
Germination under controlled conditions
Pregermination treatments
■
prechilling (12 weeks)
Nursery production
Sowing season
■
autumn, without treatment or spring,
with treatment
Vegetative propagation
Type of cutting
■
■
hardwood
semihardwood
Position along
the shoot
indifferent
indifferent
Conditions
■
■
Germination Viability
30 / 20 ºC
light
■
Nursery practice
■
Number of
internodes Size
Chastetree is readily reproduced by vegetative propa
gation from cuttings collected in winter as well as in
summer (Dirr and Heuser, 2006; Mac Carthaig and
Spethmann, 2000).
6070 %
Emergence
forestpot 300 cm3:
1/0 or 2/0
15 cm
23
T: 4 ºC
MC: 48 %
■ airtight container
■
■
the first spring
Collecting time
winter
summer
Auxin
concentration
none or < 0.5 %
none or < 0.5 %
Economou et al. (2000) and Varma et al. (1991) have
reproduced this species by in vitro cultivation of axillary
buds.
139 Vitex agnus-castus
Tolerance to desiccation: ORTHODOX
References
General references
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea. ANPA, Roma
Tutin TG (1972) Vitex L. In: Tutin TG et al. (eds). Flora Euro
paea. Vol 3. Cambridge University Press, Cambridge
Young JA, Young CG (1992) Seeds of woody plants in North
America. Dioscorides Press, Portland
Specific references
Dirr MA, Heuser CW (2006) The Reference Manual of Woody
Plant Propagation: From Seed to Tissue Culture. A Practical
Working Guide to the Propagation of over 1100 Species, 2nd.
ed. Varsity Pr Inc, Athens
Economou A, Hatzilazarou S, Karahalios V, Ralli P (2000) Pro
pagation of Vitex agnuscastus by tissue culture. Acta Horti
culturae 541:147151
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns
Gehölzvermehrung. Parey Buchverlag, Berlin
Varma PN, Vikramaditya, Sarka M (1991) A preliminary report
on the in vitro culture of Vitex agnuscastus. Journal of Eco
nomic and Taxonomic Botany 15:687694
Vitex agnus-castus 140
General distribution: Southwestern, Southeastern,
Middle and Eastern Europe, Caucasus, Western and
Middle Asia, Northern Africa
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia,
BosniaHerzegovina, Montenegro, Albania, Greece,
Diagnostic traits
The wild grapevine is a deciduous, woody climber that
can reach 30 m in height. In old stems, the bark peels
off in long shreds. Its leaves exhibit wide variations
within the same individual but are normally palmately
lobed, more deeply so in male plants. It is commonly a
Reproductive biology
■
dioecy
Flowering
greenish flowers,
clustered in panicles
■ from April to June
■
Pollination
■
Turkey, Syria, Lebanon, Israel, Tunisia, Algeria, Morocco
This is a typically Mediterranean species that is found
on cool soils, preferably as a climber on trees and
shrubs in riverside areas and groves. It also grows in
leafy woods and colonises rocky areas and slopes in
cooler climates.
dioecious taxon although hermaphrodite flowers have
been reported in some cases (Failla et al., 1992). This
trait distinguishes this taxon from cultivars, which pro
duce hermaphrodite flowers.
entomophilous
Fruiting
Ripening
■
■
red or black, globose
or ellipsoidal berry
■ 58 mm
from September
to November
■ dispersal by
frugivorous
vertebrates
141 Vitis vinifera
Distribution and Ecology
Sex expression
Vitis vinifera L.
subsp. sylvestris
EN: wild vine
EL: αμπέλι, κλήμα
ES: parra borde, vid salvaje
FR: lambrusque, vigne sauvage
IT: vite selvática
PT: labrusca, videira brava
Vitaceae
Variation and Hybridisation
Accurate determination of the range of distribution of
this species is difficult, since its domestication began
between 2500 and 2000 BC. Archaeobotanical studies
seem to show that domestication of the wild grape
vine took place independently in the eastern and west
ern areas of what is believed to be its natural
distribution range (Núñez and Walker, 1989), which
would imply that it was native to both regions. This hy
pothesis has been confirmed by comparative studies
which have used molecular techniques to analyse ma
terials from different points of the distribution range.
Such studies have revealed genetic differences between
western and eastern populations (Imazio et al., 2003;
ArroyoGarcía et al., 2006), possibly because these pop
ulations originated along different colonisation routes
after the glaciations (Imazio et al., 2003). Furthermore,
molecular studies of local grapevine cultivars from dif
ferent European regions have pointed to a correlation
between genetic distance and geographical distribu
tion, suggesting that grapevine domestication basi
cally took place in situ, mainly using local wild
specimens (Sefc et al., 2003; ArroyoGarcía et al.,
2006). Even if cultivars were introduced, they must
have undergone very strong introgression from local
wild provenances (Sefc et al., 2003). In any event, fos
sil grapevine leaves found in southeast France which
date from about 6900 BC (Roiron et al., 2004) are evi
dence of the spontaneous occurrence of the species in
its western distribution range.
The existence of genetic differences between regions
makes it advisable to exercise caution when collect
ing and using reproductive materials, which should be
of local provenance whenever possible. Such precau
tions are further underscored by the existence of
adaptations to cold and drought (Sefc et al., 2003)
which make it inadvisable to introduce cultivars from
other climates.
The wild grapevine was already a threatened species
by the middle of the 19th century, when pests like phyl
loxera developed. It has also suffered the constant de
struction of its habitat (Arnold et al., 1998; López et al.,
2004). Today it is difficult to find pure populations of
V. vinifera subsp. sylvestris, as they are often found to
be a mixture of wild stock, cultivars, rootstock varieties
of American species and hybrids of all these (Sefc et
al., 2003). The wild grapevine is particularly threat
ened in some natural populations with low levels of ge
netic variation, possibly as a result of habitat
destruction and a reduction in the number of individ
uals (Imazio et al., 2003). In fact, populations of this
species do not generally exceed ten individuals and in
some cases are made up of specimens of only one sex
(Arnold et al., 1998). It might therefore be of interest to
use the species in hydrological restoration projects, in
such a way as to reverse the trend towards reproduc
tive isolation of populations and thereby decrease their
degree of discontinuity.
Seed propagation
Seed handling and storage
Tolerance to desiccation: ORTHODOX
Collecting
Cleaning
1,000 seed weight
Storage
■
■
■
■
■
autumn
gathering by hand or
using longhandled tools
sequence for fleshy fruits
seed weight / kg fruit:
(data not found)
■ purity: (data not found)
■
31 g
T: 4 ºC
MC: 8 %
■ airtight container
■
Vitis vinifera 142
Germination under controlled conditions
Pregermination treatments
■
prechilling (812 weeks)
Conditions
■
■
30 / 20 ºC; 30 / 15 ºC; 25 ºC
light
Germination Viability
■
(data not found)
Coldstratification time can be reduced if seeds are
treated with a 1,000 ppm solution of gibberellic acid
for 24 hours (Ellis et al., 1983).
Sowing season
■
autumn, without treatment, or
spring, with treatment
Vegetative propagation
Type of cutting
■
hardwood
Position along
the shoot
basal or middle
Nursery practice
■
forestpot 300 cm3:
1/0 or 2/0
Number of
internodes Size
20 30 cm
Traditionally, hardwood cuttings are used for vine
multiplication. The material is normally harvested be
tween December and January and stored temporarily.
In the spring, when the temperatures begin to rise
(Muñoz and Villalobos, 1976), the cuttings are planted
after overnight rehydration (Alley and Christensen,
1974; Balo and Balo, 1968). The effects of rooting
hormones vary greatly between individuals and in
some cases can even have negative effects (Alley,
1979). It is also possible to propagate vines in sum
mer, using softwood cuttings from one to two intern
odes in length (Muñoz and Valenzuela, 1978;
Stefanini and Iacono, 1998; Thomas and Schiefelbein,
2004). Mac Cárthaig and Spethmann (2000) recom
mend the use of bud cuttings. The material is col
lected in late November and stored until it is
Emergence
■
(data not found)
Collecting time
DecemberJanuary
Auxin
concentration
< 0.1 %
processed and planted in February. In this technique,
the bestdeveloped lateral buds are chosen and the
upper and lower internodes are cut through transver
sally at a distance of 1.5 to 2 cm from the bud. In the
opposite side of the stem from the bud, a sloping cut
is made lengthwise from top to bottom. It is recom
mended that cuttings be treated with 1% indolbutyric
acid. The cuttings are then planted at a slight angle,
leaving only the bud protruding from the substrate,
in boxes containing a mixture of peat and sand. Vines
can also be propagated by layering.
The methods for in vitro propagation of vines are well
developed. Stem segments with a lateral bud are often
used as explants. (Mhatre et al., 2000; Singh et al.,
2004).
143 Vitis vinifera
Nursery production
References
General references
Ellis RH, Hong TD, Roberts EH (1985) Handbook of Seed Tech
nology for Genebanks Volume II. Compendium of Specific
Germination Information and Test Recommendations Hand
books for Genebanks: No. 3. IPGRI, Rome
Piotto B, Di Noi A (eds) (2001) Propagazione per seme di al
beri e arbusti della flora mediterranea. ANPA, Roma
Webb DA (1968) Vitis L. In: Tutin TG et al. (eds). Flora Euro
paea. Vol 2. Cambridge University Press, Cambridge
SSR analysis. Proceedings of the Eighth International Confer
ence on grape genetics and breeding. Acta Horticulturae
603:4957
López MA, Ocete R, Gallardo A, Cantos A, Troncoso A (2004)
Ecological aspects and conservation of wild grapevine popu
lations in the S.W. of the Iberian Peninsula. Acta Horticulturae
652:8185
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns Ge
hölzvermehrung. Parey Buchverlag, Berlin
Specific references
Mhatre M, Salunkhe CK, Rao PS (2000) Micropropagation of
Vitis vinifera L. towards an improved protocol. Scientia Horti
culturae 84:357363
Alley CJ (1979) Grapevine propagation. XI. Rooting of cuttings:
Effects of indolebutyric acid (IBA) and refrigeration on roo
ting. American Journal of Enology and Viticulture 30:2832
Muñoz IH, Valenzuela JB (1978) Rooting capacity of softwood
cuttings in three grape cultivars: Effect of the position on the
cane and time of collection. Agricultura Técnica 38:1417
Alley CJ, Christensen LP (1974) Rooting of ‘Thompson See
dless’ Cuttings. V. Rooting of Fresh and Stored Cuttings when
Cut November to April. American Journal of Enology and Vi
ticulture 25:168173
Muñoz IH, Villalobos AP (1976) Enraizemiento de estaces de
vid. I. Capacidad natural de dos especies de Vitis, efecto de
ubicacion en el sarmiento y de epoca de recoleccion. Agricul
tura Técnica 36:2530
Arnold C, Gillet F, Gobat JM (1998) Situation de la vigne sau
vage Vitis vinifera ssp. sylvestris en Europe. Vitis 37:159170
Nuñez DR, Walker MJ (1989) A review of palaeobotanical find
ings of early Vitis in the Mediterranean and of the origin of
cultivated grapevines, with special reference to new pointers
to prehistoric exploitation in the Western Mediterranean. Re
view of Paleobotany and Palynology 61:205237
ArroyoGarcía R, RuizGarcía L, Bolling L, Ocete R, López MA,
Arnold C, Ergul A, Söylemezoğlu G, Uzun HI, Cabello F, Ibáñez
J, Aradhya MK, Atanassov A, Atanassov I, Balint S, Cenis JL,
Costantini L, Gorislavets S, Grando MS, Klein BY, McGovern
PE, Merdinoglu D, Pejic I, Pelsy F, Primikirios N, Risovannaya
V, RoubelakisAngelakis KA, Snoussi H, Sotiri P, Tamhankar S,
This P, Troshin L, Malpica JM, Lefort F, MartínezZapater JM
(2006) Multiple origins of cultivated grapevine (Vitis vinifera
L. ssp. sativa) based on chloroplast DNA polymorphisms. Mo
lecular Ecology 15:3707–3714
Balo E, Balo S (1968) Connection between the water level during
soaking and the water level of cuttings and their rooting when
planted in nurseries. Szoeloees Gyuemoelcsterm 4:183188
Failla O, Anzani R, Scienza A (1992) La vite selvatica in Italia:
diffusione, carateristiche e conservazione del germoplasma.
Vignevini 1/2:3746
Imazio S, Grassi F, Scienza A, Sala F, Labra M (2003) Vitis
vinifera ssp. sylvestris: the state of healt of wild Italian and
Spanish populations estimated using nuclear and chloroplast
Roiron P, Ali AA, Guendon JL, Carcaillet C, Terral JF (2004)
Preuve de l’indigénat de Populus alba L. dans le Bassin médi
terranéen occidental. Comptes Rendus Biologies 327:125132
Sefc KM, Steinkellner H, Lefort F, Botta R, Da Câmara Machado
A, Borrego J, Maletić E, Glössl J (2003) Evaluation of the gene
tic contribution of local wild vines to European grapevine cul
tivars. American Journal of Enology and Viticulture 54:1521
Singh SK, Khawale RN, Singh SP (2004) Technique for the
rapid in vitro multiplication of Vitis vinifera L. cultivars. Jour
nal of Horticultural Science and Biotechnology 79:267272
Stefanini M, Iacono F (1998) Nuove tecnologie per la produ
zione di barbatelle. Informatore Agrario 54:5356
Thomas P, Schiefelbein JW (2004) Roles of leaf in regulation
of root and shoot growth from single node softwood cuttings
of grape (Vitis vinifera). Annals of Applied Biology 144:2737
Vitis vinifera 144
3
147
Appendices
Variation
and adaptation
Trees possess hereditary systems that resemble those
of other living organisms. The variation we see both be
tween species (interspecific) and within species (in
traspecific) is attributable to two main causes:
environmental conditions, which have long been
known to man and are the basis for forestry activities,
and the genetic makeup of individuals, which is fre
quently ignored. However, this latter source of variation
is so important that we may say that no two trees are
the same.
Forests develop in highly heterogeneous environments,
in terms of both space and time, and generally exhibit
a wide range of genetic variation, both geographically
(between populations) and locally (within populations).
The transmission of this variation from one generation
to the next and its variability ensure the continuous
evolution of species and populations (Morgenstern,
1996). Geographical origin is one of the most relevant
levels of genetic variation between trees, since it de
Factors determining genetic variation
The evolution of riverside forests does not depend solely
on genetic heritage and on the surrounding environ
ment, but is also influenced by human activity. Ecosys
tem fragmentation is one of the main and most
common manmade changes, altering local conditions
and causing the isolation of tree populations. Reduc
tions in the area of the remaining local populations,
together with their increased spatial isolation and the
decrease in the number of reproductive individuals per
unit of surface, can affect genetic processes such as
the flow of pollen, fruits and seeds, crossfertilisation
and naturalselection efficiency; all of which are fac
tors that determine the degree and pattern of distri
bution of genetic diversity among species (Young,
1995). The viability of these populations can be com
promised as a result of disturbances in their reproduc
tive processes and a loss of adaptability, which is very
often associated with a loss of genetic variation.
Population evolution also depends on the gene flow
taking place through pollen or seed migration, which
attenuates the effect of selection and influences the
effective population size (Ne). When populations are
very small and gene flow is low or totally absent,
chance may prevent individuals from adapting to envi
termines adaptability (Zobel and Talbert, 1984; Müller
Starck, 1991).
The genetic quality of forest reproductive materials is
often not taken into account in ecosystem restoration
activities, including riparian habitat recovery projects;
cheaper seeds and plants are used without due regard
for their origin. These practices are revealed as partic
ularly mistaken when we consider the climate changes
that have been taking place in the Mediterranean re
gion in recent decades. Such changes have led to an
aggravation of environmental conditions due to an in
crease in temperatures, a significant drop in springtime
rainfall levels and an increase of interannual variations
in winter rains. Practice has shown that the use of
plants that are adapted to local conditions is one of
the factors that have a positive influence on the suc
cess of forestation and on the evolution of new popu
lations, which grow and develop dynamically in a
process of interaction with their environment.
ronmental conditions (Wright, 1976; Zobel and Talbert,
1984).
In Figure 1, subpopulations A’ and A” are the result of
a not too severe fragmentation of a former population
A. Here, gene flow has not been prevented and the ge
netic differentiation is lower than in subpopulations B’
and B”, shown in Figure 2, which have resulted from
the fragmentation of a riparian population B. Gene
flow between B’ and B” has been limited by the geo
graphical distance that sets them apart. In this case,
genetic differentiation is a function of the distance be
tween the two subpopulations. The effect of this dis
tance may be attenuated by the watercourse, which
provides a transfer route for fruits and seeds, or even
parts of plants. On the other hand, the small size of
subpopulation B” would have a direct effect on the
number of individuals taking part in the reproductive
process. If gene migration is insufficient, population B’’
will tend to undergo a dramatic decrease in variation as
a result of random loss of genes (genetic drift). This
could gradually become more marked in successive
generations, due to depression through inbreeding.
These two factors, drift and inbreeding, might induce a
lower degree of adaptability in subpopulation B”.
149 Variation and adaptation
Reproductive materials and adaptability
Figure 1 Fragmentation undergone by a primitive population A, divided into two
subpopulations of similar size, with a distance between them that allows gene flow.
Figure 2 Fragmentation undergone by a primitive population B leading to two
populations – one of which is very small – separated by a considerable distance.
Variation and adaptation 150
The spatial structure of these populations increases the
fragility of riverside forests, especially in the case of
dioic species or vegetatively propagated species, where
a group of trees that are close together may all be of a
single genotype.
Genetic quality of reproductive materials
An issue that frequently arises is the minimum number
of trees from which material must be collected to pre
serve adequate genetic variation in new populations.
Eriksson et al. (1995) report that, in a given population,
fifty unrelated individuals are enough to capture the
genetic variants that are most common and presum
ably offer adaptive advantages over other less frequent
variants. Reproductive materials should thus be col
lected from individuals that are separated by a certain
References
Eriksson G, Namkoong G, Roberds J (1995) Dynamic Conser
vation of Forest Tree Gene Resources. Forest Genetic Re
sources, 23:28
Morgenstern EK (1996) Geographic Variation in Forest Trees:
Genetic Basis and Application of Knowledge in Silviculture.
UBCPress, Vancouver
MüllerStarck G (1991) Survey of genetic variation as inferred
from enzyme gene markers. In: MüllerStarck G, Ziehe M (eds).
Genetic Variation in European Populations of Forest Trees. JD
Sauerländer, Frankfurt
distance, which in practice is about fifty to one hun
dred metres, in order to minimise kinship relations.
Collection of materials from many individuals is par
ticularly important in order to ensure a wide genetic
base. In the case of vegetative propagation species, the
use of a limited number of clones with high rooting
rates can be convenient from the practical and cost
point of view but makes the adaptability of the result
ing new populations unpredictable. Identifying popu
lations with a significant number of welldeveloped,
healthy individuals is of the utmost importance, since
the areas in which they are found will be potentially
good sites to collect materials.
Funding of forestation projects on private property
through regional, national or European Community ini
tiatives and the efforts of public institutions with re
gard to certification processes for forest reproductive
materials contribute to more responsible action and to
marketing seeds and plants that meet appropriate
quality standards in compliance with the law. European
Union directive 1999/105/EC highlights the importance
of forest reproductive material quality in relation to the
stability, adaptation, resistance and productivity of
forests. In the short term, adequate implementation of
these standards involves good management of mate
rialproduction areas with a view to covering market
demands through the skilful use of handling and prop
agation techniques.
Wright JW (1976) Introduction to Forest Genetics. Academic
Press, New York
Young A (1995) Forest Fragmentation: Effects on Population
Genetic Processes in Caring for the Forest: Research in a
Changing World. In: Koorpilahti E, Mikkelä H, Salonen T (eds).
Proceedings of IUFRO XX World Congress, vol. 2, Tampere,
Finland
Zobel B, Talbert J (1984) Applied Forest Tree Improvement. J.
Wiley & Sons, Inc., New York
151 Variation and adaptation
Delimitation of regions of provenance is a procedure
that allows reproductive materials to be characterised
with a view to promoting their adaptation to planta
tion areas. This system has been adopted for a great
many forest species. The delimitation is based on envi
ronmental and genetic parameters that make it possi
ble to identify the populations that are best suited to
each different area. This requires knowledge garnered
from trials and studies, which have already been con
ducted for many decades in the case of certain species.
However, the genetic variation and adaptability of most
riverside trees are not known and regions of prove
nance at species level have therefore not been deter
mined. Furthermore, limited production of plants of
these species in local and regional nurseries in the
Mediterranean region, and at a higher price than in
other production areas, is one of the factors that en
courage the use of plants of inappropriate origin in
riverine forestation operations.
Seed
handling
Seed handling and storage
One of the decisive phases for the quality of reproduc
tive materials and, consequently, for the success of the
plant production phase, is the treatment they receive
between collection and storage for future use. Fruits
and seeds are living, perishable matter and should
therefore be handled with care; whatever damage is
caused might be irreversible and in most cases will cer
tainly affect quality. Seed lot quality will depend not
only on factors that have to do with the material it
self, which is difficult to control, but also on aspects
which are more controllable, such as the presence of
microorganisms, insect attacks and mechanical and
physiological damage during handling and storage.
Once they have been harvested, the materials must be
transported as quickly as possible to the processing fa
cility, avoiding direct heat and mechanical damage and
making sure that labelling is adequate at all times in
order to ensure correct lot identification. Seeds and fruits
are transported in paper or cloth bags, or in sacks made
of jute or plastic mesh in the case of large volumes.
Seed handling and storage involve a series of sequen
tial processes (Figure 3), which require the application
of different methods, as well as the control of environ
mental conditions, which vary depending on the type of
fruit and seed.
collecting
fruit processing
packaging
storage
plant production
testing
periodic testing
conservation
seed processing
cleaning and conditioning
preprocessing storage
pregermination treatments
sowing
Figure 3 Sequence of activities from collecting to sowing
Seed handling 152
One of the most important aspects to bear in mind is
the degree of desiccation tolerance of the seeds. One
group of species has seeds that are called recalcitrant
because they lose viability if their moisture content
falls below a relatively high threshold, which varies
by species (Quercus, Aesculus, Castanea, Corylus or a
great variety of tropical taxa). Many other seeds,
known as orthodox, can stand moisture levels of less
than 10% and may be kept in cold storage for many
years. There are also seeds that display intermediate
behaviour and are classified into what is known as the
suborthodox group, which includes Juglans, Caria and
Fagus and some riparian taxa like Populus, Salix and
Ulmus.
Once the material has been taken to the processing fa
cility, operations should begin as soon as possible.
Seeds must be weighed and visually inspected to detect
the presence of fungi and insects so that lot handling
priority may be determined and the necessary meas
ures can be taken.
At this stage, the material should be kept in the shade,
in a cool place or in a coldroom, especially when it
has to wait a few days before cleaning begins. In the
case of recalcitrant seeds, this stage should be as short
Extraction and cleaning
The harvested material never arrives pure from the
field, so it must be cleaned to rid it of debris. Seed ex
traction and cleaning tasks are generally laborious, al
though some species require greater effort than others,
especially when no specific mechanical means are
available for this kind of work.
From the practical point of view, the methods used to
obtain a seed lot of suitable external quality depend to
a great extent on the shape and size of the material.
The processing stages and the methods employed for
each different species must be based on separation ef
ficiency: impurities must be eliminated without loss of
viable seed and cleaning work should be minimised to
reduce damage potential and costs.
Preliminary cleaning
When the material is fruits that must be processed for
seed extraction, preliminary cleaning is usually per
formed by flotation or by manual or mechanical siev
ing, both to eliminate large debris and other impurities
as possible and special care must be taken to avoid
both moisture content loss and the proliferation of
fungi. In general, it is also advisable to shorten the stor
age time of fleshy fruits, in the meantime making sure
that they do not lose moisture while at the same time
avoiding fermentation. Dryfruit species and orthodox
seeds can be dried out gradually; in some cases it is ad
visable to give them a preliminary airing to allow the
seed to ripen (Fraxinus angustifolia, F. excelsior), or to
ensure initial drying in the case of fruits that are to be
opened by heating.
Seeds of trees and shrubs normally used in Mediter
ranean afforestation projects can be grouped into four
large categories for the purposes of establishing com
mon extraction and cleaning protocols (Figure 4):
seeds that need not be extracted from their fruits
(Acer, Fraxinus, Quercus, Ulmus)
seeds in capsules and indehiscent dry fruits (Cistus,
Colutea)
seeds in cones and dehiscent fruits (Pinaceae, Cu
pressus, Tetraclinis, many Fabaceae, Alnus, Atriplex,
Betula, Carpinus, Carya, Casuarina, Eucalyptus, Fagus,
Liquidambar, Platanus, Populus, Tilia)
seeds in fleshy strobili and fruits (Caprifoliaceae,
Rosaceae, Rhamnaceae, Oleaceae, Juniperus, Taxus,
Cornus, Ribes)
and also to separate and collect seeds that may have
fallen free during transport and temporary storage of
the lots, as may be the case with cones and dehiscent
fruits.
153 Seed handling
Temporary storage
Seed handling 154
fruit opening
dry
dewinging
final drying
washing
liquid
separation
(recalcitrant: surface only)
final drying
sifting blowing
sifting blowing
(recalcitrant)
cleaning
drying
liquid
separation
final drying
(recalcitrant: surface only)
sifting blowing
dry
dewinging
drying
wet
dewinging
sifting blowing
cleaning
seed extraction
(cold: orthodox and
recalcitranthot: some orthodox)
(except recalcitrant)
fruit drying
cones and dehiscent
dry fruits
fruits that
can be sown
sifting blowing
cleaning
fruit cracking
fruit drying
indehiscent dry fruits
and seeds in capsules
Figure 4 Suggested sequences for handling seeds (some steps may not be necessary
for each lot; final drying may not be essential for immediate stratification or sowing)
final drying
sifting blowing
drying
washing
liquid separation
cleaning
depulping
maceration
fleshy fruits
In some cases the seeds need not be extracted from the
fruits, which can be sown directly. However, in most
species extraction is necessary, since the fruit could be a
physical impediment to germination or could contain
germination inhibitors; it is also necessary to avoid
rotting in the case of fleshy fruits.
Fruits can be opened manually by splitting the coat, with
the aid of pincers, rubber mallets, lancets or other tools if
necessary. This is an intensely laborious method and is usu
ally only employed with very small lots. Mechanised crush
ing is done with different kinds of threshers and other
mechanical tools, or with kitchen utensils such as grinders
or blenders that have been adapted for the purpose.
Dehiscent fruits and cones can be opened through spon
taneous drying and opening, by spreading the material
out in thin layers under cover or in the sun. Forced
hotair ventilation or a kiln with controlled temperature
and moisture can also be used for some species. Al
Fractioning
though Mediterranean species are well adapted to the
heat, a temperature of 30 to 40ºC should not be ex
ceeded and fluctuations in the moisture content of the
seeds should be avoided. To ensure this, the materials
must be protected from the rain and if they are spread
out to dry the seeds should be collected as the fruits
open. The use of perforated trays, to retain the fruit and
let the seeds gradually drop through, can make subse
quent cleaning easier. In the case of fruits with explosive
dehiscence, like some leguminous taxa, they should be
covered to avoid scattering. It is sometimes necessary to
shake the material after it opens in order to extract seeds
that may still remain within the open fruit; this can be
done manually or by using tumblers of different sizes,
depending on the volume of material being processed.
The pulp of fleshy fruits can be removed by manual rub
bing, using a sieve or a bag, after soaking the material in
water. Sand can be added to increase friction. Mechani
cal instruments such as mixers or shakers are also used.
This stage eliminates fruit remnants, infected, damaged
and unviable seeds and other inert matter. In some species,
wings must also be discarded. This step helps to reduce
the size and heterogeneity of seed lots, optimises sam
pling, increases testing reliability and facilitates sowing.
to use flotation, or to place the material in sieves and
wash it with pressurised water. Facilities that process
many seed lots or large seed lots use specially designed
machinery, including scalpers, blowers, aspirators, de
bearders, indented cylinders and gravity tables.
Cleaning methods are based on external characteristics
and on the physical properties that differentiate seeds
from impurities: size, length, shape, colour, density, tex
ture or rate of fall. There are many cleaning methods,
including visual inspection, screening or sieving, win
nowing, and flotation. Cleaning equipment varies ac
cordingly and may be manual or mechanical, adapted
for small or large volumes and different types of seed.
This equipment can be used alone or in combination to
improve lot quality throughout the cleaning process.
In some species, wings must be removed from the seeds
(Pinus, Abies). In some others this is not strictly necessary
but may be done in order to reduce lot volume, as in the
fruits of Fraxinus, Liquidambar or Acer. There are several
dewinging methods, ranging from the simplest, such as
manual rubbing in cloth sacks, to mechanical systems
using debearders, or rotating containers in the case of
large lots, that can carry out dewinging either dry or wet.
The simplest tools are manual screens or sieves, which
are normally used in a sequence of increasingly large or
small mesh sizes. For some species it is more practical
After wet processes like depulping or flotation frac
tioning the seeds must be dried gently for a short pe
riod of time before handling continues. It is advisable
to clean the sieves and machinery after processing each
seed lot.
155 Seed handling
Fruit opening
Drying
The moisture content of preserved seeds is of great im
portance, since it has a direct influence on longevity. Re
calcitrant seeds must only be dried briefly, spread out in
a cool area, to rid them of excess surface water. Ortho
dox seeds can be air dried in a cool, shaded place if no
appropriate purposemade facility exists. In any event,
direct sunlight, hot air or central heating is not recom
mended. If a drying room is available, it should be kept
at a temperature of 15 to 20 ºC and a relative humidity
Testing
It is advisable to test seed lots to ascertain whether
their moisture contents are suitable for storage and to
determine their external quality in terms of germina
tion capacity or viability, purity and seed size. These
test results will indicate whether the lot should be dis
carded, improved by more cleaning, or may be stored as
it is. The values determined by the tests will also give
an idea of expected performance during the plant pro
Storage
As with drying, storage conditions depend on the des
iccation tolerance of the seed. Recalcitrant seeds are
kept in porous bags such as cloth, jute or plasticmesh
sacks, or in perforated plastic containers that allow for
air circulation. They are usually mixed in with inert ma
terials, like peat, coconut fibre, sawdust or vermiculite,
which are maintained at moisture levels similar to
those of the seeds. In environments that are not specif
ically controlled, it is advisable to moisten the seeds
frequently, although they will rapidly lose their viabil
ity in such conditions. In controlled environments they
must be kept at relative humidity levels of 85 to 90 %,
of 15 to 25 %. In these conditions, the seeds of a great
many species reach a moisture content that is quite suit
able for storage. When working with small seed lots,
small drying rooms can be used; silica gel is placed in
the rooms and gradually replaced as it changes colour.
Seeds should be spread out on trays, in thin layers, to
allow for air circulation. Conventional drying limits for
orthodox seeds range from 5 to 10 % moisture content,
although it may sometimes reach 15 %.
duction stage. If the necessary facility is not available
there are laboratories, both official and private, that
provide this kind of service. When sampling and test
ing lots, it is advisable to follow the standardised pro
tocols that make it possible to compare the results of
different lots. In Europe, the protocols most commonly
used are the International Seed Testing Standards of
the International Seed Testing Association (ISTA).
at a temperature of –3 to 0 ºC for temperate species;
this will preserve seed viability for 1 to 2 years.
When relative humidity in the place of storage is not
controlled, orthodox seeds must be kept in airtight re
ceptacles made of metal, plastic or glass. If appropriate
facilities are not available, it is advisable to keep these
containers in a cool, dry place, protected from sunlight.
If coldrooms are available, the conventional storage
temperature is 4 to 5 ºC, and up to –4 ºC for some
species. For longterm conservation, the seeds of many
species can be kept at –18 ºC.
Seed handling 156
The seeds of many species germinate without difficulty
in favourable temperature and humidity conditions.
Sowing in a nursery during the appropriate season or
subjecting seeds to defined conditions in an incubator
– after soaking in water for 24 to 48 hours if the seeds
are dehydrated – is enough to obtain good perform
ance from lots with a high percentage of viable seeds.
However, many species have dormant seeds that need
to be pretreated before they can germinate. The meth
ods most commonly employed to break dormancy are
as follows:
mechanical scarification: this method is used to allow
water to penetrate seeds with an impermeable coat.
The seed coat is cut or abraded using manual tools
such as files, small pliers, knives or mechanical scar
ifiers. If treatment is manual, the ideal application
site to avoid damaging the embryo is the part of the
seed coat that is located just over the tips of the
cotyledons.
chemical scarification: the seeds are soaked in a 95%
sulphuricacid solution at room temperature (18 to
27 ºC). The treatment period will depend on the de
gree of impermeability of the seeds, which varies ac
cording to the species, the lot, and any treatment
applied when handling the material. If the seeds have
been kept in cold storage, it is advisable to first let
them return to room temperature. After treating the
seeds with acid, they must be washed carefully with
plenty of running water for 5 to 10 minutes. This
treatment is also designed to let water penetrate the
seed coats more easily.
soaking in water: the seeds are placed in a water bath
at an initial temperature of 80 to 100 ºC and left to
cool gradually. The volume of water should be 2 to
10 times greater than that of the seeds. This method
is also used to increase seed coat permeability.
cold stratification and hot stratification: cold stratifi
cation requires keeping the seeds moist, at a tempera
ture of 2 to 5 ºC, in an environment that allows a
certain degree of air circulation. The seeds are hydrated
for 24 to 48 hours and placed between layers of a
moist, inert substrate (peat, sand, vermiculite, etc.).
Containers should be covered to avoid loss of moisture,
but care must be taken to ensure adequate air circula
tion. They are then left at the required temperature for
the necessary length of time. Stratification is also pos
sible without a substrate, by placing the moist seeds in
plastic bags, Petri dishes or other containers which must
be opened periodically to let in some air. However,
stratification in such containers should be carried out
with great care in order to avoid not only seed desic
cation but also the proliferation of fungi, especially in
the case of long treatment periods. This type of cold
stratification often makes it possible to break a dor
mancy which is due to physiological causes. When mor
phological dormancy is present in the seeds, in species
in which the embryo has not developed at the time of
fruit ripening, it may be advisable to apply hot stratifi
cation before cold stratification in order to stimulate
embryo development. Hot stratification is also carried
out in a moist environment, but at a temperature of no
more than 30 to 35 ºC; 15 to 20 ºC is normally enough.
use of hormones and other chemical compounds:
some cases require the use of hormones or com
pounds such as gibberellic acid (GA3) or ethylene, or
other substances that have a positive or stimulating
effect on germination. These products should be em
ployed at appropriate concentrations for periods of
time that are not harmful to the seeds.
157 Seed handling
Pregermination treatments
Nursery
culture
Nursery cultivation regime
Generally, each nursery develops a system of cultivation
based on its own experience, which leads to different,
sometimes very different, cultivation regimes for the
same species depending on the nursery in question. Al
though this is inevitable, since each nursery has its own
particular characteristics, it is also necessary for the
various systems to result in a final product which is rel
atively uniform and in accordance with a specific qual
ity of plant for a particular purpose. To this end, the
nursery needs to know the influence of the different
cultivation variables on the development and final
quality of each species. Therefore, the main variables
in nursery cultivation and the basic relationship of each
with plant development need to be identified.
Each variable of the nursery cultivation system must
be considered individually in relation to the whole
regime. The variables or, rather, the groups of variables
which define the cultivation of plants in containers are:
forest reproductive material (seeds, cuttings, etc.), en
Environmental conditions
The environmental conditions to be managed in nurs
eries are temperature, humidity and light (CO2 can also
be controlled). Substrate temperature plays a role in
the absorption of water and transpiration and the as
similation of essential nutrients. Environmental tem
perature (and its diurnal variation or periodicity) affects
metabolic processes such as photosynthesis and respi
ration and biophysical processes such as transpiration
(Landis et al., 1992). Also of interest are the degree of
light (affects plant growth), its intensity (required to
stimulate photosynthesis and prolong active growth),
its duration (involved in the induction of dormancy),
and its quality (different wave lengths activate differ
ent plant functions) (Landis et al., 1992).
Containers
Nursery culture 158
The receptacle or container is one of the variables with
more obvious effects on the crop. The size of the con
tainer (volume, height, diameter and shape) and its den
sity (number of cells per unit area) are the two basic
factors that control plant growth in nurseries as well as
the possible posttransplant response (Landis et al.,
1990a). Low density containers control the balance be
tween the aerial and radical parts of the plant better, thus
vironmental conditions, containers, substrates, fertil
ization, irrigation and pest and disease control (Bris
sette et al., 1991; Landis et al., 1989, 1990a, 1990b,
1992, 1994, 1998; Peñuelas and Ocaña, 1996).
The cultivation regime for a species, a group of species,
or plants of a particular type consists of a definition of
the groups of characteristics or classes to be satisfied
by these variables, giving a detailed description of each
and of plant development throughout the regime (Lan
dis et al., 1998). These features should be included in a
calendar or overall schedule of the cultivation regime,
which in its simplest version is a chart of the condi
tions to be maintained and operations to be carried out
in the nursery from planting until the eventual release
of the plant (Brissette et al., 1991). The growing period
is divided according to the different stages of plant de
velopment in the nursery, normally restricted to three:
the germination and/or establishment phase, the ac
tive (or rapid) growth phase and the hardening phase.
The climatic conditions of the area in which the nursery
is located will have a determining influence on the type
of cultivation and the planting timetable. The most com
mon facilities for controlling environmental factors are
greenhouses and shade screening. Providing shade re
sults in decreased levels of radiation (preventing possi
ble leaf burning), reduced air and leaf temperatures
(enhancing photosynthesis), lower substrate tempera
tures (reducing irrigation requirements) and altered
shoot/root ratio relationships, which can cause morpho
logical imbalances (Svenson, 2000). Some species can be
grown without shade while others need some protec
tion (especially in the early stages of cultivation) to avoid
possible damage to the photosynthetic system.
preventing the tendency of some species to grow tall and
branchless due to the density of planting. Another im
portant aspect of the container is its influence on the sub
strate properties, especially its waterretention capacity
(Ansorena, 1994). Further considerations are that the con
tainer should avoid root malformations by means of anti
spiralling systems, encourage good aerial root pruning and
have sufficient depth for deeprooting species.
The substrate supplies water, air, nutrients and support
to the plant and determines the practical implementa
tion of other important variables such as irrigation and
fertilization (Landis et al., 1990a). Not without reason
is it said that this is the main factor which determines
the success of cultivation in containers (Ansorena,
1994). In general, the composition of substrates used in
both horticulture and forest cultivation is mainly peat,
to which an aerator component such as perlite or ver
miculite is added (Fonteno, 1993; Burés, 1997). In ad
dition to these components, nurseries often use pine
bark, sand, Litonita, topsoil or mulch, whose properties
and characteristics are well reflected in the literature
(Burés, 1997).
Irrigation
Unlike containers, irrigation is not a fixed variable but
a part of daily management in the nursery. It is one of
the most important and sensitive variables in the whole
process of nursery cultivation, due to the small volume
of the containers, the difficulty of rehydrating com
monly used substrates and the influence of irrigation
on plant nutrition and on the properties of the sub
strate (Landis et al., 1989). There are two main factors
to be considered with regard to this variable: the quan
tity and the quality of the water. Both are influenced
by the growth stage of the crop and must conform to
the requirements of the plant at all times.
Water quality can vary with the source, but irrigation
with water of good quality is a requirement for high
quality plant production (Will and Faust, 1999). The
main water quality parameters are salinity, pH, hard
ness and macroand micronutrient content. All of
these parameters can affect plant growth either di
rectly, by creating toxicity (such as that caused by
Na+, Cl, B and heavy metals) and deficiencies, or in
directly, by altering the availability of other plant nu
Although they will depend on cultivation requirements,
the properties of an ideal substrate may be summarised
as: slightly acidic pH (5.56.5); high cation exchange
capacity; low inherent fertility (which presupposes that
the nursery uses fertilisation); proper balance of pore
sizes (macropores and micropores) and a sterile envi
ronment (freedom from pests and diseases) (Landis et
al., 1990a). Aeration porosity (which depends on the
macropores), is considered the most important prop
erty of a cultivation substrate or growing medium
(Bernier and González, 1995).
trients. The most important factor in irrigation water
is its alkalinity, which affects the pH of the substrate.
Carbonates and bicarbonates cause an increase in the
pH of the substrate over time due to their ability to
neutralise H+ ions.
The quantity of irrigation water depends on two dis
tinct factors: watering frequency and the amount used
at each watering. The two are closely intertwined and
depend on the season (evaporationtranspiration) and
the growing phase (Landis et al., 1989). Crucially for
the growth and development of the plant, repeated ex
posure of the substrate to intensive irrigation followed
by no less intense desiccation significantly affects the
availability of water and oxygen to the roots in the sub
strate (Timmer and Miller, 1991; Miller and Timmer,
1994; Heiskanen, 1993). Irrigation needs may be es
tablished by tactile and visual examination of the sub
strate, TDR or gravimetry. The difficulty of managing
this variable often leads nursery operators to irrigate
too much, which results in a loss of efficiency (Karam
and Nieman, 1994).
159 Nursery culture
Substrate
Fertilization
The addition of nutrients to the plant, or fertilization,
is one of the most important factors in the entire
process of cultivation. This variable, along with irriga
tion, makes it possible to manipulate the amount and
quality of growth and accelerate or delay it, and may
also alter the nutrient composition of the plant tissues,
with effects on the level of reserves, rooting capacity
and resistance to water stress, cold and disease.
Nutrients are added by implementing a programme
that sets out the following basic characteristics (Oliet,
1998): the type and composition of the fertilizer, the
way in which it is applied (irrigation, incorporation into
the substrate, etc.), the relative proportion of nutrients
and the timing of fertilizer application (periodic, con
stant or exponential) (Landis et al., 1989). In practice,
the relative proportion of nutrients should be main
tained by means of appropriate concentrations of the
different minerals in solution in the medium, varying
with the phase of plant growth (Ingestad, 1979; Lan
dis et al., 1989; Van den Driessche, 1991).
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of container grown red pine seedlings. New Forests 5:335348
van den Driessche R (1991) Effects of nutrients on stock per
formance in the forest. In: van den Driessche R (ed). Mineral
nutrition in conifer seedlings. CRC Press
Whitcomb CE (1988) Plant production in containers. Lacebark
Inc, Stillwater
Will E, Faust JE (1999) Irrigation water quality for greenhouse
production. PB 1617 Agricultural Extension Service. Univer
sity of Tennessee, Knoxville (online URL http://www.utexten
sion.utk.edu/publications/pbfiles/pb1617.pdf)
161 Nursery culture
pests and mycorrhizae. Agric. Handbk. 674. U.S.D.A, Forest Service,
Washington (online URL http://www.rngr.net/Publications/ctnm)
Cutting
propagation
Vegetative propagation by cuttings
Vegetative propagation, as a production method for
plants that are intended to be used for restoring natu
ral areas, should be considered with precaution because
of the risk of reducing the genetic diversity of new pop
ulations. Nevertheless, in certain taxa with seeds that
are delicate to manage, such as Salicaceae, or when
the production of viable seeds is low, as frequently hap
pens with Ulmus minor; or simply because for certain
taxa this is the cheapest way of producing plants, veg
etative propagation may be a useful alternative.
Cutting propagation is a method commonly used for
mass producing plants of many riparian species, taking
advantage of their suitability for this type of propaga
tion owing to the fact that they require high moisture
content soils and are adapted to periodic flooding. In
any event, if this method is used, special attention
Types of cutting
In highly simplified terms, three types of aerial cuttings
can be differentiated:
softwood cuttings: in woody plants, cuttings obtained
from branches or flexible apexes that have still not
lignified (normally in the months of May and June, or
July). In general, cuttings of this type root rapidly, but
it is important not to let them dry out.
semihardwood cuttings: partially lignified, rigid cut
tings obtained from the current year’s growth in
hardwood plants during the period of vegetative ac
tivity (in general, from midJuly to the beginning of
the autumn).
hardwood cuttings: lignified cuttings obtained from
the previous year’s growth in hardwood plants dur
ing the rest period (end of autumn, winter or the be
ginning of spring). There are three types of hardwood
cutting: straight cuttings, mallet cuttings and heel
cuttings. The latter two are used for propagating
species that root with difficulty. Mallet cuttings re
should be paid to dioic species and to producing mate
rial of both sexes in order to maintain the balance of
male and female individuals.
The determining factor in propagation by cuttings is
correct adventitious root formation. This is a complex
process in which a number of factors come into play.
The success of the rooting process and the survival of
the new plants depend on the combination of those
factors. The aptitude of the species, the genetic fac
tors of the individual, the physiological conditions of
the parent plant, the type of cutting and its position
on the plant, the time when the material is obtained,
the treatments it undergoes and rooting conditions
are the main factors to take into account (Hartmann
and Kester 1987; Mac Cárthaig and Spethmann
2000).
tain, at their base, a segment of the branch into
which they are inserted, whereas the heel cutting
only retains at its base a small portion of that branch,
in the form of a heel, as its name indicates.
Mention should be made of the ease with which some
species can be multiplied using root segments. This
type of material can be used in species that resprout
from the root naturally, such as Populus tremula or
Ulmus minor. However, the drawback to this propaga
tion approach is that root segments are harder to ob
tain than aerial cuttings, added to which it is not
possible to extract a large quantity of material from
the same individual.
Some species can be propagated using more than one
type of cutting, although this might pose the need for
appropriate facilities, as cuttings taken in spring or
summer are more delicate and require an environment
with controlled humidity and temperature conditions.
Cutting propagation 162
Material gathered from parent plants in a juvenile state
usually forms roots more easily than material taken from
adult plants. In turn, in adult individuals, cuttings taken
from lower branches take root more easily than those
taken from upper branches (Mac Cárthaig and Speth
mann, 2000). This difference in behaviour is due to the
phenomenon known as cyclophysis, which is the loss of
juvenile potential – such as the capacity to form adven
titious roots – which tissues suffer as their cells undergo
a greater number of divisions. The lower branches in a
tree come from tissues with a younger physiological age,
despite the fact that they are chronologically older than
the upper branches. It should be borne in mind, more
Preparation of the cuttings
over, that this variable behaviour according to the posi
tion and hierarchy of the branches at the top persists for
a time in the material gathered from them, a phenome
non known as topophysis.
As a result, in order to ensure successful cutting prop
agation, parent plants in which tissues with an adult
physiological age predominate should undergo rejuve
nating treatments. The most common way to achieve
rejuvenation is by severe pruning, although there are
other alternatives, such as serial cutting propagation
or etiolation, among others (Davis and Hartmann,
1988; Howard et al., 1988).
Cuttings should be taken from healthy, vigorous plants,
as far as possible avoiding or eliminating any buds or
flowers on the stems or shoots that are gathered. The
material that has been collected should be kept per
manently in a cool, moist atmosphere, particularly in
the case of softwood or semihardwood cuttings, which
are highly susceptible to drying out. If the shoots are
not processed immediately, they should be wrapped in
plastic bags and kept at a low temperature (14 ºC).
transmitting diseases. Making a bevel cut at the base
of the cutting is recommended, as it helps to enlarge
the surface area of bare tissue with the potential to
send out roots and makes it easier to insert the cutting
into the substrate. It is advisable to make a straight cut
in the upper part of material removed from the middle
and basal part of shoots, i.e. material that does not
have an apical bud, to ensure that the correct orienta
tion is maintained when the cutting is planted.
Pruning shears or sharp knives should be used to take
the cuttings, to produce clean cuts. The tools should be
sterilised fairly frequently by submerging them in al
cohol or a mixture of bleach and water (1:9) to avoid
In cuttings with leaves, those in the middle or basal
third are eliminated to avoid excess water loss from
transpiration; likewise, half of the remaining leaves are
cut if they are very big.
Treatment with hormones
Hormones are applied for the purpose of promoting or
speeding up the production of roots or improving their
quality. The hormone most often used is indolebutyric
acid, in powder or in solution. When the powder form
is used, the cuttings should be lightly shaken to elimi
nate surplus hormone. When the liquid form is used,
the immersion time for the cuttings will depend on the
concentration of the solution.
The hormone preparation that is used should be free
from residues, so it is advisable to prepare the solutions
just before using them and always dispose of any so
lution that is left over once the cuttings have been
treated.
163 Cutting propagation
Topophysis and cyclophysis
Rooting conditions
The most appropriate conditions for stimulating root
formation vary according to the type of cutting. Semi
hardwood and softwood cuttings are kept at high rel
ative humidity in a tunnel with mist, with bed heating
at a temperature of approximately 20 ºC, using a sub
strate that allows good aeration of the roots, such as a
mixture of peat and perlite in a 1:1 proportion (Hart
mann and Kester, 1987). Hardwood cuttings, in most
cases, are planted directly into the growing substrate in
a container. However, the most difficulttoroot species
also require bed heating.
Acclimatisation
One of the most delicate processes in vegetative prop
agation using cuttings is the acclimatisation stage.
Once the material has taken root, it should gradually be
introduced to more demanding temperature and hu
References
Davis FT, Hartmann HT (1988) The physiological basis of ad
ventitious root formation. Acta Horticulturae 227:113120
Hartmann HT, Kester DE (1987) Propagación de plantas. Com
pañía Editorial Continental, SA de CV, México DF
The substrate should always be sterile, with good aer
ation and low fertility. The cuttings are buried up to a
third or half of their length. They should be watered at
regular intervals, keeping the substrate permanently
moist but avoiding saturation and direct sunlight.
The planting method for root segment cuttings varies
according to the species; some are buried horizontally
in the medium and others are treated as if they were
aerial cuttings.
midity conditions. If the cuttings have been planted in
trays or boxes or in small cell pots, the plants should be
transplanted into containers of a suitable size to
enable them to develop well.
Howard BH, HarrisonMurray RS, Vasek J, Jones OP (1988)
Techniques to enhance rooting potential before cutting co
llection. Acta Horticulturae 227:176186
Mac Cárthaigh D, Spethmann W (eds) (2000) Krüssmanns Ge
hölzvermehrung. Parey Buchverlag, Berlin
Cutting propagation 164
Stool
beds
Planning and management of nursery stoolbeds
Nursery stoolbeds are plantations for producing cut
tings to be used for vegetative propagation of clones.
These cuttings are taken from stool shoots, known as
sets.
some other existing production models are different,
mainly with regard to the type of machinery and equip
ment used for cultivation operations, the principles de
scribed here are applicable to all of them. The present
model may also be applied to the production of mate
rials from other species, such as the genus Tamarix, in
which reproduction through vegetative propagation is
employed.
This section describes the basic principles for setting
up and operating a nursery stoolbed model to produce
reproductive materials from Populus clones. Although
Planning
A stool duration of 2 to 4 years is recommended. Yearly
renewal of all stools is more onerous and offers no added
advantages. On the other hand, stump removal becomes
more difficult and costly with increasing stool age.
Stoolbeds require the same ground conditions as other
forestplant production nurseries.
Stoolbed size will depend on expected levels of pro
duction, which in turn depend on the species or clone
that is used and on the growing conditions that apply.
In order to avoid soil exhaustion and allow for aera
tion, as well as for the physical recovery of soil ele
ments, it is advisable to leave areas fallow for a year.
Following the first cut back, all the viable buds on a
stool sprout the following spring, when a varying num
ber of sets may be taken. At the end of the cycle, those
that meet the cutting criteria – in terms of size (length
and diameter), straightness, adequate lignification, the
presence of welldefined buds and absence of damage
– are selected.
Production must be adequate to provide the required
amounts of cuttings for plants and stool renewal. To
this end, the first stage is to determine the necessary
quantity of stools, planting distance and stool duration
(that is, how many years the stools are to remain in
production). Allowance must be made for a percentage
of failures when planting cuttings for stools; in the
case of poplars, less than 5% of the plants fail to root
if conditions are appropriate.
Stool planting distances vary according to the man
agement methods and machinery. In general, stools are
set out very close, in rows with paths between them
for the passage of machinery and equipment.
An estimate of the annual production in number of
poplar sets and cuttings by age of stools is shown in
Table 1. Table 2 gives an estimate of the surface areas
and number of stools needed for a given output, with
several examples of stool duration.
Firstyear sets are cut level with the ground, or about
1 cm above it, to allow for adequate resprouting the
following year.
Table 1 Number of poplar sets and cuttings produced by age of stool
Number of sets
Number of cuttings
1 year old
1
3
2 years old
2
8
3 years old
3
12
4 years old
3
12
165 Stool beds
Age of the stool
Table 2 Amount of stools and surface area required to produce 1,000 poplar clone cuttings, based on nursery stoolbed rotation interval (plant
ing distance: 2 m x 0.125 m)
Stoolbed
rotation
interval
Annual number Annual number Total cuttings
of stools for
of stools for
to plant
production of
stool
allowing for
cuttings
replacement 5% failure rate
1 year
2 years
3 years
4 years
333
91
43
29
111
8
2
1
467
104
48
32
Cultivation stages
Site preparation
Soil preparation begins with deep groundworking, in
cluding one or two passes with the subsoiler, depend
ing on ground conditions. If two passes are needed, the
second pass should cross the first. This helps to aerate
the soil, making it easier for water to reach the roots
and allowing adequate drainage. Subsoiling is done in
September or October, provided the weather conditions
make it possible.
The second task to be undertaken at the same time of
year is harrowing. This should be done immediately after
subsoiling, using one pass or two crisscrossing passes to
grind down all the plant debris remaining on the land.
The next step is organic composting. The compost is
spread and ploughed in to a depth of 30 to 40 cm,
making sure that the time between spreading the com
post and burying it is as short as possible to avoid dry
ing out and evaporation.
At the end of the winter, when temperature and soil
moisture conditions are appropriate, shallow harrowing
is carried out to break up clods and lumps of soil and
provide a finer finish and a good soil structure.
In April, before setting out the cuttings, chemical com
post is added. The soil is then harrowed immediately
and left ready for planting.
Taking cuttings
Once the sets have been purchased or produced, the
cuttings are made by selecting a bud (which becomes
the terminal one) and cutting right above it; the basal
end is then cut at the predetermined length. Cuttings
should be at least 20 cm in length, with a straight
Final number Surface area Total stool Total surface
of stools
by stool bed surface area (stool
in stoolbed
age(m2)
area (m2) bed + fallow)
(m2)
444
116.75
135
12.00
198
120
26.00
8.00
116.75
52.00
36.00
32.00
233.50
78.00
48.00
40.00
upper cut and a chamfered lower cut. In poplars, such
a length ensures that each cutting has 3 or 4 viable,
well developed buds. The upper cut is made 5 to 10 mm
from the new terminal bud in order to make sure that
it does not get buried when planted, since that would
hinder sprouting and future stem growth, but in such
a way that the bud is not affected by an excessively
short cut. The cuttings should be about 10 to 20 mm in
diameter and the material should be checked for proper
lignification to ensure adequate reserves for subse
quent plant growth.
Rejects are discarded and the selected cuttings are tied
into equal bundles. They are then put into cold storage at
a temperature of 2 to 4ºC, making sure that the air circu
lation is good and the minimum relative humidity is 85 %.
The cuttings will be kept in storage until the time comes
to set them out, when soil conditions are optimum.
Cuttings may be colourcoded to identify different
species and clones.
Planting cuttings
The cuttings are taken out of cold storage and soaked
in clean water for 24 to 48 hours to ensure rehydration.
They are then taken out of the water and planted.
When planting, special care should be taken to leave at
least one bud exposed. Mechanised planting proce
dures often bury the cuttings completely, so only one
shoot is obtained for each cutting planted. The problem
of leaving only one bud exposed is that it may get dam
aged by frost. When several viable buds are left on the
cutting, it is advisable to select the strongest shoot and
remove the rest so that the sets obtained are of uni
form size and suitable for cutting production.
Stool beds 166
Cultural treatments
Control of competing vegetation
Preemergence herbicides should be applied both at
the time of planting cuttings and after the first cut
back. After setting up the nursery stoolbed, weeding is
not necessary when fallen leaves remain on the ground
for long periods of time, as they prevent competing
vegetation from growing.
Stoolbed weeding also depends on the weather, which
can cause different degrees of weed proliferation. If
herbicides have been applied, paths are normally ma
chineweeded once or twice during the vegetative pe
riod and planted areas are weeded once by hand.
Harrowing
During the first year, paths are harrowed to facilitate
aeration and irrigation or rainwater penetration. Sub
sequent harrowing operations will be performed
throughout the vegetative cycle, depending on the
amount of grass on the paths and how much the soil
has been compressed by watering.
In established stoolbeds, the initial tillage should go
deeper to break down the soil that has hardened since
the previous year. Definitive groundworking is performed
at this point, since machines will no longer be able to ac
cess pathways when the plants have grown in size.
Ridging
When floodirrigation is to be used the land is divided
Activities
up by making ridges, to make it easier to control the
watering operations. If spray or drip irrigation systems
are used, no ridging is necessary.
Chemical composting
After soil preparation, chemical composting varies de
pending on soil characteristics. It is important to re
member that excessive fertiliser use will produce sets
that are too thick and thus unsuitable for taking cut
tings. It will also cause more sprouting and reduce the
number of dormant buds.
Watering
Watering periods usually last for 5 to 7 months in
Mediterranean climates, but will always depend on the
weather and on soil characteristics.
Although water needs vary, initial watering usually
takes place every 15 days. The interval can then grad
ually be increased, since evapotranspiration decreases
as the stoolstems close the gaps between stools and
the pathways.
Treatment of pests and diseases
Stools should be monitored at all times to avoid the
appearance and proliferation of pests and diseases. All
preventive and curative treatment procedures must be
applied.
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Soil preparation Subsoiling and harrowing
Organic composting and ploughing
Harrowing
Chemical composting and harrowing
Taking material
Cutting sets and cuttings
Treatments
Applying herbicide
Harrowing
Ridging
Weeding
Watering
Plant health treatments
Planting
167 Stool beds
Figure 5 Sequence and duration of activities for the management of a poplar nursery stoolbed (operations can be brought forward or moved back
in time depending on the vegetative period, which varies from year to year and from one place to another)
Harvesting
Once the vegetative cycle is over, once the stools have
shed their leaves, the amount of available sets is
counted. Sets that are good for highquality cuttings are
selected visually on the basis of size, straightness, ade
quate lignification and bud viability. After counting the
sets, the number of cuttings produced can be estimated.
When the material is to be used at the nursery itself,
the sets are cut and the cuttings are prepared as de
scribed in the section on nursery stoolbeds. When the
material is intended for production elsewhere, it can
be stored in the form of sets or cuttings. Sets are cut,
tied in equal bundles and stored in shaded, soilcov
ered ditches until ready for dispatch. In the case of
poplars, the materials produced must comply with Eu
ropean standards (Tables 3 and 4).
Sets that are not selected for cuttings are taken from
their stools and discarded. The stools are then ready to
sprout new shoots in the next vegetative period. Clear
ing all debris from the bed concludes the work until the
next cycle.
Table 3 External quality standards for cuttings and sets of Populus spp. as required by Directive 1999/105/EC
Type of material
Cuttings
Maximum age of the wood
Sets
2
Minimum number of well formed buds
3
2
Free from necroses or damage by harmful organisms
Free from signs of desiccation, overheating, mould or decay
Free from injuries other than pruning cuts
5
√
√
√
√
√
Only a single stem
√
No excessive stem curvature
√
Table 4 Dimensions for cuttings and sets of Populus spp. as required by Directive 1999/105/EC
Type of material Class
Cuttings
Sets
Minimum length
top (cuttings) / midlength (sets) (mm)
0.20
10
3.00
15
EC1
0.20
Non Mediterranean N1
1.50
Mediterranean
3.00
EC2
regions
N2
regions
S2
S1
Minimum diameter at
(m)
4.00
8
6
25
30
Stool beds 168
Master
certificate
European regulations governing the marketing of forest reproductive material
Reproductive materials used in forestation projects
must be adapted to the conditions of the area into
which they are to be introduced. If production is envis
aged, the materials must have undergone selection, and
in some cases improvement, for traits of interest. In
connection with this, the European Union has issued a
set of standards for the marketing of forest reproductive
materials belonging to the species most commonly em
ployed for reforestation in Europe. These rules are de
scribed in Directive 1999/105/EC and related decisions
and compliance by member states is compulsory.
These standards are designed to promote transparency
in the market for reproductive forestry materials by
guaranteeing their quality, in terms of physical condi
tions, whatever genetic selection and improvement the
plants may have undergone and, in some instances,
their geographical origins. To this end, the directive in
troduces a system for the approval of basic materials
from which seeds or parts of plants may be collected
for subsequent plant production, and certificates and
control mechanisms to ensure traceability throughout
the collection, production and marketing process up to
delivery to the final user.
The sphere of application of these rules comprises re
productive materials, intended for forestry purposes, of
a series of species. They include typical riverine vege
tation taxa such as Alnus glutinosa, Alnus incana, Frax
inus angustifolia, Populus spp., Tilia cordata and Tilia
platyphyllos and species that could be used in foresta
tions in this type of habitat, such as some Quercus,
Juglans spp or Robina pseudoacacia, among others.
Each member state may also add further taxons for in
ternal marketing purposes. For instance, Spain has in
cluded other species, some of which are riparian (Ulmus
glabra, Ulmus minor or Tamarix gallica).
When the material is to be used in riparian restoration
projects and is of no interest for propagation, the
seeds are collected from basic materials which may
be taken from seed sources or, possibly, stands; that is
to say, sourceidentified or selected seeds respectively
(Table 5). If the material follows vegetative propaga
tion patterns, the Council Directive limits its produc
tion to the selected (masspropagated from plants
obtained from seeds), qualified or tested categories
(Table 6). In the case of certain species of no market
ing interest that traditionally propagate through parts
of plants, such as some native poplars, it is impossi
ble to comply with the requirements established for
these categories, which are aimed at genetically im
proved material. This difficulty may be solved by con
sidering the sustainable use of such types of
nonimproved materials as adapt naturally to local or
regional conditions. In accordance with article 4.4 of
Directive 1999/105/EC, such sustainable use could be
based on promoting in situ conservation by avoiding
nonautochthonous materials that might introgress
into local populations.
For nonregulated species, it would also be advisable to
apply some of the criteria envisaged under these rules,
especially as regards source control and traceability
though to planting out.
In any event, together with these rules and regula
tions governing basic protocols it would be highly ad
visable to consider a set of best practices for the
production of materials, such as collection from pop
ulations of a certain size and from different individ
uals more or less distant from each other, or the
promotion of clonal mixtures in the case of vegetative
propagation, as a means to guarantee a certain de
gree of genetic variation.
Table 5 Category under which reproductive material from the different types of basic material may be marketed
Category
of forest
reproductive
material
■
■
■
■
sourceidentified
selected
qualified
tested
Seed
source
√
Stand
Seed
orchard
Parents of
family
Clone
Clonal
mixture
√
√
√
√
√
√
√
√
√
√
√
169 Master certificate
Type of basic material
Table 6 Category under which different types of reproductive material may be marketed
Type of reproductive material
Listed species
(except artificial hybrids and GMO)
Category
■
■
■
Artificial hybrids
■
■
■
Genetically modified organisms
■
■
sourceidentified
Fruits
and seeds
Parts of
plants
Plants
selected
√
qualified
√
√*
√
√
tested
√
√
√
selected
√
√
√
qualified
√
√
√
tested
√
√
√
tested
√
√
√
√
√
√
* mass propagated from seeds
Master certificate 170
Plant
passport
European Union regulations regarding phytosanitary cer
tificates and plant passports include a series of direc
tives that aim to avoid the introduction into member
states and the propagation inside the Community of or
ganisms that are harmful to plants or plant products.
The basic plant health passport rules are described in
two directives:
Council Directive 2000/29/EC of 8 May 2000 on pro
tective measures against the introduction into the
Community of organisms harmful to plants or plant
products and against their spread within the Com
munity;
Commission Directive 92/90/EEC of 3 November 1992
establishing obligations to which producers and im
porters of plants, plant products or other objects are
subject and establishing details for their registration.
These regulations are based on the compilation of an
inventory of harmful organisms that are considered es
pecially hazardous and whose introduction into the
Community must be forbidden. The inventory also in
cludes harmful organisms whose introduction through
the medium of certain plants must also be forbidden.
Since the detection of some of these organisms is not
simple, the introduction into the European Union of
plants or plant products from certain countries is for
bidden in some cases, or certified proof of the applica
tion of special controls in the producing country is
demanded.
These controls are not only applicable to plants and
plant products from outside the Community: internal
production is also subject to inspection. In this case,
the plant passport is the document that certifies that
the plants have undergone the required official checks
and are free from the harmful organisms described in
the above regulations. The passport does not guarantee
that they are free from diseases and pests, only that
none of the diseases and pests listed in the regulations
has been found in the plant health inspection of the
plant materials. Plant passports, whose contents are
standardised, are issued by the responsible official
bodies of member states.
With a view to protecting crops or natural populations
that are especially sensitive or whose production or
ecological value is considered a priority, the European
regulations state that special protection must be given
to certain geographical areas. A protected zone may be
a region, a country or a group of countries of the Eu
ropean Union in which one or several harmful
organisms which are found in other EU countries are
not endemic or do not usually occur. The checks appli
cable to plant materials intended for a given protected
zone are specific to that destination and are listed
under the requirements for a special plant passport
known as the ‘ZP passport’. Inspections that do not refer
to these specific requirements are not valid for the rel
evant protected area. This passport must bear the let
ters ‘ZP’ followed by the code of the country or region
of destination of the plant materials to prove that plant
health inspection has been carried out for that
destination.
Plant passport regulations are very detailed and subject
to constant updating. Frequent reference to legislation
databases is therefore advisable to check for possible
changes. The EuroLex web page
(http://europa.eu.int/eurlex/lex/RECH_menu.do) pro
vides all recent updates.
As regards the production of plants for riverine
restoration projects, the species in this handbook that
would be affected by these regulations are: Arbutus
unedo, Humulus lupulus, Laurus nobilis, Platanus ori
entalis, Populus spp., Prunus mahaleb, Prunus spinosa,
Rubus ulmifolius, Viburnum tinus and Vitis vinifera.
Plants and other plant products of the above species
must be accompanied by a plant passport when circu
lating within the Community. For plants and part of
plants of Populus spp. to enter into or circulate within
protected areas, the ‘ZP’ passport is required.
Regarding the entry of products from outside the Com
munity, the regulations should be consulted in each in
dividual case to check for any restrictions. Depending
on the country of origin, the introduction of certain
products into the European Union may be forbidden.
171 Plant passport
European plant health regulations
Populus
spp.
Diagnostic traits for Populus alba, P. tremula and P. x canescens
Taxon
Rhytidome
Winter buds
Leaves
Petioles
Catkins
Male flowers
P. alba L.
■
■
■
not viscid
initially white tomentose,
later reddish glabrescent
short shoots: suborbicu
lar, subelliptic or
subpentagonal, entire or
sinuatedentate
■ long shoots: palmately
lobed, deltoid or ovate
oblong; base generally
cordate
■ initially whitetomen
tose; later dark green,
glabrous above, white or
greengreyish, tomentose
beneath
■
short shoots: 23 cm;
long shoots: up to 17 cm
■ not much flattened
■
■
■
■
■
Stigmas
white or greyish
■
■
female: up to 12 cm long
scales in female catkins
crenate or subentire,
hairy; scales in male
catkins: irregularly
crenatedentate or
subentire, hairy
(3)8(10) stamens
anthers purple at first,
later yellow
greenyellowish
bipartite
P. tremula L.
■
■
■
light greygreenish
short shoots: ovate
orbicular, obtuse, base
truncate or cordate, irre
gularly dentatecrenate
■ long shoots: larger,
ovatetriangular, apex
acute, base truncate or
cordate
■ initially more or less
hairy; later both sides
green, quite discolorous,
glabrous
■
■
■
■
■
■
■
■
■
whitish
frequently slightly viscid
initially rather pubes
cent, later brownish
glabrescent or glabrous
■
■
P. x canescens (Aiton) Sm.
■
■
■
short shoots: veins
prominent, glandulose
(2,5)46(8) cm
strongly flattened
laterally
512 cm long
scales hairy,
palmatelaciniate
(4)8(12) stamens
anthers purple
purple
bifid
■
■
■
■
■
short shoots: oval or suborbi
cular, sinuatedentate
long shoots: deltoidovate to
cordate
short shoots: initially greyish
pubescent; later more or less
concolorous, glabrous or
glabrescent; long shoots:
greytomentose beneath
short shoots: more than 5 cm
strongly flattened laterally
female: 46 cm long
scales in female catkins
irregularly laciniate
815 stamens
Populus 172
Diagnostic traits for Populus nigra, P. deltoides and P. x canadensis
Trunk
Twigs
Leaves
Female catkins
Male flowers
Capsules
P. nigra L
■
■
soon blackish, furrowed
often with large bosses
with epicormic shoots
terete or slightly angled
at apex
■ first yellowish, later
greyish
■
long shoots: 510 x 48
cm; short shoots: smaller
and broader
■ short shoots: rhombic,
base broadly cuneate or
more or less rounded;
long shoots: trullate
■ margin not ciliate
■ without glands at base
■
■
■
■
715 cm long
625 stamens
2 valves
P. deltoides Marshall
■
■
■
■
■
■
■
■
■
■
without bosses
strongly angled
P. x canadensis Moench.
■
■
without bosses
usually slightly angled at apex
first greenish, later
greenishbrown to
greyishbrown
1018 cm, longer than
wider
ovatecordate or deltoid,
base usually truncate
margin densely ciliate
glands at base
■
■
■
deltoid or ovate
margin shortly ciliate
usually glands at base
1520 cm long
3060 stamens
■
1525 stamens
34 valves
173 Populus
Taxon
Salix
spp.
Distribution
Salix alba L.
General distribution: Southwestern, Southeastern,
Middle and Eastern Europe, Caucasus, Western and
Middle Asia, Siberia, China, Northern Africa
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia, Bosnia
Herzegovina, Montenegro, Albania, Greece, Cyprus, Tur
key, Syria, Lebanon, Israel, Tunisia, Algeria, Morocco
Salix amplexicaulis Bory
General distribution: Southwestern Europe, Northern
Africa
Mediterranean region: Italy, Montenegro, Albania,
Greece, Turkey
Salix atrocinerea Brot.
General distribution: Southwestern, Middle and North
ern Europe, Northern Africa
Mediterranean region: Portugal, Spain, France (incl.
Corsica), Italy (Sardegna), Tunisia, Algeria, Morocco
Salix 174
Distribution
Salix eleagnos Scop.
General distribution: Southwestern, Southeastern,
Middle and Eastern Europe, Western Asia, Northern
Africa
Mediterranean region: Spain, France (incl. Corse), Italy,
Croatia, BosniaHerzegovina, Montenegro, Albania,
Greece, Turkey, Morocco
Salix fragilis L.
General distribution: Southwestern, Southeastern,
Middle and Eastern Europe, Caucasus, Western Asia
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia, Bosnia
Herzegovina, Montenegro, Albania, Greece, Turkey
Salix pedicellata Desf.
General distribution: Southwestern and Southeastern
Europe, Northern África
175 Salix
Mediterranean region: Spain, France (Corse), Italy
(Sardegna and Sicilia), Tunisia, Algeria, Morocco
Distribution
Salix purpurea L.
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus, West
ern Asia, Mongolia, Northern Africa
Mediterranean region: Portugal, Spain, France (incl.
Corse), Italy (incl. Sardegna and Sicilia), Croatia, Bo
sniaHerzegovina, Montenegro, Albania, Greece, Tur
key, Syria, Lebanon, Tunisia, Algeria, Morocco
Salix salviifolia Brot.
General distribution: Southwestern Europe
Mediterranean region: Portugal, Spain
Salix triandra L.
General distribution: Southwestern, Southeastern,
Middle, Northern and Eastern Europe, Caucasus, West
ern, Middle and Eastern Asia, Siberia, Russian Far East,
Mongolia, China, Northern Africa
Mediterranean region: Portugal, Spain, France, Italy,
Croatia, BosniaHerzegovina, Montenegro, Albania,
Greece, Turkey, Israel, Tunisia, Algeria
Salix 176
7 mm
1 cm
Salix alba L.
177 Salix
5 mm
5 mm
3 mm
1 cm
Salix amplexicaulis Bory
Salix 178
5 mm
1 cm
1 cm
179 Salix
Salix atrocinerea Brot.
1 cm
1 cm
5 mm
Salix eleagnos Scop.
Salix 180
5 mm
1 cm
5 mm
181 Salix
Salix fragilis L.
1 cm
1 cm
1 cm
Salix pedicellata Desf.
Salix 182
5 mm
5 mm
1 cm
183 Salix
Salix purpurea L.
1 cm
5 mm
1 cm
Salix salviifolia Brot.
Salix 184
5 mm
1 cm
5 mm
185 Salix
Salix triandra L.
Salix 186
▪ trunk: greyishbrown, deeply
longitudinally furrowed
▪ 210 x 12 cm
▪ linear or linearlanceolate
▪ base cuneate
▪ margin revolute, finely
glanduloseserrate
▪ glabrous above; densely
tomentose beneath
▪ base cuneate
▪ margin serrate
▪ silky or glabrescent,
rarely glabrous, glaucous
▪ glabrescent or glabrous, rarely
pubescent
▪ terete, very fragile at junctions
▪ branches: glabrous, orange,
greenish or reddishbrown
▪ trunk: greyish or greyishbrown,
becoming fissured
▪ tree to 825 m
S. fragilis L.
▪ alternate
▪ 210 x 0.52 cm
▪ lanceolate, oblonglanceolate,
obovatelanceolate, oblongobovate
or linearlanceolate
▪ glabrous
▪ angled
▪ glabrous
▪ smooth, reddishbrown, sometimes
almost black, greenish or brown,
flaking off in irregular patches
▪ shrub to 46(10) m
S. triandra L.
…/…
▪ base rounded, rarely cuneate
or quite attenuate
▪ margin glanduloseserrate,
▪ margin not revolute,
sometimes subserrate
glanduloseserrate
▪ glabrous, shiny above;
▪ glabrous, rarely quite hairy beneath
glaucescent or pale green beneath
▪ 516 x 13 cm
▪ lanceolate to ovatelanceolate
▪ glabrescent, slightly pubescent ▪ initially glabrescent, then
at base and/or apex
glabrous, shiny
▪ alternate
▪ alternate
▪ pubescent or glabrescent,
orange or yellow
▪ glabrous, reddishbrown
▪ shrub up to 6 m, rarely tree
S. eleagnos Scop.
▪ up to 10 x 12.5 cm
▪ lanceolate
▪ alternate
▪ silky
Buds
Leaves
▪ silky
Twigs
▪ branches: smooth, brown,
reddishbrown or orangeyellow
▪ tree up to 25 m
Bark
Habit
S. alba L.
Taxon
Leaves linear, lanceolate or ovatelanceolate
Diagnostic traits for species of Salix L. distributed in the Mediterranean region
187 Salix
27 x 1 cm
peduncle long
loose
appearing at the same time
as leaves
▪
▪
▪
▪
filaments free
filaments hairy
2 stamens
2 nectaries
▪ outer side glabrescent;
inner side pubescent
▪ caducous
▪ apex acute
▪ uniformly yellowish
▪ margin entire or serrate
▪
▪
▪
▪
▪ 25(7) mm
▪ pubescent
▪ margin dentade
▪ caducous
▪ linearlanceolate
S. alba L.
Female flowers ▪ pistil glabrous, sessile or
shortly stipitate
▪ style short
▪ stigmas bifid
▪ 1 nectary
Male flowers
Floral bracts
Catkin bracts
Catkins
Petioles
Stipules
Taxon
…/…
filaments free
filaments hairy
3 stamens
2 nectaries
▪ persistent
▪
▪
▪
▪
▪ style medium
▪ stigmas bifid
▪ 2 nectaries
▪ style short
▪ stigmas bifid
▪ 1 nectary
filaments free
filaments hairy at base
2 stamens
2 nectaries
▪ style long
▪ stigmas bifid
▪ 1 nectary
▪
▪
▪
▪
▪ appearing at the same time
as leaves
▪ (2)3.5(7) x 1 cm
▪ pistil glabrous, stipitate
filaments united at lower third
filaments hairy
2 stamens
1 nectary
▪ 510 mm
▪ glabrous or glabrescent, sometimes
glandulose at the junction of leaf
blade
▪ pistil glabrous, shortly stipitate ▪ pistil glabrous, stipitate
▪
▪
▪
▪
▪ uniformly pale yellow, brownish ▪ uniformly coloured
when adult, sometimes apex red
▪ apex obtuse, rarely emarginate
▪ margin hairy
▪ margin hairy
▪ outer side hairy; inner side
glabrescent
▪ persistent
▪ caducous
▪ margin entire
▪ > 5 mm
▪ glabrescent or glabrous
▪ semireniform, large
S. triandra L.
▪ caducous; present in young shoots ▪ persistent
▪ semicordate, very asymmetric,
broad
S. fragilis L.
▪ 27 x 0.51 cm
▪ peduncle long
▪ dense
▪ appearing just before the leaves ▪ appearing at the same time
or at the same time
as leaves
▪ up to 3 x 1 cm
▪ sessile or peduncle very short
▪ < 5 mm
▪ hairy
▪ absent or reduced to glands
S. eleagnos Scop.
Salix 188
▪ pubescent
▪ alternate
▪ 210 x 12 cm
▪ 510 x 13 cm
▪ 210 x 12 cm
▪ oblongobovate, elliptic, ▪ elliptic, oblongelliptic, ▪ oblongobovate, oblong
obovateelliptic, lanceolate lanceolate or obovate
lanceolate, obovate
or obovatelanceolate
lanceolate
lanceolate or linea
lanceolate
▪ base rounded, cuneate
▪ base rounded or shortly ▪ base rounded or
or shortly attenuate
attenuate, rarely cuneate
shortly attenuate
▪ margin revolute, entire, ▪ margin revolute, entire, ▪ margin revolute, dentade
paucidentate or
paucidentate or
serrate, sometimes
dentateserrate
dentateserrate
paucidentate or entire
▪ glabrous or tomentose
▪ glabrescent above;
▪ tomentose or glabrescent
above, reddish and white
thinly pubescent or
sometimes glabrous,
hairs; tomentose beneath,
glabrescent, glaucous
above; densely
reddish and white hairs,
beneath
tomentose beneath
sometimes glabrous,
glaucous
▪ veins prominent beneath ▪ veins prominent beneath
▪ pubescent
▪ alternate
▪ shrub up to 6 m
S. purpurea L.
…/…
▪ base rounded, sometimes
shortly attenuate or cuneate
▪ margin not revolute,
dentate towards apex,
entire towards base
▪ glabrous above;
glaucescent beneath
▪ base cordate
semiamplexicaul
▪ margin serrate towards
apex, entire towards base
▪ glabrous, dark green
above; glabrous,
glaucous beneath
▪ 57 x 11.5 cm
▪ linear, linearlanceolate,
oblongobovate or
spatulate
▪ glabrous
▪ opposite, sometimes
alternate
▪ up to 3050 x 816 mm
▪ oblonglanceolate
or oblong
▪ glabrous
▪ opposite or suboposite,
rarely alternate
▪ glabrous, very shiny
▪ yellowishbrown to reddish ▪ glabrous, shiny greyish,
brown or dark brown
yellowish, reddishbrown or black
▪ decorticate wood
smooth, unridge
▪ shrub to 3(5) m
S. amplexicaulis Bory
Leaves generally opposite
▪ pubescent or glabrescent ▪ glabrous
Leaves
Buds
▪ tomentose
▪ pubescent
▪ shrub up to 6 m
Twigs
▪ shrub or small tree
up to 10 m
▪ glabrous, reddishbrown ▪ glabrous, reddishbrown ▪ glabrous, reddishbrown
or greyishbrown
or greyishbrown
or greyishbrown
▪ decorticate wood with
▪ decorticate wood with ▪ decorticate wood with
numerous prominent
numerous prominent
numerous prominent
ridges
ridges
ridges
▪ shrub, sometimes small
tree up to 12 m
S. salviifolia Brot.
Bark
Habit
S. pedicellata Desf.
S. atrocinerea Brot.
Taxon
Leaves broadly lanceolate, oblongelliptic, elliptic or obovate
Diagnostic traits for species of Salix L. distributed in the Mediterranean region
189 Salix
▪ ± 5 mm
▪ hairy
Petioles
▪ up to 7 x 12 cm
▪ up to 7 x 1(2) cm
▪ sessile or peduncle short ▪ peduncle short
▪ appearing before the
▪ appearing before the
leaves
leaves
Female
flowers
Male
flowers
Floral
bracts
▪ pistil glabrous,
longstipitate
▪ style medium
▪ stigmas entire or bifid
▪ filament glabrous
or glabrescent
▪ 2 stamens
▪ filaments more or less
hairy at base
▪ 2 stamens
▪ 1 nectary
▪ pistil tomentose,
stipitate
▪ style short
▪ stigmas entire or bifid
▪ 1 nectary
▪ 2 stamens
▪ 1 nectary
▪ filaments free
▪ filaments free
▪ filaments hairy
near base
▪ filaments united
▪ apex darker
▪ usually opposite
▪ peduncle short
▪ (0.3)0.53.2 mm
▪ caducous
▪ small
S. amplexicaulis Bory
▪ pistil pubescent, stipitate ▪ pistil pubescent, sessile
or shortly stipitate
▪ style short
▪ generally stigmas entire
▪ 1 nectary
▪ filaments united at
base or free
▪ filaments hairy
▪ hairy
▪ apex darker
▪ greyish, densely tomentose
▪ 37 x 12 cm
▪ peduncle short
▪ appearing at the same
time as leaves
▪ < 5 mm
▪ tomentose
▪ persistent
▪ margin serrate
▪ tomentose
▪ semicordate
S. salviifolia Brot.
▪ apex darker
▪ usually apex darker
▪ apex obtuse
▪ apex obtuse
▪ more or less densely hairy ▪ hairy
Catkin bracts
Catkins
▪ ≥ 5 mm
▪ hairy
S. pedicellata Desf.
▪ semicordate or
semireniform, broad
▪ margin not much dentade ▪ margin dentate
▪ glabrescent above;
tomentose beneath
▪ caducous; present in
▪ caducous
young shoots
Stipules ▪ semicordate or reniform
S. atrocinerea Brot.
Taxon
…/…
▪ style short, glabrous
▪ stigmas entire
▪ 1 nectary
▪ pistil pubescent, sessile
▪ 1 nectary
▪ filaments hairy
▪ filaments united
▪ usually apex darker
▪ apex obtuse
▪ hairy
▪ 3 x 1 cm
▪ sessile or peduncle short
▪ appearing before the
leaves
▪ < 5 mm
▪ glabrous
▪ caducous
S. purpurea L.
Tamarix
spp.
Distribution
Tamarix africana Poiret
General distribution: Southwestern and Southeastern
Europe, Northern Africa, Macaronesia
Mediterranean region: Portugal, Spain (incl. Baleares),
France (incl. Corse), Italy (incl. Sardegna and Sicilia),
Tunisia, Algeria, Morocco
Tamarix boveana Bunge
General distribution: Southwestern Europe, Northern
Africa
Mediterranean region: Spain, Libya, Tunisia, Algeria,
Morocco
Tamarix canariensis Willd.
General distribution: Southwestern and Southeastern
Europe, Northern Africa, Macaronesia
Mediterranean region: Portugal, Spain (incl. Baleares),
France, Italy (Sardegna and Sicilia), Libya, Tunisia, Al
geria, Morocco
Tamarix 190
Distribution
Tamarix dalmatica Baum
General distribution: Southeastern Europe
Mediterranean region: Italy, Croatia, Bosnia Herze
govina, Montenegro, Albania, Greece
Tamarix gallica L.
General distribution: Southwestern and Southeastern
Europe, Macaronesia
Mediterranean region: Portugal, Spain (incl. Baleares),
France (incl. Corse), Italy (incl. Sardegna and Sicilia)
Tamarix hampeana Boiss. & Heldr.
General distribution: Southeastern Europe, Western
Asia
191 Tamarix
Mediterranean region: Greece, Turkey, Israel
Distribution
Tamarix parviflora DC.
General distribution: Southeastern Europe, Western
Asia, Northern Africa
Mediterranean region: Croatia, BosniaHerzegovina,
Montenegro, Albania, Greece (incl. Kriti), Turkey, Israel,
Algeria
Tamarix smyrnensis Bunge
General distribution: Southeastern and Eastern Europe,
Caucasus, Western and Middle Asia
Mediterranean region: Greece (incl. Kriti), Cyprus,
Turkey, Syria, Lebanon, Israel
Tamarix tetranda Pallas ex Bieb.
General distribution: Southeastern and Eastern Europe,
Western Asia
Mediterranean region: Albania, Greece, Cyprus, Turkey,
Lebanon
Tamarix 192
193 Tamarix
▪ 1.54 mm long
▪ smooth or minutely papillose
▪ margin scarious
▪ 26 mm long
▪ papillose
▪ reddishbrown or brownish
T. boveana Bunge
▪ synlophic to parasynlophic
▪ muticous or slightly apiculate
▪ (2.7)34 x 1.32 mm
▪ narrowly ovate to unguiculate
▪ muticous or slightly apiculate
Nectariferous ▪ synlophic
disc
▪ 23.3 x 12 mm
▪ trullate to ovate
Anthers
Petals
▪ 11.8 mm long
▪ 1.73 x 1.52.4 mm
▪ trullate, acute; outer slightly
▪ outer trullate, acute; inner
longer, narrower and more acute
ovate, slightly shorter, obtuse
▪ margin subentire
▪ margin outer entire;
inner denticulate at apex
Sepals
▪ tetramerous, rarely tetramerous
and pentamerous
▪ margin densely papillose
▪ equal or longer than calyx
▪ linear, acute
▪ pentamerous
▪ longer or shorter than calyx
▪ narrowly oblong, shortly acute,
to triangular, acuminate
▪ margin usually papillose
Flowers
Bracts
Inflorescences ▪ (15)3070(80) x (5)69 mm
▪ 40150 x 712 mmq
▪ usually on previous year's branches ▪ usually on previous year's branches
▪ rachis sometimes papillose
▪ usually simple
▪ usually simple
▪ black or darkpurple
Leaves
Bark
T. africana Poiret
Taxon
▪ glaucous
▪ green
▪ synlophic
▪ fleshy
▪ apiculate
▪ 1.21.6(1.7) x 0.51 mm
▪ obovate
▪ margin densely denticulate
▪ 0.61 mm long
▪ trullate, acute
▪ pentamerous
▪ equal or longer than calyx
▪ lineartriangular, long
acuminate to subulate
▪ margin papillose
▪ synlophic
▪ not very fleshy
▪ slightly apiculate
▪ (1.6)1.72 x 0.81 mm
▪ elliptic to ovate
▪ 0.71.8 mm long
▪ trullate to ovate, acute; inner
somewhat longer and more obtuse
▪ margin not very denticulate
▪ pentamerous
▪ margin denticulate
▪ usually shorter than calyx
▪ narrowly triangular, acuminate
▪ 1050 x 35 mm
▪ 1050 x 35 mm
▪ usually on current year's branches▪ usually on current year's branches
▪ rachis usually papillose
▪ rachis usually glabrous
▪ densely compound
▪ loosely compound
▪ 1.32.5 mm long
▪ few or without saltsecretory glands
▪ brownishblack or deep purple
T. gallica L.
▪ 1.32.5 mm long
▪ many saltsecretory glands
▪ purple or reddishbrown
T. canariensis Willd.
Diagnostic traits for species of Tamarix L. distributed in the Western European Mediterranean region
Tamarix 194
▪ 24 mm long
▪ 2060(130) x (8)1012 mm
▪ usually on previous
year's branches
▪ simple or loosely compound
▪ 1.754 mm long
▪ brown or reddishbrown
T. hampeana Boiss. & Heldr.
▪ fleshy
▪ paralophic
▪ 2.54 mm long
▪ ovateelliptic
T smyrnensis Bunge
▪ shorter than calyx
▪ longer than pedicels
▪ synlophic
▪ 1.82.5 mm long
▪ parabolic or oblong
▪ 22.5 mm long
▪ outer trullate, keeled, acute;
inner ovate, shorter, obtuse
▪ tetramerous, occasionally
▪ oblong, herbaceous in
proximal half, obtuse
▪ shorter than calyx
▪ longer than pedicels
▪ hololophic, lobes entire to ▪ paralophic to parasynlophic
faintly emarginate, hypodiscal
insertion of stamens
▪ usually fleshy
▪ fleshy
▪ 22.75mm long
▪ 2.23 mm long
▪ovate to suborbicular, strongly ▪ ovate to ovateelliptic
keeled
▪ persistent
▪ margin irregularly denticulate ▪ margin entire
▪ 1 mm long
▪ trullate to ovate, obtuse
▪ pentamerous
pentamerous
▪ margin almost entirely scarious
▪ tetramerous
▪ 35 mm long
▪ margin scarious
▪ black to greyishblack
T. tetrandra Pallas ex Bieb.
▪ up to 40 x 4 mm
▪ 3060 x 67 mm
▪ usually on previous year's
▪ usually on previous
and/or current year's branches
year's branches
▪ loosely compound
▪ simple or loosely compound
▪ 23.5 mm long
▪ reddish brown to brown
▪ triangular acuminate, obtuse
▪ shorter than calyx
▪ longer than pedicels
▪ 1040 x 36 mm
▪ usually on previous
year's branches
▪ simple
▪ 1.62.5 mm long
▪ reddishbrown to purple
T. parviflora DC.
▪ 11.5 mm long
▪ joined at base; outer
trullate, keeled, acute;
inner ovate, obtuse
▪ margin outer subentire; inner ▪margin irregularly
denticulate
slightly denticulate
▪ 22.5 mm long
▪ trullate, acuminate; outer
more acute
▪ muticous or slightly apiculate
▪ 2.34.5(5) x 1.41.8 mm
▪ narrowly ellipticobovate,
unguiculate
▪ subpersistent
▪margin entire or scarcely
denticulate
▪ 1.53.5 x 1.52.4 mm
▪ trullate, more or less
keeled; outer more acute
▪ tetramerous, sometimes few ▪ tetrapentamerous
pentamerous
▪ equal or longer than calyx ▪ shorter than calyx
▪ shorter than pedicels,
sometimes equal to or longer
▪ broadly triangular,
obtuse to acuminate
▪ margin scabridpapillose
Nectariferous▪ paralophic
disc
Anthers
Petals
Sepals
Flowers
Bracts
Inflorescences ▪ 2070 x 712 mm
▪ usually on previous
year's branches
▪ simple
▪ reddishblack,
brownishblack to black
Leaves
Bark
T. dalmatica Baum
Taxon
Diagnostic traits for species of Tamarix L. distributed in the Eastern European Mediterranean region
Glossary
acuminate
tapering gradually to a protracted point
acute
ending in a point
adaptability
the ability of an individual or a population to change to fit
changed environmental conditions
allotetraploid
an organism having four sets of chromosomes, produced
by the union of genetically distinct chromosome sets (usu
ally form different species)
alternate
not opposite or whorled; placed singly at nodes
amplexicaul
clasping the stem
androdioecy
sex expression that occurs when both male and hermaph
rodite individuals are present in a population
andromonoecy
sex expression that occurs when both male and hermaph
rodite flowers occur on the same individual
anemophilous
pollinated by wind
anther
the terminal portion of a stamen, which bears the pollen
sacs
anthesis
the period during which a flower is fully open and functional
apex
the distal part of a leaf
apical bud
a bud at the tip of a shoot
apiculate
ending in a mucro
apomixis
development of an embryo without the occurrence of fer
tilisation. Since this process does not involve the forma
tion of gametes or meiosis, the progeny are genetically
identical to their parent
attenuate
tapering gradually
autochthonous
originating in the region where it is found
autogamy
selffertilisation
auxin
a class of hormones that promotes and regulates
the growth and development of plants, including cell
elongation
axillary bud
a bud at the junction of a stem and a petiole
backcrossing
a cross of an hybrid with one of the two parental indi
viduals or species
berry
a fleshy, indehiscent, manyseeded fruit containing no
hard parts except the seed
bifid
divided into two parts along less than half its length
bipartite
divided into two parts along at least half its total length
bipinnate
twice pinnate; a pinnate leaf, having also pinnately di
vided segments
bract
a modified leaf, generally small, growing just below a
flower or an inflorescence
bracteole
A small modified leaf on the flower stalk, above the bract
and below the calyx
caducous
dropping off or shedding early
calyx
all the sepals of a flower
cambium
a layer of actively dividing cells responsible for the in
crease in thickness of stems and roots
capsule
a dry dehiscent fruit that develops from two or more
carpels
carpel
modified leaf forming a pistil
cation
an ion or group of ions having a positive charge
cation exchange
a chemical process in which cations of like charge are ex
changed equally between a solid and a solution
195 Glossary
achene
a small, dry, indehiscent, usually singleseeded fruit
catkin
a long inflorescence, usually pendulous, with tiny, unisex
ual, petalless flowers
caudate
bearing a taillike appendage
centre of origin
a geographical location or local region where a particular
group of organisms is believed to have originated
chromosome
a structure in all living cells, consisting of a molecule of
DNA bonded to various proteins, that carries the genes
ciliate
bearing hairs on the margin
clone
cells, groups of cells or organisms that are produced asex
ually from a single ancestor, to which they are genetically
identical
compound inflorescence
an inflorescence with more than one flower per branch
compound leaf
one having two or more distinct leaflets
concolorous
of uniform colour above and below
connate
united with a structure of the same kind
cordate
heartshaped
coriaceous
having a leathery texture
corolla
the petals of a flower considered as a group, often coloured
corymb
an inflorescence whose outer flowers have longer pedicels
than the inner ones, so that together they form a round,
rather flattopped cluster; the outer flowers open before
the inner ones
corymbiform
a round, rather flattopped inflorescence, resembling a
corymb
crenate
with rounded teeth
crossbreeding
breeding from individuals of different species, varieties or
breeds
crossfertilisation
fertilisation by the union of a male gamete and a female
gamete from different individuals of the same species
cultivar
a cultivated plant that has been selected because it has de
sirable characteristics that distinguish it from other plants
of the same species; a cultivar receives a unique name
cuneate
wedgeshaped, with the narrow end at base
cutting
a part of a stem, root or other part removed from a donor
plant to produce a new plant by inducing roots
cyme
inflorescence in which the main axis terminates in a
flower, as do the secondary axes that arise alongside it
cytokinin
a class of plant hormone involved in cell division and growth.
deciduous
shedding leaves at the end of the growing season
decussate
opposite arrangement on a stem, at right angles to the
pairs immediately above and bellow
dehiscent
opening spontaneously at maturity
deltoid
in the shape of a capital delta or equilateral triangle
dense
having parts very close together
dentate
with toothlike projections along the margin
denticulate
with small teeth
dioecy
sex expression that occurs when either female or male in
dividuals are present in a population
diploid
having two sets of chromosomes in each cell. In organisms
that reproduce sexually, one set of chromosomes is inher
ited from each parent
discolorous
of different colour or shade above and beneath
distal
the region of an organ that is furthest from the point by
which it is attached to the plant
Glossary 196
dormancy
a period of cessation of growth or development
double flower
a flower having many more than the usual number of
petals, usually in a crowded or overlapping arrangement
doubletoothed
dentate with each tooth bearing smaller teeth
follicle
dry fruit derived from a single carpel which splits along
one side only
gamete
a reproductive cell
garriga
(pl. garrigues) scrubland of low shrubs with a liking for al
kaline reaction soils
gene
the hereditary fundamental unit, that occupies a specific
location on a chromosome
drupe
a fleshy fruit with a stony endocarp that encloses one or
few seeds, such as a plum
gene flow
the movement of genes among different populations that
results from dispersal of gametes and seeds
elliptic
narrowed at ends and widest at or near the middle
genetic drift
fluctuations in the frequency of the appearance of a gene
among generations; these changes occur by chance rather
than by natural selection and their effects are most marked
in very small, isolated populations
effective population size
the average number of individuals in a population that ac
tually contribute genes to succeeding generations by mat
ing; this number is generally rather lower that the census
number because there is a large differences in reproduc
tive success between individuals
emarginate
with a shallow notch at the apex
endocarp
the innermost layer of the fruit wall
entire
not broken up into teeth or lobes
entomophilous
pollinated by insects
etiolation
phenomenon that occurs when plants or part of plants are
grown in either partial or complete absence of light, char
acterised by much more rapid elongation and pale yellow
coloration of the organs
evapotranspiration
the process of moisture transfer from the earth to the at
mosphere through the evaporation of water and transpi
ration from plants
explant
an organ or tissue excised from a donor plant that is used
to initiate an in vitro culture
fascicle
cluster of shortstalked leaves or flowers growing closely
together
filament
a thready structure that bears the anther in the stamen
genetic diversity
the totality of different genes in a group of individuals or
species
genetic transformation
modification of the genome through the addition of DNA
from a cell of a different genotype
genetic variation
the differences observed among individuals within a partic
ular population or between populations, due to their genes
genome
complete set of genes in a set of chromosomes
genotype
the genetic constitution of an organism
germination capacity
percentage of seeds which produce normal seedlings in
relation to the total number of seeds of a sample taken
from a seed lot
gibberellic acid
a hormone obtained from the fungus Gibberella fujikuroi
glabrescent
nearly glabrous
glabrous
smooth, lacking hairs
glandule
a minute secreting part or appendage
glandulose
having glandules
197 Glossary
DNA
deoxyribonucleic acid consists of two long chains of nu
cleotides linked together in a structure resembling a lad
der twisted into a spiral
glaucous
slightly bluish shade of light green in colour
glaucescent
tending towards glaucous
globose
spherical
glomerule
a compact, globose inflorescence with many sessile or sub
sessile flowers
gynodioecy
sex expression that occurs when female and hermaphro
dite individuals are present in a population
hairy
having hairs
haploid
having a single set of chromosomes
hardwood cutting
a lignified cutting, taken from the previous season’s
growth of in dormant woody plants
head
a dense cluster of sessile flowers attached to a receptacle
herbaceous
not woody or membranous. Leaflike texture
hermaphroditism
having flowers with both male and female reproductive parts
hispid
with short, stiff or bristly hairs
hololophic
nectariferous disc divided into five lobes, each lying be
tween two stamens that may be either free from or united
to the disc; the lobes may be entire or have an obtuse,
truncate, retuse or emarginate apex
hybridisation
crossbreeding between individuals of different species
hypodiscal
beneath the nectariferous disc
in vitro
aseptic culture under laboratory conditions
inbreeding
the breeding or mating of related individuals
inbreeding depression
the reduction in vigour often observed in progeny from
mating between close relatives
indehiscent
not splitting open at maturity
indolebutyric acid
a synthetic hormone promoting elongation of stems and
roots (IBA)
indumentum
a covering of hairs, scales, glandules, etc. found on the
surface of different organs
inflorescence
a branching flowering system, its branches ending in flowers
infructescence
a fruiting structure that consists of more than a single
fruit, produced by an inflorescence
inoculation
the process of introducing a microorganism into a plant
internode
the portion of a stem between two successive levels of in
sertion of leaves or leaf pairs
interspecific
occurring between different species
intraspecific
occurring between individuals or populations of the same
species
introgression
the incorporation of genes of one species into the genetic
information of another through hybridisation followed by
backcrossing
involucre
a series of bracts below or around a flower or an
inflorescence
keeled
having a ridge or shaped like a ridge or the keel of a ship
laciniate
deeply cut into irregular, narrow segments
lanate
having woolly hairs
lanceolate
like the blade of a lance, tapering towards the apex and
towards the base
lateral bud
axillary bud
layer
a shoot which, while still attached to the plant, forms
roots when it comes into contact with a substrate. The
layered section, with its roots, can then be separated from
the original plant
leaflet
one of the blades or leaflike divisions of a compound leaf
Glossary 198
lignify
to turn into wood or become woody
linear
long and narrow with parallel or almost parallel margins
lobe
a rounded projection
lobule
a subdivision of a lobe
long shoot
major shoot that has long internodes
loose
having parts widely separated from one another
macronutrient
a mineral used by plants in a relatively large amounts
mallet cutting
a hardwood cutting that includes a short section of stem
of the previous season’s growth
margin
edge of a laminar organ (such as a leaf)
maquis
droughtadapted Mediterranean scrubland composed of
shrubs and small trees with thick, coriaceous, perennial
leaves or spiny foliage.
meiosis
the process of cell division in sexually reproducing organ
isms that involves the reduction in the number of chro
mosomes and some exchange of genetic material, leading
to the production of genetically different, haploid repro
ductive cells
meristem
Plant tissue whose cells actively divide to form new tissues
that cause the plant to grow
micronutrient
a mineral used by plants in very small amounts
micropropagation
plant tissue culture
molecular marker
a gene or a specific fragment of DNA that can be used to
identify an organism, species or strain or a phenotypic trait
associated with it
monoecy
sex expression that occurs when the same individual has
separate female and male flowers
mucro
a short point
muticous
lacking a mucro
natural selection
the process by which favourable heritable traits become
more common in successive generations
navicular
shaped like a boat
nectariferous disc
a nectarsecreting disklike or ringlike glandulose out
growth from the receptacle inside the flower
nectary
a gland that secretes nectar, usually located at the bases
of insectpollinated flowers
node
the point on a stem where a leaf is attached
oblanceolate
reverse lanceshaped, broadest in upper third and taper
ing from the middle towards the base
oblong
longer than broad with margins nearly parallel for most of
their length
obovate
ovate but with the terminal half broader than the basal
half
obtuse
blunt, not acute
opposite
arrangement in which leaves arise in pairs on either side
of each node
orbicular
circularshaped
ortet
the original plant from which a clone is started through
vegetative propagation
oval
having the shape of an ellipse
ovary
the ovulebearing lower part of a pistil that ripens into a fruit
ovate
with the outline of an egg having the broader end at base
palmate
having three or more leaflets or lobes radiating from one
point, like the fingers of a hand
199 Glossary
legume
dry fruit derived from a single carpel splitting along both
sides into two valves when ripe
palmatifid
palmately divided up to halfway to the base
panicle
a raceme composed of racemes, frequently pyramidshaped
paniculiform cyme
a cyme resembling a panicle
papilla
tiny conical outgrowth from an epidermal cell
papillose
having papillae
paralophic
nectariferous disc divided into four or five lobes with trun
cate apex, united to one stamen
parthenocarpy
the production of fruit without fertilisation
patent
spreading from the stem at an open angle
paucidentate
with few teeth
pedicel
stalk of each flower in a compound inflorescence
peduncle
stalk of a solitary flower or of an inflorescence
pentagonal
resembling a pentagon, having five sides
pinnate
having divisions or segments arranged on each side of a
common axis
pinnatisect
deeply divided pinnately to nearly the middle vein
pistil
the female organ of a flower, comprising the stigma, style
and ovary; it consists of a single carpel or of several
carpels fused together
plasticity
the ability of a single genotype to give rise to a spectrum
of phenotypes. Plasticity may occur as different pheno
types among different individuals of the same genotype,
different phenotypes within the lifetime of a single indi
vidual, or different phenotypes in response to specific en
vironmental conditions
poly
prefix denoting many
pollen sac
structure where pollen grains are produced
polyploid
having three or more sets of chromosomes
pome
a simple fruit that has several seed chambers developed
from a pluricarpelar ovary embedded in a fleshy part de
veloped from the receptacle
pentamerous
having flower parts, such as petals, sepals or stamens, in
sets of five
population
a group of individuals of the same species, occupying a
particular geographic area, that experiences total or great
reproductive isolation
perianth
the envelope of a flower, consisting of the calyx and the
corolla
pruinose
having a pale grey waxy dustlike coating
perennial leaf
leaf lasting for more than one vegetative period
persistent
not falling off
pH
potential of hydrogen; a measure of the acidity or alka
linity of a solution
phenology
the relationship between a periodic biological phenome
non and climatic conditions; the time frame of any sea
sonal biological phenomenon
phenotype
the observed characteristics of an individual, determined
by its genotype interacting with the environment in which
it is growing
provenance
a specific geographical location within the distribution
range of a species
puberule
minutely pubescent
pubescent
having short soft hairs
pulverulent
covered with a fine powder
purity
the percentage composition by weight of pure seeds in
relation to the total weight of a sample taken from a
seed lot; the sample is separated into three components:
pure seeds, seeds of other species and inert matter
Glossary 200
raceme
a simple inflorescence having stalked flowers arranged al
ternately along a single axis
racemiform cyme
a cyme resembling a raceme but with the axis ending in
a flower
rachis
the main axis of an inflorescence or of a pinnately com
pound leaf
ramet
an individual member of a clone
receptacle
the expanded tip of a flower stalk that bears the floral
structures or the group of flowers in some inflorescences
region of provenance
an area demarcated for a species or group of species in
which stands or seed sources showing similar phenotypic or
genetic characters are found, or an area or group of areas
with uniform or similar ecological conditions
reniform
kidneyshaped
reproductive material
fruits, seeds, plants and part of plants (buds, cuttings, ex
plants, embryos, layers, roots, scions, sets and other parts)
for the production of planting stocks
retuse
notched slightly at an obtuse apex
revolute
with the margins curling under
rhizomatous
having rhizomes
rhizome
a horizontal, usually underground stem that often sends
out roots and shoots
rhytidome
dead tissues, generally rough and cracked, that can cover
the trunk, branches and roots of trees and shrubs
samara
a one seeded indehiscent dry fruit with a winglike structure
scarious
thin, dry and membranous, not green
scion
part of a plant for grafting into another plant
scabrid
rough to touch due to minute, hard projections
seed orchard
a plantation of selected clones or families which is iso
lated or managed so as to avoid or reduce pollination from
outside sources, and managed so as to produce frequent,
abundant and easily collected crops of seeds
seed source
trees within an area from which fruits and seeds are
collected
selfcompatible
capable of selffertilisation
selffertilisation
fertilisation by the union of male and female gametes
from the same individual
selfincompatible
incapable of selffertilisation
semi
prefix denoting a half or that something is halfdone
semihardwood cutting
a stiff, partially lignified cutting taken from the new
growth of woody plants during the growing season
sepal
one of the parts of the outer whorl of a flower, usually
greenish
serial cutting
a method of propagation by which cuttings, once rooted,
are divided the following year to produce a greater num
ber of cuttings
sericeous
covered with fine, short hairs, somewhat shiny, resembling
silk
serrate
sawtoothed, with acute teeth pointing towards the apex
serrulate
minutely serrate
sessile
lacking a stalk and attached directly at the base
set
oneyearold shoot from a stool, used to obtain cuttings
set of chromosomes
the group of qualitatively different chromosomes inher
ited as a unit from one parent
201 Glossary
quantitative feature
a characteristic that shows continuous phenotypic varia
tion; it often depends upon the cumulative action of many
genes, each of which produces only a small effect, and may
also be influenced significantly by environmental effects
seta
a stiff hair
subspecies
taxonomic category immediately below species
short shoot
lateral shoot that has very short or no internodes
sulcate
having long, narrow furrows or grooves
setose
bearing setae
simple inflorescence
an inflorescence with singleflowered branches
simple leaf
not lobed or divided
sinuate
having a wavy margin
softwood cutting
a cutting taken from a not already lignified soft stem or tip
of a woody plant
solitary flower
flower not arranged on an inflorescence
spatulate
shaped like a spatula
spike
a raceme in which the flowers are all sessile
stamen
the male organ of a flower, comprising the filament and
the anther
stand
delineated population of trees possessing at least a mod
erate degree of uniformity in composition, structure and
quality
stellate
starshaped
stigma
top part of the pistil, on which pollen is deposited at pol
lination
subulate
tapering very gradually upwards to a fine point
sympatry
the occurrence of species or subspecific taxa in the same
area or in overlapping areas
synlophic
nectariferous disc deeply divided into four or five, some
times three, lobes with attenuate apexes united to one
stamen, giving the impression that the stamen itself has a
broad base
taxon
(pl. taxa) a group of organisms forming any taxonomic
category (e.g. family, genus or species)
TDR
a probe used to measure soil water content (time domain
reflectometry)
terete
circular in crosssection
terminal
appearing at the end of a stem or a similar part
tetramerous
having flower parts, such as petals, sepals or stamens, in
sets of four
tetraploid
having four sets of chromosomes
thalweg
line connecting the lowest parts of a river bed or valley
thermophile
native of warm climes; unable to withstand cold
stipe
stalk of a pistil
thyrse
a dense inflorescence comprising a racemose central axis
with lateral branches ending in cymes
stipule
a leafy appendix at the base of a petiole, usually occurring
in pairs
trichome
a hair, bristle, papilla or scalelike outgrowth from an epi
dermal cell
stipitate
carried on a stipe
style
the slender part of a flower pistil, extending from the ovary
to the stigma
sub
prefix denoting somewhat, slightly, rather, or under
tomentose
totally covered with hairs
tripinnate
three times pinnate; a pinnate leaf with bipinnately di
vided segments
trullate
ovate but margins straight; trowelshaped
Glossary 202
truncate
appearing to terminate abruptly, as though an end or point
has been cut off
turion
a thick fleshy young shoot or sucker rising from a subter
ranean bud
umbel
a usually umbellashaped inflorescence in which all the
pedicels arise from about the same point at the tip of the
peduncle; the outer flowers open before the inner ones
unguiculate
narrowed into a claw
unisexual
having either stamens or pistils but not both
valve
one of the sections into which a legume or other dehiscent
fruit splits
variety
a taxonomic subdivision of a species consisting of a group
of individuals that differ from others of the same species
in minor but heritable characteristics
vein
one of the vascular bundles or ribs that form the branch
ing framework of conducting and supporting tissues in a
leaf or other expanded part
viability
the capacity of a seed to germinate under favourable con
ditions; it is usually expressed as the percentage of seeds
having a live embryo in relation to the total number of
seeds of a sample taken from a seed lot
vicariant
each of the species in different, more or less distant geo
graphical areas that play the same ecological role and dis
play only limited morphological differences from each
other
whorl
the arrangement of three or more leaves, petals or other
organs radiating from a single node
203 Glossary
zonal
having a geographic distribution that is mainly
determined by climate
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